rogs of the Genus Eleutherodactylus in Western Ecuador ;?v j^"--* ■*),' 'l»tv ^^.^^^v.^*' \ l^a. -T'»«*'?^S!tr>f>: Sj^stematics, Ecology, and Biogeography John D. Lynch and William E. Dpd Ernst Msyr Lfbrary HlMMtwTt r?f nnmrjarstn/e Zoology THEUl Museum of Comparative Zoology HERPETOLOGY LIBRARY { MUSEUM The Universi ling with Volume 1 in 1946, was disco nerly published in the above series wt Museum Occasional Papers until Number 180 in December 1996. The University of Kansas Natural History Museum Miscellaneous Publications began with Number 1 in 1946 and ended with Number 88 in February 1996. Monographs of the The University of Kansas Natural History Museum were initiated in 1970 and discontinued with Number 8 in December 1992. The foregoing publication series now are combined in a new series entitled The University of Kansas Natural History Museum Scientific Papers, begun in January 1997. 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Include expiration date. Cover design by Linda Trueb. Photograph of Eleutherodactylus labiosus by the late Kenneth Miyata. The University of Kansas Natural History Museum Special Publication No. 23 21 February 1997 Frogs of the Genus Eleutherodactylus (Leptodactylidae) in Western Ecuador: Systematics, Ecology, and Biogeography John D. Lynch Professor, School of Life Sciences The University of Nebraska, Lincohi, Nebraska 68588 USA, Associate, Division of Herpetology Natural History Museum, The University of Kansas Lawrence, Kansas 66045, USA William E. Duellman Curator, Division of Herpetology Natural History Museum Professor, Department of Systematics and Ecology The University of Kansas Lawrence, Kansas 66045, USA Natural History Museum The University of Kansas Lawrence, Kansas SPECIAL PUBLICATIONS Editor: Linda Trueb Managing Editor: Joseph T. Collins Special Publication No. 23 pp. i-iv, 1-236 Plates 1-8 Published 21 February 1997 ISBN 0-89338-054-7 © 1997 BY Natural History Museum Dyche Hall The University of Kansas Lawrence, Kansas 66045-2454 USA Printed by University of Kansas Printing Service Lawrence. Kansas CONTENTS ABSTRACT 1 RESUMEN 1 INTRODUCTION 1 Acknowledgments 3 MATERIALS AND METHODS 4 WESTERN ECUADOR 4 Physiography 8 River Systems 9 Climate 9 BiocLiMATic Regimes 12 Vegetation 17 Geologic and Climatic History 18 Human Environmental ModiAcations 23 SYSTEMATICS 24 Description of Characters 24 Subgenera and Species Groups 41 Key to Species 53 Clave de las Especies 58 Accounts of Species 63 Eleutherodactylus achatinus (Boulenger) 63 Eleuthewdactylus actites Lynch 66 Eleutherodactylus anatipes Lynch and Myers 68 Eleutherodactylus anonudus (Boulenger) 69 Eleutherodactylus apiculatus Lynch and Burrowes 70 Eleutherodactylus appendiculatus (Werner) 71 Eleutherodactylus babax Lynch 72 Eleutherodactylus cajamarcensis Barbour and Noble 73 Eleutherodactylus calcarulatus Lynch 74 Eleutherodactylus caprifer Lynch = 76 Eleutherodactylus celator Lynch 76 Eleutherodactylus cerastes Lynch 78 Eleutherodactylus chalceus (Peters) 79 Eleutherodactylus colomai new species 81 Eleutherodactylus crenungiiis Lynch 83 Eleutherodactylus crucifer (Boulenger) 85 Eleutherodactylus degener new species 86 Eleutherodactylus dissimulatus new species 88 Eleutherodactylus duelhnani Lynch 90 Eleutherodactylus eremitus Lynch 91 Eleutherodactylus eugeniae new species , 93 Eleutherodactylus floridus new species 94 Eleutherodactylus gentryi new species 97 Eleutherodactylus gularis (Boulenger) 99 Eleutherodactylus hamiotae Flores 102 Eleutherodactylus hectus Lynch and Burrowes 103 Eleutherodactylus helonotus (Lynch) 104 Eleutherodactylus illotus new species 104 Eleutherodactylus lahiosus Lynch, Rui'z, and Ardila 106 Eleutherodactylus laticlavius Lynch and Burrowes 107 Eleutherodactylus latidiscus (Boulenger) 108 Eleutherodactylus leoni Lynch 1 10 Eleuthewdactyliis longirostris (Boulenger) 1 1 1 Eleutherodactylus loustes Lynch 1 12 Eleuthewdactylus luteolatemlis Lynch 113 Eleutherodaciylus lyinani Barbour and Noble 1 14 Eleuthewdactylus muricatus Lynch and Miyata 115 Eleutherodactylus necerus Lynch 116 Eleutherodactylus nyctophylax Lynch 118 Eleutherodactylus ocellatus Lynch and Burrowes 119 Eleutherodactylus ornatissimus (Despax) 120 Eleutherodactylus parvillus Lynch 121 Eleutherodactylus phoxocephalus Lynch 122 Eleutherodactylus pteridophilus new species 124 Eleutherodactylus pyrrhomerus Lynch 126 Eleutherodactylus quincpiagesimus Lynch and Trueb 127 Eleutherodactylus rosadoi Flores 129 Eleutherodactylus ruidus Lynch 130 Eleutherodactylus scolodiscus Lynch and Burrowes 131 Eleutherodactylus simonbolivari Wiens and Coloma 132 Eleutherodactylus siopelus Lynch and Burrowes 133 Eleutherodactylus sobetes Lynch 133 Eleutherodactylus subsigillatus (Boulenger) 134 Eleutherodactylus sulculus Lynch and Burrowes 135 Eleutherodactylus surdus (Boulenger) 136 Eleutherodactylus taeniatus (Boulenger) 138 Eleutherodactylus teuebrionis Lynch and Miyata 138 Eleutherodactylus thymalopsoides Lynch 139 Eleutherodactylus truebae new species 140 Eleutherodactylus unistrigatus (Giinther) 142 Eleutherodactylus verecundus Lynch and Burrowes 143 Eleutherodactylus vertebralis (Boulenger) 144 Eleutherodact\'lus w -nigrum (Boettger) 145 Eleutherodactylus walkeri Lynch 148 ECOLOGY 150 The Eleutherodactylus Way of Life 150 Eleutherodactylus communities 150 Discussion 159 BIOGEOGRAPHY 165 Patterns of Distribution 165 Patterns of Speciation 176 Historical Biogeography 182 Comparisons with Other Regions 185 Comparisons with Other Taxa 188 FUTURE RESEARCH 190 LITERATURE CITED 191 APPENDICES 197 I. Specimens Examined 197 II. Gazetteer 208 III. Species of Eleutherodactylus 219 IV. Distributional data 235 PLATES 1-8 Following page 76 IV ABSTRACT Western Ecuador encompasses the Pacific lowlands, a series of low coastal mountains, and the Andes, which rise to heights of more than 6000 m. Within this region, bioclimatic regimes range from humid tropical rainforest to desert on the lowlands, from subtropical dry and subtropical humid environments on the lower slopes of the Andes to humid temperate and subtemperate forests, subparamo, and paramo at high elevations in the Andes. This region is inhabited by 61 species of Eleiitherodactylus, each of which is treated in an account that includes a diagnosis, description or reference to a description, coloration in life, natural history, and distribution. Nine new species are described, and E. alberchi is placed in the synonymy of £. surdus. Records for E. taeniatits in Ecuador are based on misidentifications. All morphological characters used in the identifica- tion and systematicsof£'/£^//r//eraJ«crv/t<5 are defined and illustrated, and a key to the identification of the species in western Ecuador is provided. Nine communities containing six to fifteen species of Eleiitherodactylus were analyzed with respect to species diversity, rank and relative abundance of species, body size, and microhabitat untilization. Fidelity of species to bioclimatic regimes is rather low; the highest degree of endemism is 26% in the humid subtropical regime. Among the Eleutherodactyhis in western Ecuador, 32 species are endemic to the region, and 24 other species reach the southern limits of their distributions there. Altitudinally, the greatest number of species is between 1 200 m and 2200 m. Sizes of distributions are large at low elevations compared to smaller ranges of species in the Andes. Patterns of speciation include upland vicariants of lowland relatives, latitudinal replacement of sister taxa in the Andes, and trans-Andean relatives. Key words: Ecuador; Pacific versant; Leptodactylidae; Eleiitherodactylus; Taxonomy; New species; Commu- nity ecology; Biogeography. RESUMEN La parte oeste de Ecuador abarca las tierras bajas del Pacifico, una serie de montafias bajas costefias, y los Andes que suben a elevaciones mayores de los 6000 m. Dentro de esta region los regimenes bioclimaticos varian desde selva hiimeda tropical hasta desierto en las tierras bajas, desde ambientes secos subtropicales hasta humedos subtropicales en las pendientes bajas de los Andes, y hasta bosques templados y subtemplados, subparamo, y paramo en las elevaciones altas de los Andes. Esta region es habitada por 61 especies de Eleiitherodactylus, cada una de estas se cubre en un sumario que incluye diagnosis, descripcion o referenda a una descripcion, coloracion en vida, historia natural, y distribucion geografica. Se describe nueve especies nuevas y se coloca E. alberchi en la sinonimia de E. surdus. Los registros de E. taeniatus en Ecuador han sido basados en identificaciones erroneas. Todos los caracteres morfologicos utilizados en la identificacion y sistematica son definidos e ilustrados, y se provee una clave para la identificacion de las especies en el oeste del Ecuador. Nueve comunidades que contienen de seis a quince especies de Eleiithe rodactylus fueron analizadas con respecto a diversidad, grado de abundancia y abundancia relativa de especies, tamaiio del cuerpo, y utilizacion de microhabitat. La fidelidad de especies a regimenes bioclimaticos es algo bajo; el grado mas alto de endemismo es 26% y se encuentra en el regimen hiimedo subtropical. De las especies de Eleutherodactyhis en el oeste del Ecuador, 32 son endemicas a la region, y las otras 24 especies llegan a su Ifmite sur en esta region. Altitudinalmente el mayor niimero de especies se encuentran entre 1100 m y 2200 m. El tamaiio de las distribuciones es grande en las tierras bajas en comparacion con las extensiones mas pequehas de las especies en los Andes. Los patrones de especiacion incluyen vicariantes en las tierras altas que tiene parientes en las tierras bajas, reemplazo latitudinal de especies hermanas en los Andes, y parientes transandinos. Palabras claves: Ecuador; region Andina-Pacffica; Leptodactylidae; Eleiitherodactylus; Taxonomia; Especies nuevas; Ecologia de comunidades; Biogeografia. INTRODUCTION Anyone who has ventured into the humid low- tion and rocks along streams with a headlight land rainforests of northwestern Ecuador on into immediately reveals to the visitor a variety of frogs the cloud forests on the Pacific slopes of the Andes of the genus Eleiitherodactylus, the most speciose in Ecuador at night has been greeted by a variety of group of vertebrates in the neotropics, with more "chirps" and "peeps." Inspection of the low vegeta- than 500 species recognized (Duellman, 1 993). In UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 many places in the neotropics, especially in Andean cloud forests and adjacent lowland rainforests, these are the most diverse and abundant anurans. As such, they should be ideal subjects for ecologi- cal studies and monitoring of populations. How- ever, such has not been the case, principally be- cause these frogs have been a notoriously difficult group taxonomically. We anticipate that this di- lemma can be remedied by thorough and precise taxonomic treatments of members of the genus in various geographic regions. We began our investigations of the Ecuadorian herpetofauna in the mid-1960s. Duellman worked sporadically in western Ecuador from 1967-1984. Lynch's field work there in 1967-1979 focused almost exclusively on Eleuthewdactyhis, and al- though his interests shifted to the rich Colombian fauna about 1 5 years ago, he retained an obligation to summarize the Eleiitherodactylus fauna of Ecua- dor. Four summaries of the Eleuthewdactylus of geographical regions of the country have ap- peared— the Andes of southern Ecuador (Lynch, 1979a), the Amazon Basin (Lynch, 1980a), the Andes of northern Ecuador (Lynch. 1 98 1 a), and the Amazonian slopes ofthe Andes of Ecuador (Lynch and Duellman, 1980). Most recently, we (Lynch and Duellman, 1995) described E. philipi from paramos in southern Ecuador. The present work is a summary of the Eleiitherodactylus of the Pacific lowlands and the western slopes oftheAndesofEcuador. Herein, the upper altitudinal limit is arbitrary; it is defined not by elevation but, rather, by assemblages of Eleuthewdactylus. Some species (e.g., E. curtipes) from the paramos do encroach into the upper cloud forests, but we do not include these species; they were treated by Lynch ( 1 98 1 a). Our intent here is to treat the species confined to forested regions of western Ecuador. This small fragment of the total range ofthe genus contains more than 12% ofthe species oi Eleuthewdactylus. In addition to a thor- ough taxonomic treatment, we examine patterns of distribution with respect to climate, vegetation, topography, and historical biogeography. We also focus on several eleuthero-dactyline communities in western Ecuador and compare these with com- munities elsewhere. In so doing, we hope to pro- vide the ground work for future studies on the ecology of eleutherodactyline frogs in northwest- ern South America. A seminal paper on Eleuthewdactyhis in west- ern Ecuador was that in which Boulenger ( 1898) described six new species (as Hylodes and Syrrhophus) collected by W. F. H. Rosenberg for the British Museum. Of the 61 species now known from western Ecuador, only 16 of these were de- scribed prior to our work in the region. Lynch (and coworkers) described 32 species, and nine are named herein; three other species {E. hamiotae Flores. E. wsadoi Flores, and E. sunonholivari Wiens and Coloma) have been named in recent years. Eight ofthe 1 1 species of Eleuthewdactylus now known from cloud forest at Tandapi, Ecuador, were treated by Lynch ( 1 976a). Lynch and Buirowes (1990) reported on the Eleuthew-dactylus in the cloud forest at La Planada in southwestern Colom- bia; ofthe eight species described as new (and then known only from Colombia), we have found six in collections from Ecuador. Some of the most com- mon species on the Pacific lowlands of Ecuador are members of the paraphyletic Eleuthewdactylus "fitzingeri " group of Lynch (1976) and Lynch and Myers (1983), the Chocoan members of which were treated monographically by Lynch and Myers (1983). Most ofthe species known from western Ecua- dor have been described, and we do not repeat descriptions in the accounts of those species; rather, we provide references to adequate descriptions. Complete descriptions are given for the nine spe- cies named herein. We provide a redescription of Eleuthewdactylus gularis (Boulenger) because existing descriptions (Boulenger, 1898; Cochran and Coin, 1970) are inadequate and because more than one species has been confounded under that name. For all species, we provide lengthy, num- bered diagnoses, including comparisons and com- ments, even when doing so results in duplication of material published elsewhere; we have done this to facilitate comparisons and identifications. Accounts of two species, E. siopelus and E. sulculus that are known from extreme southwestern Colombia but yet to be recorded from Ecuador, are included because we are confident that they will be found in Ecuador. Our other departure is the inclusion ofthe only other eleutherodacty lines (Barycholos pulcher ELEUTHERODACTYLUS IN WESTERN ECUADOR and Phyllonastes sp.) known from western Ecua- dor in the key and in comparisons in appropriate diagnoses. This is the last in a series of papers on Ecuador- ian eleutherodactylines, and we take this opportu- nity to review, and expand on, the characters we find to be most useful in the identification of spe- cies. This concern for well-defined characters re- flects Lynch's interest in the phylogenetic relation- ships within the genus and the desire to have groups of species defined by shared derived characters rather than by phenetic similarity. The initial under- standing of relationships within Eleiithewdactylus came with the recognition that the subgenus Craugastor could be diagnosed by a derived con- dition of the m. adductor mandibulae (Lynch, 1 986). That discovery caused JDL to question (and ulti- mately to reject) many of his earlier groupings of Eleutherodoctylus and to continue to search for synapomorphies for various units within the genus. Acknowledgments More than a quarter of a century has passed since we initiated our work in Ecuador. During that time we have built up a debt of gratitude to many persons who contributed directly or indirectly to the completion of this work. We are especially grateful to our field companions whose efforts greatly enhanced the collection of specimens and field data. At various times, JDL was accompanied by Thomas J. Berger, David C. Cannatella, Thomas H. Fritts, Robert W. Hen-derson, Marsha C. Lynch, and Gerald R. Smith. Duellman variously was accompanied by Patricia A. Burrowes, Dana T. Duellman, David M. Hillis, Bruce MacBryde, Alan H. Savitzky, John E. Simmons, and Linda Trueb. Duellman's field work was accomplished as part of research supported by the National Science Foun- dation (Grants DEB 74-02998, 76-09986, and 82- 1 9388 ) and the National Geographic Society (Grant 1304); the biogeographic analysis herein is part of a project on patterns of anuran speciation and biogeography supported by the National Science Foundation (Grant BSR 8805920). Lynch's field work was made possible by grants from the Society of the Sigma Xi, Watkins Fund of the Natural History Museum, and the University of Nebraska Research Council. Funding for the color plates was provided by the School of Biological Sciences, University of Nebraska, Lincoln. For the loan of specimens and/or providing working space in their respective institutions, we are grateful to Darrel R. Frost, Charles W. Myers, and Richard G. Zweifel, American Museum of Natural History: Alice G. C. Grandison, British Museum (Natural History); Alan E. Leviton, Cali- fornia Academy of Sciences; Robert F Inger. Field Museum of Natural History; Jose R Rosado and Ernest E. Williams, Museum of Comparative Zool- ogy, Harvard University; Josef Eiselt, Natur- historisches Museum, Wien; Dorothy Smith, Uni- versity of Illinois Museum of Natural History; Arnold G. Kluge, Ronald A. Nussbaum. and the late Charles F. Walker, University of Michigan Museum of Zoology; the late Doris M. Cochran. W. Ronald Heyer, Roy W. McDiarmid, the late James A. Peters, and George R. Zug, National Museum of Natural History. We acknowledge the cooperation ofAnaAlmendariz(EscuelaPolitecnicaNacional), Luis A. Coloma (Pontificia Universidad Catolica del Ecuador), and Alexandra Quiguango (Eco- ciencia) for loaning specimens from Ecuadorian collections and providing data on localities in Ec- uador. Coloma and Martha L. Crump, who acted as liaison with Ecociencia, provided color slides, as did Patricia A. Burrowes and Roy W. McDiarmid. Credits for photographs in Plates 1-8 are noted in the legends as: JDL = John D. Lynch, KM = Kenneth Miyata, LAC = Luis A. Coloma, MLC = Martha L. Crump, PAB = Patricia A. Burrowes, RWM = Roy W. McDiarmid. WED = William E. Duellman. We thank Patricia A. Burrowes and Erik R. Wild for helpful suggestions on the analysis of the ecological data, Joseph R. Mendelson III for checking specimens against our list of specimens examined, Santiago F Burneo and Ignacio de la Riva for providing the Spanish version of the key. and Rafael Joglar for assistance with the Spanish Resumen. We also thank Eugenia del Pino and Fernando Ortiz C, who generously provided facili- ties for our use at the Universidad Catolica in Quito, and to Sergio Figueroa and Abel Tovar V., Ministerio de Agricultura y Ganaderfa. for issuing permits for collection and exportation of speci- mens. UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 An earlier version of this manuscript benefitted from thoughful comments by Martha L. Crump, throrough review by Luis A. Coloma. and critical evaluation by Jay M. Savage, who encouraged us to eschew certain hectoring terms in our discussion of anatomy. To these three persons in particular we extend our profound thanks. We especially thank Linda Trueb. whose careful editing and assistance with the electronic transfer of illustrations resulted in the improvement of our otherwise benign ef- forts. Lastly, we dedicate this paper to the memo- ries of two deceased friends and colleagues — Kenneth Miyata and James A. Peters; they loved Ecuador and collected many of the specimens on which this work is based. MATERIALS AND METHODS Specimens are identified by standardized mu- seum codes designated by Leviton et al. (1985), except for the addition of four collections housed in Quito, Ecuador: ECO = Ecociencia (specimens to be deposited in a museum), EPN = Escuela Politecnica Nacional, MECN = Museo Ecuatori- ano de Ciencias Naturales, and QCAZ = Museo de Zoologfa, Pontificia Universidad Catolica del Ec- uador. Some specimens collected recently by Ana Almendariz are identified by field numbers (e.g., EPN- A A). References are made to some speci- mens in Colombian collections: ICNMHN = Instituto de Ciencias Naturales, Museo de Historia Natural, Universidad Nacional de Colombia, Bogota; IND- AN = amphibian collection, Instituto Nacional de los Recursos Naturales Renovables y del Ambiente, Bogota; UVC = Universidad Valle de Cauca, Call. All specimens from western Ecuador that have been studied are listed in Appendix L Localities from which specimens have been examined are listed with their geographic coordinates and eleva- tions (when known) in Appendix IL Localities that have been located are plotted on distribution maps in the Accounts of Species; because of crowding of symbols, some closely approximated localities are not shown. Methods for taking measurements and ascer- taining structural features are defined in a follow- ing section. Description of Characters. Measure- ments were taken with dial calipers to the nearest 0. 1 mm; measurements, and proportions are given for each sex when their means differ significantly (P< 0.05); otherwise these data are combined. The following abbreviations are used: E-N = eye-nos- tril distance; HL = head length, HW = head width, lOD = interorbital distance, SVL = snout-vent length. Areas of distribution were measured from maps using a Micro-Plan II image analysis system (Laboratory Computer Systems, Inc., Cambridge, MA). Throughout the text, we have avoided using the terms senior and junior author, because in this case the junior is senior chronologically; therefore, the authors are noted simply as JDL and WED. Al- though we both assume responsibility for the ma- terial presented herein, JDL is the primary author for the sections on systematics, and WED is the primary author for the sections on western Ecua- dor, ecology, and biogeography. All morphological illustrations were drawn by JDL using a dissecting microscope equipped with a camera lucida. All graphs and maps were generated by WED using Aldus Freehand® on a Macintosh computer. The color photographs were reproduced electronically from color transparencies. WESTERN ECUADOR Western Ecuador encompasses the area from Colombia to Peru and from the Pacific Ocean to the western crest of the Andes; this region includes all of the provinces of Bolivar, El Oro, Esmeraldas, Guayas, Los Rios, and Manabi, and the western parts of the provinces of Azuay, Carchi, Cafiar, Chimborazo, Cotopaxi, Imbabura, Loja, and Pichincha (Fig. 1). In order to provide a working basis for the ecological and biogeographic analyses of the Eleutherodactylus of the region, we provide ELEUTHERODACTYLUS IN WESTERN ECUADOR Fig. 1. Western and central Ecuador showing provinces and major cities. UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 2. Western Ecuador showing physical features. ELEUTHERODACTYLUS IN WESTERN ECUADOR Fig. 3. Profiles of western Ecuador at 0°S to 4°S Latitude. Note the varying widths of the Pacific lowlands, the precipitous slopes of the Andes, and the complex lower topography at 4°S Latitude, which is the northern part of the Huancabamba Depression. Based on Mapa Pisco 1:2,000,000, 1971, Instituto Geografico Militar, Quito, Ecuador. Vertical exaggeration = 20x. 8 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 synopses of geography, climates, and bioclimatic regimes of the region, as well as a brief synopsis of its geological and climatic history and comments on changes during historic times. The complexities of physiography and climate cannot be appreciated readily by someone who has not descended the precipitous slopes from the lofty Andes, where many peaks within one degree of the Equator have permanent snow, to the Pacific lowlands. In fact, the snow-covered summit of one of the highest peaks in South America, Volcan Chimborazo at a height of 6310 m, is only 50 km airline from the 300-m contour at the base of the Andes. In such a transect, mean annual temperatures increase from below 0°C to more than 25°C. Likewise, on the Pacific Coastal Plain, majestic rainforests receiv- ing more than 3500 mm of rain annually gradually give way southward to deserts receiving less than 200 mm of rainfall annually. Physiography Western Ecuador can be divided into three physi- ographic regions, each of which is described below (Fig. 2). Pacific Lowlands. — The lowlands between the Pacific Ocean and the base of the Andes at an elevation of approximately 600 m vary in breadth from about 200 km at Punta Santa Elena (02° 1 1 ' S ) to about 40 km in northern Provincia Esmeraldas; however, the lowlands are even narrower — in places no more than 20 km wide — south of the Golfo de Guayaquil. The terrain is nearly flat, especially in the south and northwest, whereas low, rolling ter- rain is encountered in southern Provincia Esmeraldas and northwestern Provincia Pichincha. The coastal region is swampy along the borders of the Golfo de Guayaquil southward nearly to the Peruvian border. Cordillera de la Costa. — Nearly paralleling the coast and inland for distances of 5-60 km from just south of Esmeraldas to just west of Guayaquil is a range of low mountains that is interrupted at about 01 °S Lat. Local names are applied to various higher parts of this system of eroded ridges. Most of these highlands are less than 600 m in elevation (Fig. 2). Although there are some higher regions to the north and south, the largest continuous high- land area (>600 m) extends from 00° 12' N Lat. to 00°25' S Lat. in northern Provincia Manabi; in this range, Cerro de Pata de Pajaro rises to 800 m, and Cerro San Sebastian to 780 m. To the north in the Montaiias de Muisne south of Esmeraldas, the highest ridges are at elevations of 600-800 m, whereas in the extreme south in the Cerros de Chongon, few ridges are above 400 m. Andes. — The Andes are the dominant physi- ographic feature in Ecuador; this high volcanic range extends the length of the country and is divided into eastern (Cordillera Oriental) and west- ern (Cordillera Occidental) ranges separated by inter-Andean basins and connected, or nearly so, by transverse ridges. For a detailed description, see Duellman (1979). The Andes along the western versant of Ecuador consist of three distinct regions: 1 . To the north of the Rio Chota, the Andes in extreme northwestern Ecuador are part of a mass, the Nudo de Pasto centered in southern Colombia; the highest peak is Volcan Chiles (4723 m) on the Ecuadorian-Colombian border. 2. South of the Rio Chota Valley, the major range of the Cordillera Occidental extends south- ward to the Rio Jubones Valley. The entire range exceeds elevations of 3000 m, and there is one continuous area more than 100 km long that ex- ceeds 4000 m. The highest volcanoes in this range (from north to south) are Yanahurco de Pinan (4535 m), Cotacachi (4944 m), Rucu-Pichincha (4324 m), Guagua-Pichincha (4675 m), Atacazo (4463 m), Corazon (4788 m), Illiniza Norte (5126 m), Illiniza Sur (5248 m), Quilotoa (39 14 m), Sagoatoa (4153 m), Carihuayrazo (5020 m), Chimborazo (6310 m), Chanlor (4300 m), PatuI (4163 m), and Minas (4096 m). Except in the northern part of this range, the western slopes are extremely precipitous (Fig. 3). 3. To the south of the Rio Jubones southward into northern Peru, the mountains are lower and fragmented into complex series of north-south or northeast-southwest ranges that are separated by valleys, many of which are at elevations below 1500 m (Fig. 3). This complex system of low ranges and basins collectively is referred to as the Huancabamba Depression, wherein exists the low- est breach in the Andes (Abra de Porculla at 2145 m in northern Peru) between Colombia and south- ELEUTHERODACTYLUS IN WESTERN ECUADOR ern Chile. Along the western versant of the Huancabamba Depression in Ecuador, the highest mountains are in the Cordillera de Chilla (Cerro Chillacocha, 3580 m) and in the Cordillera de Celica with one long ridge above 3000 m. River Systems As might be expected from the topography, drainage systems in western Ecuador are complex (Fig. 2). In the lowlands in northwestern Ecuador, the major rivers flow north or northwest. One of the major drainages is the Rio Mira system, the mouth of which is in Colombia. In the headwaters of this system is the Rio Chota originating in the Ibarra Basin and forming a depression between the Nudo de Pasto and the Cordillera Occidental. Other ma- jor affluents of the Rio Mira are the Rio San Juan on the Colombian-Ecuadorian border and the Rio Lita, tributaries of which drain the northernmost part of the Cordillera Occidental. The broad, meandering Rio Esmeraldas has many affluents draining the northwestern Pacific lowlands. Two of these, the Rio Guayllabamba and the Rio Blanco, drain the Pacific slopes of the Andes from about 0°30' N Lat. to 1°S Lat. Major rivers flowing into the Rio Guayllabamba include the Rio Nanegal, Rio Pachijal, and Rio Pitsara. The headwaters of these streams are narrowly separated from the Rio Blanco, which has many tributaries (e.g., Rio Mindo, Rio Saloya, Rio Toachi, and Rio Pilaton) draining the slopes of the Andes. Between the Cordillera de la Costa and the Andes southward from about 0°15' S Lat., the major rivers flow southward to the Golfo de Guayaquil. The major rivers are the Rio Daule and the Rio Babahoyo, which converge just north of Guayaquil to form the broad Rio Guayas. Most of the rivers draining the adjacent precipitous slopes of the Andes are rather short; major exceptions are south of 2°S Lat. and include the Rio Chimbo-Rio Coco and the Rio Chanchan. Southward and drain- ing directly into the Pacific Ocean are numerous small rivers and two major rivers, the Rio Jubones and the Rio Puyango, both of which extend far inland in deep valleys in the Andes. The western slopes of the Cordillera de la Costa are drained by many small rivers flowing into the Pacific Ocean. The largest ofthese, Rio Calceta and Rio Portoviejo, separate the northern and southern parts of the Cordillera de la Costa. Climate Irrespective of the effects of altitude, the climate in western Ecuador is influenced by two drastically different ocean currents. The cold Humboldt Cur- rent originating in the Antarctic Ocean sweeps northward along the west coast of South America to about 2°S Lat. where it swings westward away from the land. Most of the moisture in onshore winds condenses as fog offshore, so that rainfall is slight on the Pacific lowlands and only moderate at higher elevations. Contrariwise, the warm equato- rial current. El Nino, flows southward along the coast of Colombia and northern Ecuador before turning westward. The onshore winds are heavily laden with moisture, which condenses over the lowlands and continues to do so as the air masses rise along the face the Andes. This results in heavy rainfall throughout the year in northwestern Ecua- dor and moderately heavy rainfall throughout the year on the Andean slopes north of 1°S Lat. In January-March, the equatorial current shifts farther south; consequently, during these months, rain falls more frequently and heavily throughout the region, and it is at this time that most, if not all, of the rain falls in the lowlands south of 2°S. Lat. (Fig. 4). However, even on the Andean slopes, there is a marked decrease in rainfall in June-August; for example, at Lita (00°50' N, 78°27' W; 570 m), rainfall exceeds 300 mm per month in October- April, but it drops to less than 200 mm per month in June-August. Likewise, farther south and also at higher elevations the same annual pattern prevails, but the rainy period is shorter; for example, at Pilalo (00°56' S, 79°00' W; 2400 m), rainfall ex- ceeds 125 mm per month in December- April, but it declines to less than 40 mm per month in July and August. The western slopes of the Andes are character- ized by a humid adiabatic lapse rate that, because of lower evaporation rates at higher elevations, re- sults in high humidity even though rainfall is less than at lower elevations (Table 1). Dense cloud banks are almost of daily occurrence at various 10 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 500 400 300 200 100 / / ^ V ( / \ Cayapas (3804 mn- ) \ Sto. Domingo de los Colorados (2906 mm) \ \ ly y » < h— "" "* K « '— —1 ^-H •■ — -1 1 \ /> Rocafuerte (519 mm) \ \ / / ^ \ \ / ^ < \ --•^ ^ ^ \ \ > ►• — —4 » "^ t '^ ^ \ r Salinas (68 mi n) " ^ < *"'" > — ~ —4 1 ^ _^ i»« » JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Month Fig. 4. Annual patterns of rainfall on the Pacific lowlands of Ecuador. Note the decrease in rainfall from north to south. Cayapas is at 00°50' N, 79°01' W, 65 m; Santo Domingo de los Colorados is at 00° 15' S, 79° 10' W, 600 m; Rocafuerte is at 00°55' S. 80°27' W. 10 m; Salinas is at 02° 11' S. 80°59' W, 6 m. Data from Cafiadas (1983). elevations along the western slopes of the Andes. Climatic data are not available for sites in the Cordil- lera de la Costa. However, at elevations of 700-800 m on Cerro Sebastian, just inland from Machalilla, Provincia Manabi, Foster ( 1 992a: 36) commented on the moisture in the fog forest in January: "With every passing cloud that comes up against the ridge, the forest starts dripping heavily, very much as if it were raining. Stepping out into the clearing, however, one immediately finds that there is no rain at all." Throughout all but the southern part of western Ecuador the great physiographic relief results in rapidly declining temperatures from average annual temperatures of more than 25°C in the lowlands to below 0°C on the highest snow-covered volcanoes (Fig. 5). Annual variation in monthly mean tem- peratures is less than 3°C in the humid tropical lowlands (e.g., 25-27°C at Cayapas) but some- what greater in arid tropical lowlands (e.g.. 21- 26°C at Salinas). Likewise, the annual variation in monthly means is minuscule at sites on the western slopes of the Andes — for example, 21 -23°C at Lita (570 m), 16-18°C at Chiriboga (1910 m), and 13- 1 4°C at Pilalo (2400 m). In contrast, daily fluctuations are greater than the variation in monthly means. Temperatures obtained from a maximum-minimum thermometer at actual collecting sites in 1975 are given in Table 2. ELEUTHERODACTYLUS IN WESTERN ECUADOR 11 Table I. Climatic data for selected sites in western Ecuador (exerpted from Cafiadas, 1983). Mean Mean annual annual Days temp. precip. without Ecological Site Lat. & Long. Elev. (m) (0°C) (mm) rain regime Salinas, Guayas 02°irS, 80°59'W 6 23.0 68 233 Desert Guayaquil, Guayas 02°12'S, 79°53'W 6 25.0 820 172 Dry tropical Salinas, Imbabura 00°30' N, 78°06' W 1730 19.4 307 205 Dry subtropical Ibarra, Imbabura 00°21' N, 78°06'W 2228 15.4 589 60 Dry temperate Muisne, Esmeraldas 00°37'N, 80°01' W 6 24.7 1574 43 Suhumid tropical Cayapas. Esmeraldas 00°50'N, 79°01'W 65 25.6 3804 — Humid tropical Lita, Imbabura 00°50' N, 78°27' W 570 22.4 3678 — Humid subtropical Pilalo, Cotopaxi 00°56' S, 79°00' W 2400 18.0 1408 18 Humid subtemperate Otavalo, Imbabura 00°14'N, 78°16' W 2556 14.4 735 41 Humid temperate 3000 2000 c o > 1000 Tuican Guanujo •• Aloag Guaranda \ Calacali Nono — • • Pilalo Chiriboga Apuela Pallatanga • Nanegalito Maldonado • ^indo | Tandapi • •. * Gualea Facto Alluriquin Lita • / Sto. Dom. d 1 Colorados • Buenavista • Bucay Quevedo Machala • Cayapas v^^—'\^ ^Naranjal 10 12 14 16 18 20 22 Mean Annual Temperature (°C) 24 Fig. 5. Mean annual temperatures at selected sites over an elevational range in western Ecuador. See Appendix II for locations. Data from Caiiadas (1983) and Indice Toponimico de la Repiiblica del Ecuador (1972). 12 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Table 2. Daily fluctuations in air temperatures (°C) at collecting sites in western Ecuador in 1975. Site Elevation Dates Min. temp. Max. temp. C. C Rio Palenque, Los Rios 8 km ESE Pto. Quito, Pichincha I8km WPiiias, El Oro 4 km E Dos Rios. Pichincha 3.5 km NE Mindo. Pichincha 14 km W Chiriboga, Pichincha Quebrada Zapadores, Pichincha 9 km SE Tandayapa, Pichincha 9.5 km NW Nono, Pichincha 14 km SW Tulcan, Carchi 220 m 27-31 Mar. 20.0 29.5 530 m 06-07 Apr. 20.5 27.5 780 m 06-07 Mar. 19.0 25.0 1140 m 01-02 Apr. 14.5 20.5 1 540 m 07-08 Apr. 15.0 25.0 1960 m 08-09 May 8.5 23.0 2010 m 03-04 Apr. 10.5 22.0 2150 m 09- 10 Apr. 10.0 23.0 2530 m 11-12 Apr. 10.5 19.5 3340 m 18-19 May 0.0 18.0 BiocLiMATic Regimes The concept of life zones based on temperature, precipitation, and evapotranspiration rates proposed by Holdridge ( 1947) for recognizing plant forma- tions and elaborated upon by Holdridge ( 1 967 ) was applied to Ecuador by Canadas ( 1 983). Although in theory the life zones are defined by abiotic factors, in practice, they are recognized by characteristic flora. A weakness of the system is that edaphic differences are not taken into consideration; conse- quently, areas having similar climates but drasti- cally different soils (e.g., sandy karst vs. deep alluvial) will be placed in the same bioclimatic category. For Ecuador, the Holdridge system was modi- fied and refined by Caiiadas (1983), who included data on soils and topography. We agree with Sav- age (1975) that the Holdridge system seems to be too sophisticated for determining broad vegeta- tional correlates to animal distributions. Thus, al- though we follow Caiiadas's classification (Fig. 6), we simplify it by combining many of his categories (Table 3). We use Caiiada's terms "temperate" and "subtemperate" in his sense of temperature; these terms do not refer to temperate zone climates. The 10 bioclimatic regimes that we recognize are defined below; they are arranged from tropical to temperate or cooler within dry to wet regimes (Fig. 7). The type of vegetation and characteristic plant taxa are given for each regime. For ease of comparison, we include the synonymous terms of the Holdrige system. Unless noted otherwise, data are summarized from Cafiadas (1983). Desert. — Extremely arid conditions exist in two coastal regions in southwestern Ecuador. These isolated deserts lie at elevations below 300 m and receive less than 200 mm of rainfall annually, mostly in February-May. The soils are mostly sandy, but some areas have clay, which character- istically has deep cracks throughout the long dry season. The sandy soils support the characteristic candelabra cactus (Lemairocereus cartwrighti- anus) and sparse grasses of the genera Panicium, Chloris, and Eragrostis. On the low hills, small trees, such as Bursera graveolens and Capparis angidata, occur. In the Holdridge system, this regime includes lowland desert and lowland desert scrub. Dry tropical. — Most of the Pacific lowlands south of 1°S Lat. are in this category, which also extends northward along the western lowlands to 0°15' N Lat.; discontinuous areas of this regime also occur along the northwestern coast. Rainfall is lowest (<500 mm annually, mostly in January- April) along the coastal regions and becomes pro- gressively higher inland, where annual rainfall may approach 1500 mm, most of which falls in December-mid-May. The soils are sandy; in some places along the coast the sand is heavy, dark, and deep, and in other places along the coast the soils are alkaline. Clays predominate inland and along river valleys toward the coast. There is a gradual change in vegetation from low (<3 m), scrub along much of the coast to dense scrub forest (5-10 m high) inland; in both areas the trees and bushes are primarily deciduous. In the ELEUTHERODACTYLUS IN WESTERN ECUADOR 13 Paramo 0-6° Humid Very Humid Rainy Very Rainy Subtemperate 6-12° Subhumid Humid Very Humid Rainy Very Rainy Pluvial Temperate 12-18° Dry Subhumid Humid Very Humid Rainy Very Rainy Subtropical 18-22° Very Dry Dry Subhumid Humid Very Humid Rainy Tropical 22-26° Desert Subdesert Very Dry Dry Subhumid Humid Very Humid A Regime; Temperature Precipitation <200 200-500 500-1000 1000-1500 1500-2000 2000-3000 >3000 Fig. 6. Classification of bioclimatic regimes (ecological zones) of Ecuador (Canadas. 1983). former, characteristic trees are Bursera graveolens and Jacquinia pubescens. whereas in the latter, trees (e.g., Cordia lutea, Prosopis juliflora, and Capparis crotonoides) are common. Extensive ar- eas of mangroves occur along the coast in the northwest and around the Golfo de Guayaquil; the alkaline soils along the coast support a distinctive flora dominated by bushes — Mollugo, Mesembry- antheum, and Portulaca oleracea. The areas of highest rainfall (> 1 000 mm annually) support semi- deciduous forest with some trees attaining heights of 20-25 m; characteristic trees are Ceiba trichistandra (endemic, but absent in the north- west), Bombax ruizii, Tabebuia chrysantha, and palms of the genera Astrocaryum, Attalea, and Phytelephas. In places where the water table is high, there are many false bananas (Heliconia). In the Holdridge system, this regime includes lowland thorn woodland, semidry forest, and dry forest. Dry subtropical. — The major regions occu- pied by this regime are at elevations above 300 m in the Cordillera de la Costa, the slopes of the Andes to elevations of about 2000 m south of 2°S Lat., and the Catamayo and Loj a basins in southern Ecuador. The dry subtropical regime also exists in the Rio Table 3. Comparison of bioclimatic regimes recognized herein and by Canadas (1983). Herein Cafiadas (1983) Desert Dry tropical Dry subtropical Dry temperate Subhumid tropical Humid tropical Humid subtropical Humid temperate Humid subtemperate Paramo Desert Subdesertic, dry and very dry tropical Dry, very dry, and subhumid subtropical Dry and subhumid temperate Subhumid tropical Humid and very humid tropical Humid, very humid, and rainy subtropical Humid, very humid, rainy, and very rainy temperate Humid, very humid, rainy, and very rainy temperate Humid, very humid, rainy, and very rainy paramo UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 7. Bioclimatic regimes on Caiiadas(1983). the Pacific lowlands and western slopes of the Andes in Ecuador. Adapted from ELEUTHERODACTYLUS IN WESTERN ECUADOR 15 Chota Valley in the Ibarra Basin in northern Ecua- dor. Mean annual temperatures are 1 8-22°C. Rain- fall is lowest (<500m) in the Cordillera de la Costa between 1° 15' and 1°50'S. Lat., where rain occurs primarily in January-April; however, relatively high humidity is maintained by fog (Foster, 1 992a). Areas receiving 500-1000 mm of precipitation annually include the rest of the high elevations in the Cordillera de la Costa, some parts of the Catamayo Basin, the valleys of the Rio Chota and Rio Jubones. In the Cordillera de la Costa, rainfall is concentrated in December-April, but in the other areas, there are two rainy periods — one in Febru- ary-March or April and another in September or October-NovemberorDecember. Soils range from sandy clays to reddish clays in the Sierra de la Costa to clays and soils derived from volcanic ash in the Andes. The vegetation in this tropical dry forest con- sists of semideciduous trees with the largest trees attaining heights of 15-20 m. In the drier areas, characteristic trees are Cedrela sp., Jacquinia pubescens, and Tabebuia crysantha. In the Machalilla region of the Cordillera de la Costa, important trees are Erythrina megistophylla, Phytolacca sp., Palicoiirea sp., Poiilsenia armata, Quararibaea grandifolia, and the palms Chamae- dorea polychada and Phytelephas aequatohalis (Gentry, 1992). On the Andean slopes and in the Catamayo and Loja basins, characteristic trees include Acflc/fl macracantha, Brosimum latifo-lium, Centrolohium patinensis, Coultheria tinctoria. Pi- cas sp., Juglans neotropica, and Sapindus saponaria, plus palms of the genera Bactris and Phytelephas. Ground vegetation consists of grasses. Agave americana, and cacti — Cereus and Opuntia. Agave and Opuntia are especially prevalent in the Rio Chota Valley, and a peculiar cactuslike euphorb is common in the Rio Jubones Valley. Climbers of the families Bignoniaceae and Malpighiaceae are common, and the most abundant epiphytes are bromeliads of the genus Tillandsia. In some areas on the lower slopes of the Andes, dense cloud banks provide considerable moisture and low evapotranspiration rates, so that humidity and temperatures are more like that of the humid subtropical regime. Such is the case at elevations of 600-1240 m on the western slopes of Cerro San Jose Chico east of Manta Real (2°34' S, 79°2 1 ' W), Provincia Canar. The largest trees in this forest are Huertera glandulosa, Landen-bergia pavonii, Otoba sp.,Sapium sp., and Tetragastris panamensis; epiphytes are common and woody vines are un- common (Foster, 1992b). In the Holdridge system, this regime includes premontane desert scrub, thorn woodland, dry for- est, and moist forest. Dry temperate. — This bioclimatic regime is typical of many inter-Andean basins at elevations of 2000-3050 m, where mean annual temperatures are 1 2- 1 8°C. Annual rainfall is 200- 1 000 mm; the heaviest rains are in March-April and October- November, and the driest months are June-Sep- tember. Soils are sandy, volcanic ash, or a mixture thereof. The vegetation consists mostly of xerophytic, deciduous shrubs and low (<10 m) trees. Charac- teristic plants are Croton sp.. Datura stramonium, Inga sp., Juglans neotropica, Mimosa quitensis, Nicotania rustica, and Puya sp. In drier areas. Agave americana and Opuntia tuna are common, and the reed Scirpus totora occurs at the margins of lakes and rivers. In the Holdridge system, this regime includes lower montane thorn scrub and dry forest. Subhumid tropical. — This regime is interme- diate climatically, geographically, and floristic- ally between dry and humid tropical regimes. It occurs continuously up to elevations of 460 m between 0°20' and 2° 15' S Lat. and has isolated patches in the northwestern lowlands. Mean annual temperatures are 23-26°C. Annual rainfall is 1 500- 2000 mm, with the rainy season being in Decem- ber-June. The soils mostly are sandy gray or red clays, but in some inland areas volcanic sands also are present. This quasi-rainforest is composed of evergreen, broadleaf trees usually reaching heights of more than 20 m, with some emergents attaining heights of 30 m or more. Characteristic trees are Bombax sp., Centrolobium patinensis, Ficus sp., Myroxylon balsamum, Octoteasp.,Poulsenia armata, Pouteria sp., Tabebuia ecuadorensis, and palms — Astro- caryum sp., Phytelephas aequatori-alis, and Attalea colenda. Dominant climbers are members of the Bignoniaceae and Malpighiacea. False bananas 16 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 {Heliconia), elephant-ear plant.s (Xaiithosonia). and the large terrestrial, spiny bromeliad {Aechmea magdalemw) (iccur in wet areas. In the Holdridge system, this regime includes lowland moist forest. Humid tropical. — This regime supporting the Chocoan lowland rainforest extends southward from Colombia to about 1°40' S Lat.; in Ecuador, this regime reaches the Pacific coast only in the extreme northwest. A small area supporting this regime is at the base of the Andes at about 1 °50' S Lat. The maximum elevation attained is 600 m. Mean annual temperatures are 23-26°C. Through- out much of the region the mean annual rainfall is 2000-3000 mm, with the driest months (all of which receive more than 35 mm of rain) being August-November. Inland and north of the Equa- tor, rainfall is higher, with as much as 3804 mm recorded at Cayapas, where in the driest months (October-November) rainfall exceeds 1 20 mm per month. Soils are derived primarily from volcanic ash. The vegetation is classical lowland tropical rainforest having three principal strata with many woody vines, epiphyes, mosses, and lichens. The canopy is discontinuous at about 30-35 m above the ground; some emergents may exceed 40 m in height. This upper stratum is made up of many species of trees, including Brosimum utile, Dacwydes occidentalism Huberodendron pationoi, Humiriastnim procerum, Virola dixonii, and vari- ous Lauraceae. The middle stratum of the forest contains trees such as Callophyllum longifolium, Eschweilera pittieri, Platymiscium pinnatum, Terminaliaamazonica, various species of firownea, Nectondra, and Ocotea, and two dominant palms — Guilielma gasipaes and Iriartea corneto. The low- ermost layer of trees is discontinuous and com- posed of many species, among the most conspicu- ous of which are Herrania balaensis. Piper sp., Rollinia mucosa, Tetra-thylacium macrophyllum, Vismia obtusa, and Urea sp. In the Holdridge system, this regime includes lowland wet forest and rainforest. Humid subtropical. — This regime exists in the northern part of the Cordillera de la Costa and on the slopes of the Andes at elevations of 300-1800 m from the Colombian border southward to 1 °55' S Lat. Mean annual temperatures are 18-24°C. An- nual rainfall generally is 20()()-30(J() mm, but an area on the lower slopes of the Andes between the Equator and 1 °N Lat. receives more than 3000 mm. Lita, at an elevation of 570 mm, has an annual mean rainfall of 3678 mm. Seasonality of rainfall in- creases from north to south. At Lita (0°50' N Lat.) and Santo Domingo de los Colorados (0°15' S Lat.), only July is relatively dry — 120 mm of rain at Lita and 43 mm at Santo Domingo de los Colorados— but at Caluma ( 1°37' S Lat.) only 10- 22 mm of rain falls monthly during July-Novem- ber. Mostly, the soils are derived from volcanic ash and have the capacity to retain water. The forest in this regime commonly is referred to as cloud forest or lower montane rainforest. The forest is characterized by many ferns, vines, epi- phytes, mosses, and lichens. In many areas, three strata of vegetation are recognizable, but on steep slopes stratification is difficult to recognize. There is no continuous canopy; in many places palms (Attalea colenda and Iriartea corneto) are the high- est trees with fronds reaching 20 m above the ground. Somewhat lower are treeferns and a vari- ety of trees, including Brosimum utile, Clarisia racemosa, Clavija eggersi, Dactroydes sp., and Virola sp. Among the lowermost trees are Apeiba membranacea, Matista coloradorum, Poulteria sp., and Pouruma chocoana, among others. In the Holdridge system, this regime includes premontane moist forest, wet forest, and rainforest. Humid temperate. — With the exception of in- trusions by the dry valleys of the Rfo Chota in the north and the Rio Jubones in the south, this regime is a continuous narrow zone along the west face of the Andes at elevations of 1 800-3000 m from the Colombian border to 3°30' S Lat. Annual rainfall is 2000-3000 mm on the lower slopes but declines to 1000-1500 on the upper slopes. July-August are the driest months, but at Chiriboga (1910 m) more than 40 mm of rain falls in the driest month (July), and at Uyumbicho (2725 m) more than 30 mm of rain falls during the driest month (July). However, to the south the dry season is more intense and extends from June to November; for example, at Celica (4°7' S Lat., 2700 m), only about 10 mm of rain falls in June and August. However, at least on the western slopes of the Andes, high humidity is ELEUTHERODACTYLUS IN WESTERN ECUADOR 17 maintained by dense fogs that are almost of daily occurrence. The soils are primarily derived from volcanic sands and ash, but some clays are present; the soils characteristically contain a high amount of organic matter. This regime usually is referred to as high cloud forest or upper montane rainforest. Evergreen broa- dleaf trees attain heights of about 20 m, but at higher elevations trees may be dwarfed; trees com- monly are festooned with bromeliads, mosses, li- chens, and other epiphytes. Among the characteris- tic trees are Cedrela rosei. Cinchona sp., Croton sp., Didymopanaxmowtotoni, Eugenia sp., Guarea sp., Podocarpus sp., Polylepis sp., and Weinmannia descendens. Understory bushes include Baccharis sp., Bocconia frutescens, and in the south Emhothrium grandifoliwn. Ferns of the genera BlecJiniim and Dicksonia commonly are the most abundant ground cover. In some places, especially along streams, the vinelike bamboo, Chusquea scandens, forms almost impenetrable thickets. At the edges of the forest, a large-leafed herb. Gunnera, is conspicuous. In the Holdridge system, this regime includes lower montane moist forest, wet forest, and rainforest. Humid subtemperate. — This regime includes the uppermost forests and subparamos at eleva- tions of 3000^000 m throughout the Andes to about 3°30' S Lat. Mean annual temperatures are 6- 12°C. Throughout most of the region, annual rain- fall is 500-1000 mm, but at places on the upper western slopes of the Cordillera Occidental annual rainfall is 1 550-2000 mm; on these slopes atO°20'- 0°30' N Lat.. annual rainfall is 2000-3000 m. Throughout the regime rainfall is lowest in July and August. Most soils are volcanic sands and ash. At lower elevations, sometimes referred to as eltin forest, dwarf trees of the genera Clusia, Podocarpus, and Weinmannia, together with the bush Loricaria thuyoides, are present; commonly they are festooned with mosses. At higher eleva- tions, commonly called subparamo, grasses {Festuca and Stipa) and low herbaceous plants {Hypericum laricifolium and Vaccinium mortinia) are common, as are ferns of the genus Blechnum and low bushes of the genus Baccharis. In the area south of Tulcan, Senecio abietinus, Espeletia hartwegiana, and the spiny terrestrial bromeliad Puya are present. In the Holdridge system, this regime includes montane moist forest and rainforest. Paramo.— At the high elevations (4000-5000 m) on the volcanoes along the Cordillera Occiden- tal exists the highest life zone in the Andes. This regime is highly fragmented into zones on the slopes of the higher volcanoes and on some of the connecting ridges; the largest continuous area of paramo in the Cordillera Occidental extends from about 1° to 2°S Lat. Mean annual temperatures are 3-6°C, and annual rainfall is 500-1500 mm. The soil is mostly volcanic ash. The low vegetation in paramo consists mostly of low grasses, Bromus and Poa, and taller grasses, Festuca. Scattered among the grasses are Senecio and in some places extensive areas of cushion plants, a life form consisting of closely packed individuals of several unrelated species. Cushion plants in the central part of the Cordillera Oriental usually consist of five species — Arenaria dicranoides, Azorella sp., Draba bethomaniana, Plantago rigida, and Werneria humilis (Acosta- Solis, 1984). In wet areas mosses, Lycopodium and Sphagnum, also may be parts of cushion plants. In the Holdridge system, this regime includes wet paramo and Pluvial paramo. Vegetation Characteristic species of plants have been listed in the definition of the bioclimatic regimes. Herein, we attempt to provide comparisons of the life forms of vegetation in different parts of Ecuador with respect to elevation and rainfall (and its seasonal- ity). The flora of western Ecuador is best known, but albeit incompletely, on the lowlands. Dodson and Gentry ( 1991 ) estimated that as many as 6300 species of vascular plants (20% endemic) occur in this region; many of the endemics are known from small areas of only a few square kilometers (Gen- try, 1 986). The flora of three areas has been studied intensively — humid tropical forest at Rfo Palenque (Dodson and Gentry, 1978), subhumid tropical forest at Jauneche (Dodson et al., 1985), and tropi- cal dry forest in the Cordillera Chongon-Colonche in Guayas (Valverde, 1991). 18 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Table 4. Diversity of woody plants having diameters of >2.5 cm in 0. 1 -ha plots in forests in western Colombia (C) and Ecuador (E) (from Gentry, 1988). Site Lat. & long. Elevation Rainfall Species Regime Tutunendo, C. 05°46' N, 76°35' W 90 m 9000 mm 258 Humid tropical Bajo Calima, C. 03°55' N, 77°02' W 100 m 7470 mm 265 Humid tropical Centinela. E. 00°34'S. 79°18'W 550 m 3000 mm 140 Humid tropical Rio Palenque, E. 00°34' S, 79°20' W 200 m 2980 mm 121 Humid tropical Juaneche, E. 01°16'S, 79°42'W 60 m 1855 mm 96 Dry tropical Capeira, E. 02°00' S, 79°58' W 50 m 804 mm 60 Dry tropical There is a conspicuous change from deciduous to evergreen forests at the lower end of the precipi- tation gradient from coastal southwestern Ecuador inland and northeastward (Fig. 7). However, equally important biologically but less conspicuous to the casual observer is the dramatic increase in species diversity of woody plants along the north-south rainfall gradient. This phenomenon was demon- strated effectively by Gentry (1988), who com- pared diversity among equal-sized plots in western Ecuador and Colombia. Diversity decreases by more than 50% in humid tropical forests from the north with 9000 mm of rainfall annually to the southern limits of these forests where rainfall is only about 3000 mm (Table 4). Diversity decreases by another 50% in the dry tropical forest. Unfortunately, comparable data are not avail- able from the Andean slopes. However, Valencia and J0rgensen (1992) reported 32 species of trees in a 1-ha plot at an elevation of 3260-3300 m on Volcan Pasochoa. In addition to the deciduous-evergreen nature of the woody vegetation along the lower part of the rainfall gradient, which is at least in part influenced by the length of the dry season, there is a noticeable change in the presence of a canopy. A continuous canopy is important in providing shade and reduc- ing evaporation in the lower strata; thus, the soil and leaf litter retain high moisture contents. Al- though the canopy is discontinuous and rainfall generally declines with altitude in the humid sub- tropical and temperate regimes on the slopes of the Andes, these environments are characterized by high humidity. In addition to rainfall, moisture is achieved by fog, low dew points, and low evapora- tion rates. The vegetation in these regimes is char- acterized by many ferns and moisture dependent herbs in the lowermost strata. Trees commonly are heavily laden with epiphytes, including bromeli- ads, orchids, ferns, and mosses. At the highest elevations with low temperatures, species of trees characteristic of lower elevations are either absent or dwarfed and commonly festooned with epi- phytic mosses. One of the most striking vegetation formations in the Andes is the paramo. These supra-treeline regions generally lack woody plants and are pre- dominately grasslands with some areas having characteristic Espeletia or cushion plants. Thus, in western Ecuador, vegetation ranges from hot deserts with candelabra cacti through dry to wet tropical, subtropical, and temperate forests to the tundralike paramos. These highly diverse vegetation types provide highly different kinds of habitats for ani- mals. Geologic and Climatic History The dominant physiographic feature in western South America, the long Andean chain, was not always so prominent. Orogenic events associated with plate tectonics along the west coast of South America in the Late Cretaceous first uplifted the Andes to elevations probably not greatly exceeding 1000 m (Simpson, 1979; Zeil, 1979). According to Galvis (1982) and Galvis and Mojica ( 1 994), in the early Tertiary, an island arc existed to the west of the present Cordillera Occidental in Colombia and remnants persist today as the Serrania de Baudo in northern Departamento de Choco and possibly also the Serrania de Gallinazo in Departamento de Nariiio. Following the reasoning of Galvis and Mojica, we suggest that the Cordillera de la Costa in Ecuador represents remnants of the southern part of this island arc. ELEUTHERODACTYLUS IN WESTERN ECUADOR Volcanism and uplift of this island arc resulted from tectonic activities, particularly the subduc- tion of the Pacific plate, so that sections of the island arc were uplifted to elevations much greater than exist today — 1810 m at Alto del Buey in the Serrania de Baudo to 800 m at Cerro de Pata de Pajaro in Ecuador. The erosion of these elevated islands provided sediments to fill in the basins of what are now the Rio San Juan and Rio Atrato in Colombia and presumably the lowlands between the Cordillera de la Costa and the Cordillera Occi- dental of the Ecuadorian Andes. According to Sauer ( 1971 ), the uplifted terrain resulting from the Cretaceous orogeny subsequently was eroded into 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 is evidenced by many active volcanoes in Colombia and Ecuador. These stupen- dous orogenies and subsequent erosion resulted in the deposition of deep volcanic-ash soils on the slopes of the Andes and ash and alluvium on the lowlands. However, geological evidence points to some volcanic activity and associated uplift, erosion, and deposits in the Cordillera Occidental of Colombia in the Oligocene (Galvis and Mojica (1994). If sediments originating from erosion of the early Tertiary island arc covered the region now occu- pied by the Cordillera Occidental and the volcan- ism reported for Colombia during the Oligocene extended into Ecuador, elevations of the Ecuador- ian Cordillera Occidental may have been much higher in the mid-Tertiary than formerly presumed. In the Quaternary, three periods of glaciation occurred in the Andes north of the Huancabamba Depression (Sauer, 1971). The first of these prob- ably was insignificant because few peaks had been raised to elevations that were suitable for glacia- tion; in fact, the absence of evidence of glaciation on some high peaks strongly indicates that they were uplifted to their present heights after the last glaciation. However, glacial striae and moraines provide evidence that during the last glaciation snow line was depressed 1 500-2000 m. We can only speculate about the climate in what is now western Ecuador during the Cenozoic. but several models and supporting evidence provide an insight into Quaternary climates. The first of these is the climatic model proposed by Fairbridge ( 1 972) that predicts a southern shift in the equatorial current during glacial maxima; this would result in higher rainfall farther south along the Pacific coast (Fig. 8). The second model (Vuilleumier, 1971; Fig. 8. Climatic models of a maximum glacial (A) and interglacial (B) cycle proposed by Fairbridge ( 1 972) showing ocean currents, pressure systems, and wind directions. (Solid arrows are July; dashed arrows are January.) Continental outlines and extent of glacial ice ( shaded areas in A ) are based on S impson (1971). Adapted from Simpson (1979). 20 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Simpson, 1975) predicts altitudinal oscillations in climate and vegetation in the Andes; this model is strongly supported by palynological data from the Andes of Colombia, which show that the Recent vegetation of the high Andes did not come into existence there until the Late Pliocene (consistent with orogenic evidence) and that changes in veg- etation correspond to cool glacial and warm inter- glacial phases during the Quaternary (van der Hammen, 1974); during glacial maxima, tempera- tures in the high Andes were depressed 6-7°C, but in the lowlands temperatures were depressed only about 3°C (Haffer, 1979). According to Simpson ( 1979). major glaciations occurred about 250,000 and 130.000 years ago. and the last minor glacial advance took place only 14,000 years ago. The once-popular conception of the stability and antiq- uity of lowland tropical forests was dispelled by Haffer (1969; 1974), who proposed the refuge theory that forest and nonforest biomes changed continuously in distribution, fragmenting and coa- lescing during the varying humid and arid climatic conditions in the Quaternary. How do the events proposed in these models apply to the western Andean slopes and Pacific lowlands of Ecuador? Four aspects of Pleistocene glaciation and associated climatic oscillations con- tribute to biological distributions: 1 . Temperatures were lower during glacial phases and higher during interglacials. 2. Sea level was lower during glacial phases and higher during interglacials. 3. Rainfall decreased during the height of glacia- tion and increased during interglacials. 4. The water table in the lowlands was lower during glacial phases and higher during inter- glacials. With the exception of temperature fluctuations, each of these phenomena is associated with an increased amount of the earth's water being frozen during glacial periods; thus, rainfall diminished and sea level and water tables dropped. Therefore, during glacial maxima, the lowlands of western Ecuador were only slightly broader than they are now, probably had a lower water table, and most likely received less rainfall than at present. These phenomena are not inconsistent with proposed forest refuges in Chocoan Colombia, where rain- fall today is significantly higher than in the low- lands of Ecuador; refuges were proposed for birds (Haffer, 1974). Heliconius butterflies (Brown. 1982), and woody plants (Gentry, 1982; Prance. 1982). The models for depressed temperatures and decreased rainfall during glacial phases predict climatic depression to such a degree that lower bioclimatic regimes would either be eliminated or compressed. Climatic compression was addressed by Duellman ( 1 983a), who applied adiabatic lapse rates to climatic data from four elevations on the Amazonian slopes of the Andes and the upper Amazon Basin in Ecuador to predict climates and zones of vegetation on those slopes during glacial maxima. Because moist air cools more slowly than dry air, there is a range of adiabatic lapse rates from a loss of 6°C per 1000 m of elevation for saturated air to a loss of 10°C per 1000 m of elevation for dry air (Trewartha. 1943). This concept was applied to data on tempera- tures at 17 stations between 6 and 3560 m and within one degree of the Equator in western Ecua- dor When these data are plotted against the theo- retical humid adiabatic lapse rate, it is evident that at elevations below 1 500 m, the rate of temperature reduction with respect to elevation is less than 6°C, whereas the opposite is true at elevations of more than 1500 m (Fig. 9). Using van der Hammen's ( 1 974) and Haffer 's ( 1 979) predicted depression of temperatures by 6°C in the highlands and 3°C in the lowlands, temperature was lower for each of the sites proportional to a 6°C depression at 4000 m and a 3°C depression at sea level, and the humid adiabatic lapse rate was depressed accordingly. Again, at elevations below 1500 m. the rate of temperature reduction is less than that predicted by the humid adiabatic lapse rate, and at higher eleva- tions the opposite is true. This discrepancy can be explained by change in moisture content of the air masses as they rise along the front of the Andes. Although some moisture is lost at lower elevations, the air masses retain great quantities of moisture at higher elevations, as is evidenced by heavy fog banks above 1500 m. The oscillations in temperature and changes in adiabatic lapse rates during the Quaternary resulted in altitudinal shifts in vegetation zones in the Andes. ELEUTHERODACTYLUS IN WESTERN ECUADOR 21 4000 3500 3000 2500 c o ■■g 2000 > ID 1500 1000 500 5 10 15 20 25 Mean Annual Temperature (°C) 30 Fig. 9. Mean annual temperatures at sites at different elevations in western Ecuador ( • ) and hypothesized mean annual temperatures at those same sites during glacial maxima (o). The humid adiabatic lapse rate is shown as a solid line; the hypothesized humid adiabatic lapse rate during glacial maxima is shown as a dashed line. 22 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 These were calculated for the eastern slopes of the equatorial Andes by Duellman (1983a). Using a depression of 6°C and a corresponding lowering of vegetation zones in the high Andes and a depres- sion of 3°C at low elevations, it is possible to postulate the elevational ranges of vegetation zones during a glacial maximum. Likewise, the same can be postulated for a maximum interglacial, when, according to van der Hammen (1974), tempera- tures in the high Andes were elevated about 3°C; an increase of 1.5°C was used at low elevations (Fig. 10). Interglacial times are predicted to be much the same as at present, except all vegetation zones were shifted upward slightly at lower elevations and more so at higher elevations; the snow line was at approximately 5500 m and the upper limits of forest and therefore the frost line was at about 4000 m, in contrast to elevations of 4900 m and 3400 m today, respectively. This upward climatic shift re- sulted in paramos being more restricted and iso- lated than at present and much of the area covered by paramo today being dominated by subparamo. Elhn forest and high cloud forest would have breached many Andean passes, and in places, high cloud forest may have been continuous between the Pacific and Amazonian slopes of the Andes. Otherwise, vegetation zones were nearly as con- tinuous as they are today. In contrast, glacial times were characterized not only by depression of temperatures, more so at higher than lower elevations, but lower rainfall, which probably was most reduced at lower eleva- tions. These climatic modifications resulted in a lowering and compression of montane vegetation zones and drier conditions that eliminated or greatly restricted lowland tropical rainforest in western Ecuador. Cool-air drainage in river valleys deep- ened by rushing waters from melting glaciers would have resulted in cooler temperatures and lower evaporation rates and allowed the existence of cloud forest at moderately low elevations. The intervening ridges would have been more suscep- tible to insolation and dry winds off of the lowlands and probably supported tropical dry forest. Pockets of lowland tropical rainforest probably persisted in areas of high rainfall, such as in the region between Cayapas and Lita at the base of the Andes. 6000 Snow Snow Snow 5000 Paramo Paramo Subparamo 4000 Elfin Forest Subparamo Paramo High Cloud Forest g Elfin Forest .2 3000 > Subparamo High Cloud Forest w Elfin Forest High Cloud Forest 2000 Cloud Forest Cloud Forest 1000 Cloud Forest Transition Transition Rainforest Transition Rainforest A- Rainforest Maximum Maximum Glacial Present Interglacial Fig. 10. Present and postulated vegetation zones during maximal glacial and interglacial times with re- spect to elevation in equatorial western Ecuador Admittedly, this scenario assumes that during glacial periods the Andes were not significantly higher than at present and also is based on a restricted amount of evidence, some of which has been interpreted in support of a contrary position ELEUTHERODACTYLUS IN WESTERN ECUADOR 23 (e.g., Collinvaux, 1993). Most of this evidence is extrapolated from the paleoecological work of van der Hammen (1974) and van der Hammen and Cleef ( 1986), and the hypotheses of Haffer (1969) and Vuilleumier (1971). However, it provides a basis for refinement as new paleoclimatic and pa- leoecological data become available and a model for interpretation of ecological and geographic effects on patterns of speciation and distribution of organisms inhabiting western Ecuador. Human Environmental Modifications Prior to the early 1960s, the human population on the Pacific lowlands and on the Pacific slopes of the Andes in Ecuador was relatively low. Indig- enous peoples, such as the Cayapas and Colorados, were primarily hunter-gatherers, and the few colo- nists who inhabited the region were mostly subsis- tence farmers. For decades, the only town of any size on the coastal lowlands was Guayaquil, esti- mated to have about 28,000 inhabitants in 1879 (Whymper, 1 892). In the vicinity of Guayaquil and less so in smaller towns and villages, adjacent forests were cut for firewood and construction materials, and palm trees were cut for their fronds for thatched roofs. Most rural inhabitants were located along the rivers that provided transit to towns. As Guayaquil became a port more fre- quently visited by freighters, timber cruisers began removing economically important trees, such as caoba (Perseo theo-bromifolia), balsa (Ochroma lagopus) and Colorado (Pouteria sp. ) that were near rivers and easily accessible; during the period of World War II, rubber collectors worked the native rubber trees (Castilla elastica) (Dodson and Gen- try, 1978). Narrow "avenues" of colonization emerged along the two railroads that crossed the Andes and penetrated the lowlands to the coast; the northern railroad extended from Ibana to San Lorenzo, and the southern from Alausi to Guayaquil. Although these railroads provided important trade routes, they had only limited impact on the natural re- sources of the region. The first commercial crops of any significance were banana and cacao planta- tions at lower elevations and coffee plantations in the shade of native trees at higher elevations; most of these plantations were within a short traveling distance to the railroad. Prior to the mid- 1 900s, few roads penetrated the region; the most important of these was the "old road" from Quito via Chiriboga to Santo Domingo de los Colorados. Aside from subsistence farming and some dairy cattle, coffee was the major agricultural product. Two coincident events in the early 1960s dra- matically affected the Pacific lowlands. The first was the initiation of a large road-building pro- gram— a new paved road from Quito to Santo Domingo de los Colorados. paved roads from there to Esmeraldas and to Guayaquil, and later, paved roads from Cuenca to Guayaquil. These roads provided the routes for commerce and coloniza- tion, which were bolstered by opportunities for the second event, commercial agriculture involving cacao, sugar cane, oil palms, and especially ba- nanas (the "oro verde" or green gold), and forest extraction for commercial purposes. Within a de- cade hundreds of thousands of hectares of forest were cut, burned, and planted. Chemical pesticides (DDT and Bordeaux) were used with abandon; the latter sprayed from planes gave banana plants a bluish hue. During our various trips to the lowlands between Esmeraldas and Babahoyo between 1967 and 1984, we witnessed this wanton destruction and pollution of the lowland tropical forest. The natural resources have been pressured fur- ther by the great increase in population by burgeon- ing colonization and birth rates. Guayaquil now has more than 2.5 million inhabitants, and Provincia Manabi more than 1 million (Mejia et al.. 1995). Small towns or villages of two decades ago now are major metropolises, some with more than 100,000 people. According to Dodson and Gentry (1991). the forests of western Ecuador are among the most severely threatened of the world's ecosystems; more than 90% of the forests on the lowlands in the foothills have been converted to agriculture. This rapid destruction in the lowlands and on the higher slopes of the Andes was documented further by Mejia et al. (1995). Although there is notably less destruction of natural forests on the steep slopes, an 24 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 increasing number of roads (and the accompanying colonists) results in ribbons ot destruction, even in protected areas, such as the Bosque Protector de Molleturo. Two large natural reserves on the west- em versant in Ecuador, the Reserva Ecologica Cotacachi-Cayapas and the Reserva Etnica y For- estal Awa totaling about 280,000 ha, face destruc- tion as squatters establish themselves within the boundaries of the reserves and economically valu- able trees are removed. At higher elevations, mo- nocultures of Eucalyptus and Pinus form a barren, inhospitable habitat for native animals. Over a short period of time, the changes wrought by humans equal or exceed those by natural causes during the Pleistocene. Certainly, within the past few decades, many species have become extinct, as noted for plants by Dodson and Gentry (1991 ). We regret that this publication is a requiem for extirpated and surely threatened species of Eleutherodactylus. My antecedents were Eleutherodactylus Long ago when there were just a few of us But as the Andes arose Some of us nearly froze So we dispersed in response to this stimulus We differentiated without much of a fuss Preferring climates that others would cuss Where it rained, it would pour So we'd speciate more For us, others' nonplus was a plus But all did not remain thus For the diversity of Eleutherodactylus The forests were cleared Our habitats disappeared So now there are fewer species left of us SYSTEMATICS Description of Characters In many museums, frogs of the genus Eleutherodactylus often are among the last speci- mens to be identified in incoming collections. This is owing not only to the plethora of species, but also the confusion about many taxonomic characters. In an attempt to rectify this situation and hopefully to standardize terminology, we clearly define and illustrate the morphological characters that we have found to be useful in eleutherodactyline systemat- ics; examples are provided from species from west- ern Ecuador. However, "definition" of some char- acters (e.g., skin texture) is difficult, because al- though the character states are not discrete, their characterization in descriptions may suggest that they are discrete. We realize that this may confuse nonspecialists; for this we apologize. The charac- ters are presented in approximately the same order in which they appear in the numerical diagnoses. Texture of skin. — In several species (e.g., E. achatinus, anomalus, and w-nigrum) in western Ecuador, the skin on the ventral surfaces is smooth, except for being granular on the proximal surfaces of the thighs. In most species in the region, the skin on the belly is granular ("areolate"); the throat and chest may be less strongly granular or smooth. In a few of these species (e.g., E. crenunguis, labiosus, and ocellatus), the granules are poorly defined; the belly in individual specimens might appear to be smooth or weakly granular in separate observation events. The texture of the skin on the venter (if granular) is affected by preservation fluids and techniques; thus, a dead specimen that is later preserved may appear to have smooth skin on the belly, even under high magnification. The skin on the dorsum, flanks, and upper sur- faces of the limbs tends to have the same texture in any given species, but some departure from that generality occurs. In many species (e.g., E. hamiotae, laticlavius, quinquagesimus, and vertebralis), the texture of the skin on the posterior part of the dorsum is coarser than that on the anterior part. Among those species with tubercles on the dorsal surfaces, the tubercles usually are smaller on the limbs than on the dorsum of the body, except in those taxa (e.g., E. appendiculatus) in which the tubercles are closely associated with the color pattern on the limbs. Transitional zones between the texture of the dorsum and that of the venter occur on the flanks, but the texture of the flanks is not always intermediate between the dor- ELEUTHERODACIYLUS IN WESTERN ECUADOR 25 sal and ventral textures; in some species in the E. fitzingeh and E. conspicillatiis groups, the dorsum is finely granular and the venter is smooth, but the flanks bear larger granules. The extreme conditions of texture on the dor- sum are: (1) smooth (e.g., E. apiciilatus, caprifer. gularis, and loustes), and (2) coarsely tuberculate, and usually bedecked with folds (e.g., E. anomalus, cerastes, andnecerus). Most species (e.g., £. actites, crenunguis, leoni, nyctophylax, ornatissimus, and siopelus) have dorsal skin that is described as uniformly finely granular ( shagreen). If some granu- lations are notably higher than others, the skin is described as tuberculate if the granules are conical or subconical (e.g., E. hectus and E. latidiscus) or warty if the granules are flattened (e.g., E. babax, dueUmani, and ruidus). The descriptors of dorsal skin texture are intended to define the average condition into a few categories (e.g., smooth, shagreen, weakly tuberculate, or coarsely tubercu- late) qualified with reference to folds and/or regu- lar (i.e., named) tubercles. The dorsal skin texture is strongly affected by numerous factors — inad- equate preservatives, death by desiccation and/or heat stress, early decomposition, and desiccation of a preserved specimen followed by some effort at rehydration. Commonly, juveniles are more tuber- culate than conspecific adults. The specific terms describing elevations on the skin are defined as follow. Tubercles are relatively large elevations that may be elongate, conical, or subconical (in order of decreasing elevation). Warts also are relatively large, but flattened or rounded in contrast to tubercles; the base of a wart is broader than its elevation. The terms pustules and spicules are applied to smaller structures; pw^m/^^ are small warts, and spicules are minute tubercles. Pustules and spicules are difficult to distinguish from some descriptors of skin texture (e.g., shagreen). There- fore, a comparsion is made between the general texture of the skin and the relative sizes and shapes of protuberances on a given individual. Tubercles. — Apart from being used to describe texture, certain tubercles regularly appear in differ- ent individuals and species of eleutherodactyline frogs. These tubercles have specific names because they occur in particular places on the frog. Savage (1987) used the term "accessory" for what we call supernumerary tubercles. He used the term "super- numerary" to describe tubercles other than subarticular tubercles on the undersides of digits. Body: Aside from scapular tubercles associated with postocular folds, few body tubercles have specific names, at least among species in western Ecuador. (See Folds.) However, the pattern of tubercles on the dorsum of several species (e.g., E. anomalus and E. necerus) is nonrandom, but the details of tubercular arrangements have not been investigated as a source of taxonomic characters. (See Skin Texture.) Eyelid: If the posterolateral quadrant of the upper eyelid bears an enlarged tubercle, the struc- ture is termed the eyelid tubercle. In some frogs, this tubercle is elongate (i.e., much longer than wide; e.g., E. cerastes) or conical (i.e., longer than wide; e.g., E. appendiculatus). The upper eyelid usually bears a single enlarged tubercle (e.g., E. labiosus), but some species (e.g., E. leoni and E. verecundus) have three or four enlarged tubercles. If the upper eyelid is simply warty or tuberculate with none of the tubercles being larger than others, the frog is described as lacking an eyelid tubercle. Eoot: The plantar surface of the foot bears several tubercles having particular names. Toes I and II each have a single subarticular tubercle; Toes III and V each have two subarticular tu- bercles, and Toe IV has three subarticular tubercles. The inner (preaxial) metatarsal tubercle is longer than wide; it is usually oval, but it is elongate in some species (e.g., £". anomalus and E. longirostris) and laterally compressed in a few species (e.g., E. necerus).The outer ipostaxial) metatarsal tubercle usually is much smaller than the inner metatarsal tubercle and round or slightly longer than wide; in a few species (e.g., E. eremitus, hectus. and qidnquagesimus) it is conical or subconical. Super- numerary plantar tubercles are absent in some species (e.g., E. anatipes, cerastes, and hamiotae); they are present as low, diffuse structures in a few species (e.g., E. chalceus and E. loustes), but more commonly (e.g., E. achatinus, labiosus, ocellatus, and parx'illus) only three or four tubercles are present at the bases of Toes I-IV or II-IV. In a few species (e.g., E. celator, eremitus, and hectus), supernumerary tubercles are more numerous, thereby rendering the plantar surface areolate. 26 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Forearm: The postaxial .surface of the forearm bears enlarged tubercles in many species. The most constant of these is the anfchrachial titherclc that Hcs just proximal to the wrist, but several species (e.g.. E. appendiculatiis, cerastes, and verecundus) have a row of equally large tubercles between the wrist and elbow. These may be low or conical and are termed ulnar tubercles, unless they are com- pressed and Juxtaposed, in which case they are termed an ulnar fold. (See Folds: Ulnar.) Hand: The tubercles on the palmar surface of the hand closely parallel those on the foot. Suharticular tubercles occur beneath the joints between phalanges; thus, there is one suharticular tubercle on the thumb and second finger and two on the third and fourth fingers. Suharticular tubercles vary in elevation (low and rounded vs. conical and projecting) and shape (base rounded vs. base elon- gate). In a few eleutherodactylines (e.g., E. chalceus), the suharticular tubercles are broad and some (usually distal ones on outer digits) are bifid. Other prominent tubercles are on the proximal edge of the palm. The thenar tubercle at the base of the thumb is oval, whereas the palmar tubercle on the outer part of the palm usually is bifid (partially divided distally ). In older literature, the palmar and thenar tubercles were called the metacarpal and carpal tubercles. On the fleshy part of the palm, supernumerary (palmar) tubercles (also called ac- cessory tubercles) may be present. These usually are low and difficult to see clearly, but in some species (e.g., Barycholos pulcher), the supernu- merary tubercles are prominent and conical. Head: Apart from the eyelid, postrictal, and snout tubercles discussed separately, there are a few tubercles that occur sporadically on the head. Some species (e.g., E. labiosus) possess an interocular tubercle between the eyes; others (e.g., E. latidiscus and E. muricatus) have tubercles on the upper surface (usually on the midline) of the snout. A few species (e.g., E. ocellatus) have small conical tubercles along the margin of the lower jaw. Hind limb: Many species (e.g., E. anomalus. cerastes, latidiscus, and necerus) have obvious, enlarged tubercles associated with features of the color pattern on the shank, but these tubercles are not named. The upper edge of the heel commonly bears one or more tubercles variously called tu- bercles or calcars. We term those that are enlarged, but not longer than wide, heel tubercles. If the tubercle is elongate and pointed, it is identified as a conical heel tubercle. A conical heel tubercle may be only twice as long as wide (e.g., E. calcarulatus, crucifer muricatus, i\ndverecundus) or much more than twice as long as wide, as in the extralimital E. calcaratus or E. inusitatus. Some authors (e.g.. Savage, 1 987) employ the term calcar for a variety of ornamentations on the heel, but we restrict the term calcar to describe an elongate tubercle having a broad, flat base (e.g., E. quinquagesimus and E. slope I us). In many eleutherodactyline frogs (e.g., E. appendiculatus, crenunguis, crucifer, quinqua- gesimus, and siopelus), the postaxial surface of the tarsus bears a series of distinguishable tubercles called outer tarsal tubercles; these display the same range of variation as ulnar tubercles. (See Tubercles: Forearm.) The inner edge of the tarsus may bear one or more tubercles, but these are homologous with the tarsal fold. (See Folds: Tar- sus.) Postrictal: Posteroventral to the tympanum and just anterior to the posterior edge of the jaw muscu- lature is a pair of postrictal tubercles. These may be low and rounded, as in E. luteolateralis and E. nyctophyla.x, or conical, as in E. pyrrhomerus. In a few species (e.g., E. babax), the tubercles are compressed and fused so as to form a short ridge that extends ventrolaterally from the tympanum. Snout: Several species (e.g., E. eremitus, pho.xocephalus. and subsigillatus) have a conical tubercle (papilla) at the tip of the snout (visible in either dorsal or lateral view) that breaks the smooth curve of the margin of the snout. This tubercle has a broad base and seldom is longer than broad, but E. appendiculatus has an exaggerated tubercle, more appropriately termed a proboscis. Dermal folds. — Among the species in western Ecuador, eight kinds of dermal folds can be found. Discoidal: This fold defines the ventral disc on the belly and is most obvious laterally and posteri- orly, but the fold is absent in a few species (e.g., E. helonotus). In most Ecuadorian, as well as other South American Eleutherodactylus, the margin of the discoidal fold is well anteriad to the groin; this is in contrast to some Mexican taxa in which the ELEUTHERODACTYLUS IN WESTERN ECUADOR 27 margin of the fold nearly reaches the bases of the hind limbs. Dorsolateral: These paired folds extend poste- riorly from the posterolateral corner of the orbit and lie lateral to the sacrum and blades of the ilia. Of the 6 1 species oi Eleutherodactyliis known from west- ern Ecuador, 43 have no traces of dorsolateral folds. In those species having folds, the folds usu- ally are most prominent anteriorly and become pustular (fragmented into a series of pustules) posteriorly before terminating above the groin. Distinct dorsolateral folds that are not fragmented into pustules posteriorly are present in only a few taxa (E. appendiculatus, hectus, and vertebralis). In 10 other species {E. achatinus, actites, gentryi, illotus, laticlavius, lymani, sobetes, thymalopsoides, and truebae), the folds are thinner or fragmented into pustules throughout their entire lengths. Four species (£. babax, duellmain, quinquagesimus, and siopelus) have short dorsolateral folds that are not evident posterior to the level of the sacrum. Eleutherodactylus anatipes has a longitudinal se- ries of elevations in the topographic position of dorsolateral folds. Although there is no evidence that these elevations are parts of a whole, we refer to them collectively as a fragmented dorsolateral fold. In E. anatipes, additional shorter, longitudinal elevations are present ventrolateral to the dorsolat- eral folds, and, with the same reservations, we also refer to these as dorsolateral folds. In three species (E. sulculus, surdus, and verecundus), the dorsolat- eral folds are evident only along the midlength of the body; the folds are difficult to see in E. surdus, in some specimens of which the folds are absent. Interocular: This transverse fold lies between the eyes on the top of the head. In western Ecuador, it is present in only one species {E. quinquagesimus). Paravertebral: These folds lie above the trans- verse processes of the vertebrae and are median to the dorsolateral folds (if present). Paravertebral folds commonly are associated with a specific pattern-polymorph, in which a broad, pale, mid- dorsal (vertebral) raphe is present (e.g., E. laticlavius, vertebralis, and walkeri): the folds are at the lateral margins of the raphe. In other species (e.g., E. cerastes and E. necerus), the paravertebral folds contribute to a pattern of dorsal ridges, espe- cially those in which there is an H-shaped fold in the scapular region. The H-shaped fold usually is associated with a pair of short paravertebral ridges above the coccyx. Postocular: These folds extend posteromedially from the eyes onto the occiput. The anterior ex- tremity of a postocular fold commonly is an en- larged tubercle on the upper eyelid, and the poste- rior extremity commonly is an enlarged scapular tubercle, as seen in E. crenunguis. In several spe- cies (e.g., E. labiosus, latidiscus, and tenebrionis), folds extend anteromedially from the scapular tu- bercles; these, together with the postocular folds, define a W-shaped occipital fold. Flores (1993) included the postocular fold as a kind of dorsolat- eral fold. Supratympanic: This fold begins at the poste- rior corner of the orbit, extends posteriorly along the upper edge of the temporal region, and usually angles posteroventrally toward the insertion of the arm. However, in E. colonial, the fold continues straight from the temporal region to a point above the insertion of the arm. In a few species (e.g., E. longirostris and E. loustes), the supratympanic fold is distinct, but in most species, its limits are difficult to define; the ambiguities obtain from color pattern (edges of supratympanic stripe) and the superposi- tion of tubercles on the fold. Tarsal: In many species in western Ecuador (e.g., E. actites, anomalus, longirostris, loustes, lymani, and ruidus), a longitudinal fold is present on the inner edge of the tarsus. This fold usually is a more or less clearly defined ridge extending a variable distance proximal to the inner metatarsal tubercle, but the fold is flaplike in E. loustes. Short inner tarsal folds sometimes have been called inner tarsal tubercles, but we term them folds when they are within one length of the inner metatarsal tu- bercle and if they appear to be connected to the inner metatarsal tubercle, as in E. hectus, laticlavius, latidiscus, leoni, muricatus, ocellatus, pyrrhomerus, siopelus, sulculus, and verecundus. In a few spe- cies (£. hamiotae, sobetes, truebae, vertebralis, and w-nigrum), we are unsure as to whether or not a short foldlike tubercle lies immediately proximal to the inner metatarsal tubercle; in some speci- mens, a structure appears to be present. In 13 species (E. achatinus. calcarulatus, cajamarcensis, crucifer, duellmani, eugeniae, gentryi, illotus, quin- 28 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 quagesimus. wsado'Lsuhsi^illatusjhynialopsoides, and na/Ai'/v). a distinct, small tubercle is present on the distal one third of the inner surface of the tarsus and is clearly separated from the inner metatarsal tubercle. We think that this structure is a homo- logue of the inner tarsal fold because of the com- mon position of the two structures and because gradation between a fold and a tubercle has been observed in the Mexican E. rhodopis. Ulnar: In none of the Eleutherodactylus in west- em Ecuador do the ulnar tubercles fuse into an ulnar fold, such as is seen in E. latens and E. ruizi in Colombia. Tympanum. — The terms "tympanum" and "ex- ternal ear" have been used to identify a combina- tion of characters rather than a single character. The "tympanum" is a combination of ( 1 ) a differenti- ated tympanic membrane and (2) a tympanic annu- lus. Wever (1985:43) defined the tympanic mem- brane as "... an area of modified skin, much thinner than ordinary skin and largely lacking subcutane- ous layers ... and its edges are attached to the flaring rim of a funnel-shaped cartilage, the tym- panic annulus... ." We concur that the skin com- prising the tympanic membrane is thinner than the skin elsewhere on the side of the head; also, the tympanic membrane usually has a smoother tex- ture, commonly has a distinctive coloration, and in many cases is slightly to obviously translucent because of the underlying tympanic cavity. When- ever a differentiated membrane is apparent, the annulus also is present, but the absence of a differ- entiated membrane does not necessarily signify the absence of the annulus. Long ago. Noble ( 193 1 :334) recognized that "... a few [frogs] have the integu- ment unmodified and separable from the underly- ing drum, ... ."" This equals the absence of a tympanic membrane but the presence of a tym- panic annulus. Such a combination frequently has been designated as "tympanum concealed," an unfortunate descriptor that confuses more than it informs. Externally, conditions vary from a prominent tympanic membrane lying within the flaring rim of the tympanic annulus to the absence of a tympanic membrane and no external evidence of a tympanic annulus (Fig. 11). Between these extremes is a gradation of conditions that may be difficult to discern owing to variations in thickness of the skin and degree to which desiccation might have drawn the skin tight against the underlying structures. Reduction of auditory structures in anurans seems to follow an obligate sequence involving loss of structures in a distal-medial sequence (Trueb, 1 993). Thus, the tympanic membrane may be absent, but the tympanic annulus and the remainder of the plectral apparatus present; subsequent loss of the tympanic annulus leaves the stapes (= columella), which also may diminish in size or disappear. Some of the problems inherent to the descrip- tion of the tympanum are artifacts of preservation; others result from investigators being unaware of the internal anatomy of the otic region. Specimens preserved in alcohol that is too strong or specimens desiccated and subsequently rehydrated might seem to have an "external tympanum," whereas living or well-preserved specimens appear to lack an "exter- nal tympanum." Some authors have reported the tympanum to be "hidden" (e.g., Flores, 1988a, for E. alberchi, a synonym off. surdus), because they could not discern an external tympanum but, curi- ously, assumed that one was present beneath the skin. Ever since his early investigation of eleutherodactyline frogs, JDL consistently has dis- sected the otic region in order to evaluate the condition of the tympanic annulus if it is not visible externally. With the reconciliation that the "tympa- num" is a combination of characters, we are forced to abandon descriptors such as "tympanum hid- den" and "tympanum concealed" and urge our colleagues to do likewise. A prominent tympanic membrane and tympanic annulus are present in most Eleutherodactylus in western Ecuador (e.g., E. calcarulatus, hecfus, longirostris, tenebrionis, and w-nigrum). Usually the skin circumscribed by the annulus is thin and in some species, it may be translucent; typically, the tympanic annulus is elevated with respect to the temporal musculature and its entire circumference is visible, although commonly, the annulus is ob- scured dorsally by a supratympanic fold. In a few species (e.g., E. gularis and E. scolodiscus), a tympanic annulus is evident through thin skin that is not connected to the annulus. In other species (e.g., E. loustes and E. tniehae), only the lower part of the tympanic annulus is evident under undiffer- ELEUTHERODACIYLUS IN WESTERN ECUADOR 29 .-.^^»STn«?^^?jP?ii?-^>,?f^.^'?^"afc:^^^^?^ Fig. 1 1. Sides of heads showing comparative distinctness of tympanic annuH and membranes in Eleuthero- dactylus. A. E. longiwstris, LACM 47170, with distinct annuH and membrane. B. E. gularis, QCAZ 4316, with annulus distinctly visible beneath skin. C. E. truebae, QCAZ 2475, with annulus partially visible beneath skin. D. E. hamiotae, USNM 239843, with no ear structures evident externally. Scale bars = 2 mm. entiated skin; otherwise, there is no external evi- dence of a tympanic annulus. Among the Eleutherodactylus in western Ecuador, a tympanic annulus is absent in six species — E. duellmani, hamiotae, ruidus, siopelus, sobetes, and surdus — and reduced or absent in one other {E. gentryi). Thus, we recognize four character combina- tions of the ear among eleutherodactyline frogs in western Ecuador, as follow: (1) tympanic mem- brane and tympanic annulus prominent; (2) tym- panic membrane absent, but most of the circumfer- ence of the tympanic annulus evident through the skin; (3) tympanic membrane absent but tympanic annulus evident only ventrally; and (4) tympanic membrane and tympanic annulus reduced in size or absent. Among eleutherodactylines in western Ecua- dor, sexual dimorphism in the diameter of the tympanic annulus is appreciable only in E. longiwstris, in which the tympanic membrane and tympanic annulus are prominent and the horizontal length (i.e., "diameter") of the annulus in females is about half the length of the eye. However, in males, the annulus occupies most of the temporal region, and the ventral rim of the annulus nearly extends to the lip. The length of the annulus in males is about three-fourths the length of the eye. Such dramatic sexual dimorphism is common, but not universal, among frogs of the subgenus Craugastor. In the other three species in the subge- nus in western Ecuador (£. anatipes, anomalus, and necerus), there is no sexual dimorphism in the diameter of the tympanic annulus, or the annuli of males are only slightly larger than those of females. Snout shape. — In most species of Eleuthero- dactylus in western Ecuador, the margin of the snout is rounded or subacuminate in dorsal view (Fig. 12). There seems to be a clear association between snout length measured from the anterior corner of the orbit to the nostril (E-N) and snout 30 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 12. Snouts oi Eleutherodoctyhis showing different shapes. Dorsal views: A. E. appendiculatus, USNM 2047 13, acuminate. B. E. longirostris. LACM 47 1 70, subacuminate. C. E. tenehrionis, MCZ 9208 1 , rounded. D. E. loustes, KU 1 79243, rounded. Profiles: E. E. anomalus, LACM 4688 1 , rounded. F. E. degener, QC AZ 1 297, truncate. G. E. colomai, QCAZ 1289, protruding. Scale bars = 2 mm. shape. Thus, those species (e.g., E. achatinus and E. ornatissimus) having longer snouts (E-N/eye length > 0.8) have subacuminate snouts, whereas those species (e.g., E. anomalus and E. duellmani) having short snouts (E-N < 0.8) have rounded snouts. The most acuminate snout among species of Eleutherodacty'lus in western Ecuador is that of the long-snouted E. appendiculatus. In lateral view (profile), the snout usually is rounded, but the profile is nearly truncate in a few species (e.g., E. subsigillatus) or protrudes well anteriad of the margin of the jaw in E. colomai (Fig. 12). The profile of snout shape reflects the orienta- tion of the alary processes of the premaxillae and the degree to which the nasal cartilages extend anteriorly. The outline curvature of the jaws in dorsal view is disrupted by the protruding nostrils if the frog has a truncate snout in profile. The only other disrup- tion is caused by the presence of a tubercle or vertical keel on the tip of the snout. In dorsal view, the canthus rostralis usually is concave in Eleutherodactylus, but in several spe- cies (e.g., E. achatinus and E. Hiatus), the underly- ing nasal bone bulges laterally, thereby causing the canthus to be an uneven curve. In these species, the curvature is concave just anterior to the orbit and just posterior to the nostrils but convex throughout most of its length. We term such canthi as sinuous in contrast to the concave or straight canthi seen in most Eleutherodactylus. In transverse cross sec- tion, the canthus rostralis may be angular (e.g., E. calcarulatus, colomai, and nyctophylax) or obso- lete (e.g., E. hamiotae, iatidiscus, and loustes). Variation exists in the breadth of the top of the snout, defined as the dorsal plane in front of the ELEUTHERODACTYLUS IN WESTERN ECUADOR 31 orbits and bordered laterally by the canthi rostrali. One extreme is exhibited by E. colomai, in which the top of the snout is broad, the canthi rostrali are angular, and the loreal regions are nearly vertical. Adult females of E. latidiscus exhibit the other extreme, in which the top of the snout is narrow, the canthi rostrali are curved concavely in dorsal view, and the loreal regions slope gradually toward the flared lips. Cranial crests. — In several species of Eleutherodactyhis in western Ecuador (e.g., E. cerastes, necerus, ruidus, and vertebralis), bony crests develop on the frontoparietal bones. These crests lie just medial to the origin of the m. pterygoideus but may extend both anterior to and posterior to that origin. As is true for many features of Eleutherodactyhis, crests are better developed in larger than in smaller individuals; therefore, they are most prominent in adult females, less so in adult males, and the least so in juveniles. When present, crests usually are lowest anteriorly and highest posteriorly, but they may be uniform in height. Crests may be thick or thin, be lateral or parasaggital on the frontoparietals, and be smooth or bear bony projections. Because there is ontogenetic and sexual variation in this character, we have characterized species based on the condition in adult females, even when that misrepresents what might be ob- served in adult males or juveniles. This places an added burden on investigators to ascertain the sex and maturity of specimens when reading this ac- count or other papers that we have authored indi- vidually and jointly. Vomerine odontophores. — There is consider- able variation in vomerine odontophores (= dentigerous processes of the vomers) in Eleuthero- dactyhis. Species of the subgenus Syrrhophus lack odontophores, as do several species in other sub- genera. All species in western Ecuador have odontophores, although they may be concealed beneath the tissue of the palate and be difficult to see in some species (e.g., E. chalceus and E. scolodiscus). Likewise, all species have odonto- phores that are posteromedial to the choanae. Frogs of the E. discoidalis group (Lynch, 1989) are dis- tinctive in having odontophores that are located more anterior than are these structures in most Eleutherodactvlus. Odontophores increase in size as the frog grows. Therefore, odontophores in females normally are larger and more easily observed than those in males; larger odontophores usually are more nar- rowly separated from one another in the midline than are smaller odontophores. These structures commonly are difficult to detect in juveniles. Historically, much of the attention to odonto- phores concerned the "widths" of the structures; this has been "measured" in terms of how far laterally the odontophores extend relative to the middle or lateral margins of the choanae. Many species of the subgenus Euhyas have extremely broad odontophores that extend laterally to the lateral margins of the choanae. In Barycholos pulcher, the odontophores are comparable to those in Euhyas. However, in speciesof Eleutherodactylus in western Ecuador, the odontophores are much narrower and do not extend laterally even to the middle of the choanae. A few species in western Ecuador (e.g., E. helonotus and E. necerus) have odontophores that might be described as arched or broad, but these are relative terms and should not be confused with similar wording used by other au- thors for frogs such as members of the subgenus Euhyas. In addition to width, the shapes of odontophores vary, as observed in ventral view (Fig. 13). For example, in adults off. achatinus, the odontophore is approximately triangular in outline with the teeth in a transverse row along the base of the triangle. In adult E. calcarulatus, the odontophores are de- scribed as oblique or slanted, because there is no development of the tooth row posterolaterally, and the teeth are either in a clump or in an oblique row. In some species with massive odontophores (e.g., E. hamiotae and E. pteridophUus), the outline of the structure is oval, and the tooth row is not so well defined as in taxa having triangular odontophores. Secondary sex characters. — Males of many species of Eleutherodactylus have vocal apparati (sac and slits) and/or nuptial pads. Eleven species {E. celator, cerastes, floridus, gentry!, hamiotae, illotus, laticlavius, pteridophilus, quinquagesimus, siopelus, and surdus) lack vocal slits and nuptial pads. Their absence makes the determination of adulthood difficult in these species, in which matu- rity in males must be determined by examination of 32 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 the testes in conjunction with field observations on amplectant pairs. Vocal slits: These slits are known to occur in 34 species oiEleuthewdactylus in western Ecuador— E. achatinus, actites, anatipes, appemUcnlatus, babax, calcarulatus, caprifer, cluilceiis. colomai, crenunguis, crucifeK eremitus, degener, eitgeniae, gularis, hectus, labiosus, leoni, longimstris, loustes, luteolateralis, muricatus, necerus, nyctophykix, omatissimus, pan'illus, phoxoceplmliis. pxrrho- merus, scolodisciis, subsigillatus, tenebrioms, verecundus, walkerL andw-nigrum. Slits are thought to be absent in 20 species— E. anomalus, diiellmani, latidiscus, hmani, rosadoL ruidus, thymalopsoides, truebae, and vertebralis, in addition to the 11 species listed above. Adult males are not known for E. dissimulatus, belonotus, ocellatus, and sobetes. Nuptial pads: White, nonspinous nuptial pads are present on the dorsomedial and medial surfaces of the base of the thumb in breeding males of 23 species of Eleuthemdactylus in western Ecuador— E. achatinus, actites, anatipes, anomalus, crucifer, degener, duellmcmi, eremitus, eugeniae, latidiscus, longirostris, lymani, necerus, nyctophylax, omatissimus, plioxocephalus, rosadoi, ruidus, subsigillatus, thymalopsoides, truebae, vertebralis, and w-nigrum. Obviously, for four species known only from adult females (E. dissimulatus, helonotus, ocellatus, and sobetes), the condition of nuptial pads is unknown. Fingers.— The groups of characters associated with the hands are the relative lengths of the fingers, especially (and usually) Fingers I and II, the devel- opment of lateral fringes or keels on the fingers, and the anatomical details (discs and pads) on the tips of the fingers. Relative lengths of digits: In most Eleuthero- dactylus in western Ecuador, the thumb (Finger I) is noticeably shorter than Finger II (when each is adpressed equally). Finger I is only slightly shorter than Finger II in E. labiosus and E. ocellatus. Finger I is longer than Finger II in species of the subgenus Craugastor (E. anatipes, anomalus, longirostris, and necerus), most of the species in the E. conspicillatus group {E. achatinus, actites, illotus, hmani, and w-nigrum), the E. sulcatus group (£. cerastes and E. helonotus), and in E. babax, crenunguis, and loustes, as well as m Fig. 13. Ventral views of palates showing shapes and separation of vomerine odontophores in eleuthero- dactyline frogs. A. Barycholos pulcher. KU 218152, arched and barely separated. B. E. crenunguis. QCAZ 1 3 10, triangular and narrowly separated. C.£. leoni. KU 1 30870, oblique and widely separated. Scale bars = 2 mm. ELEUTHERODACTYLUS IN WESTERN ECUADOR 33 Barycholos piilcher (Figs. 14, 15). The fingers of several of these species (£. babax, crermnguis, labiosus, and ocellatus) seem to be exceptionally long, because they are higher than wide and lack any trace of lateral folds. The narrowness and lengths of the fingers are accentuated in those species (e.g., E. crenunguis) that bear prominent terminal discs. Other species have short ("stubby") digits either because the fingers are short (e.g., E. helonotus) or because they are wider than high (e.g., E. gularis). Lateral fringes and webbing: In many species, fleshy keels (e.g., E. gentryi and E. lymani) or fringes (e.g., E. duellmani, siopelus, and thymalopsoides) are present on the lateral margins of the fingers; there is a continuum from keels to fringes, and in some cases the distinction is arbi- trary. Generally, the digits are narrower when bear- ing a lateral keel than when bearing a fringe. In a few species (e.g., E. loustes), the fringes partially obscure the subarticular tubercles in preserved specimens. Lateral folds and especially fringes are sensitive to desiccation. In a subset of species having lateral fringes on the fingers (e.g., E. siopelus), there is a comparable fringe on the outer edge of Finger IV; in species such as E. eugeniae and E. pteridophilus, the fringe continues proxi- mally onto the edge of the palm. No such fold occurs on the median edge of Finger I. Webbing between the fingers is absent in all species. Discs and pads: When the tips of the digits are expanded, the digit is described as having a disc. Within individuals, discs are larger and wider on the outer fingers than on the inner ones. In most species bearing discs, the disc on the thumb is only slightly wider than the digit proximal to the disc, and the disc on Finger II is intermediate in size between that on the thumb and those on Fingers III and IV. In most species of Eleiitherodactylus, the skin on the underside of the digital tip is modified to form a pad defined by a circumferential groove. Usually the pad is wider than long, and its distal border is a smooth curve. The disc cover or ungual flap is visible distal to the pad and usually is curved (= "round"), but in some species (e.g., E. crenunguis), it is emarginate (notched) and in oth- ers (e.g., E. tenebrionis), it is truncate. In some frogs of the E. diastema group, the pad is lanceolate (i.e., having a pointed tip) and bears a papilla on the underside of the disc cover or extending as a free papilla beyond the margin of the disc cover. Even when no disc is developed, the ventral pad may be present (= "narrow discs" or "discs absent" auctorum), but the disc may be incompletely de- fined, (i.e., circumferential disc evident only dis- tally). The heavy-bodied, toadlike E. anomalus, cerastes, helonotus, and necerus lack digital discs on the fingers and either lack digital pads or have poorly developed, incomplete circumferential grooves. In these frogs, the digital discs and pads are better developed on the toes. Toes. — The groups of characters associated with the feet mostly are the same ones on the hands. Relative lengths of digits: The relative lengths of Toes III-V differ (Fig. 16). In some species (e.g., E. anomalus, cerastes, and necerus). Toe V is shorter than Toe III (Fig. 16A). In several species (e.g., species of the E. conspicillatus group). Toe V is only slightly longer than Toe III, and the disc of Toe V does not reach the distal subarticular tubercle on Toe IV (Fig. 16B). In other species (e.g., mem- bers of the E. cerasinus group). Toe V is obviously longer than Toe III, but the tip of the disc does not reach the distal subarticular tubercle on Toe IV (Fig. 16C). In many species (e.g., E. apiculatus, chalceus, laticlavius, and rosadoi). Toe V is much longer than Toe III; the tip of the disc of Toe V reaches to the distal border of the distal subarticular tubercle on Toe IV (Fig. 16D). Lateral fringes and webbing: Conditions of lateral fringes on the toes are the same as those on the fingers, but in a few species (e.g., E. loustes), a fringe extends from the disc to the inner metatarsal tubercle on the median surface of Toe I. The lateral fringes coalesce to form "webbing" in a few spe- cies. We avoid using the term "webbing" to de- scribe conditions in which the lateral folds coalesce but do not encompass the basal subarticular tu- bercles; this condition has been termed "basal webbing." Webbing is measured to subarticular tubercles in the manner proposed by Savage and Heyer ( 1 967 ) as modified by Myers and Duellman ( 1 982 ). If webbing does not encompass the basal subarticular tubercles, the webbing is termed basal. Among the species of Eleutherodactylus in west- 34 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 1 4. Palmar views of hands showing lengths and thickneses of digits in Eleutherodactylus. A. E. w-nigrum, ICN (JDL 1 1265), Finger 1 > II, digits slender and long (discs present). B. E. helonotus, BM 1970.178, Finger I > II, digits slender and short (discs absent). C. E. celator, KU 1 3 1 574, Finger I < II, digits short and broad (discs present). D. E. duellmani, USNM 286044, Finger I < II, digits long and broad (discs present). Scale bars = 2 mm. ELEUTHERODACTYLUS IN WESTERN ECUADOR 35 USNM 285970 narrow. B. E chalceus, KU 119484, cuspidate bearing terminal papilla. C. E crenunguis. KU 1^0126, expanded and emarginate. D. E suhsigiUatus. KU 131567, expanded and rounded. Scale bars = ^ mm 36 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 1 6. Plantar views of feet showing variation in lengths of Toes III and V in Eleutherodactyhis. A. E. cerastes, UVC 8429. Toe V shorter than Toe III (Condition A in Appendix III). B. E. babax, USNM 285970, Toe V sightly longer than Toe III (Condition B in Appendix III). C. £. duellmam, USNM 286044, Toe V longer than Toe III (also Condition B in Appendix III). D. E. omatissimiis, KU 1 19745, Toe V much longer than Toe III (Condition C in Appendix III). Scale bars = 2 mm. Fig. 1 7. Plantar views of feet showing variation in webbing in Eleutherodactylus. A. E. anatipes, ICN 12113. B. E. anomalus, CAS-SU 10467. C. E. longirostris, USNM 150481. D. E. loiistes, KU 179243. Scale bars = 2 mm. ELEUTHERODACTYLUS IN WESTERN ECUADOR 37 ern Ecuador, measurable toe webbing occurs only in four species — E. anafipes, anoimiliis, lofii^i- rostris, and loustes (Fig. 17). Webbing extends to the discs on Toes I-III and V in E. amitipes and nearly so in E. anomaliis, but extends only to a point between the distal and penultimate subarticular tubercles on Toe IV. Webbing also is obvious in E. longiwstris, but it does not extend appreciably beyond the basal subarticular tubercles except be- tween Toes IV and V. The webbing in E. loustes is less extensive than in E. longirostris. Discs and pads: The structure of the discs and pads on the toes closely parallels that of the fingers, but in many species having little or no evidence of discs and pads on the fingers, those on the toes are obvious; this point was made by Boulenger ( 1 898) in his choice of the trivial name for E. anomalus. However, a common condition in Eleutherodactylus is that the discs on the fingers are larger than those on the toes. Commonly, the discs on the inner toes are smaller than those on the outer toes, but in species having a short Toe V, the disc on Toe V is noticeably smaller than that on Toe IV. Coloration. — Although many other kinds of frogs can be identified easily from their dorsal coloration, such is not the case with most Eleutherodactylus, most of which are various shades of brown or gray dorsally, with or without distinct or diffuse darker markings. These markings may consist of spots of varying sizes, blotches, longitudinal stripes, trans- verse bars, chevrons (V-shaped marks with the apex anteriad), or W-shaped mark in the occipital-scapu- lar region. The exceptions in western Ecuador are E. chalceus, which is creamy tan with red spots or reticulations, and E. ornatissimus, which is green with black lines and spots. This problem of identifi- cation is confounded by distinct dorsal-pattern poly- morphism in some species (e.g., E. calcarulatus, laticlavius, vertebral is, and walkeri), whereas some other species (e.g., E. achatinus, phoxocephalus, and w-nigruni ) simply are highly variable in coloration. A common pattern polymorphism in many species is the presence of a pale middorsal stripe that may be yellow, orange, red, or green in life. In most species, dark transverse bars are present on the dorsal surfaces of the forearms and shanks (also on thighs and tarsi in some species), but dark bars are absent in E. chalceus. Ventral coloration also is highly variable and usually consists of various shades of brown or gray with or without pale or dark flecks or mottling. However, a few have uniform pale venters — white in E. chalceus and E. eremitus, creamy-yellow in E. achatinus and E. anatipes, and greenish-yellow in E. ornatissimus. Two species have distinctive pat- terns on the throat and/or chest; in E. caprifer a pair of dark longitudinal lines is present on the throat, and in E. longirostris, pairs of dark spots are present on the throat and chest. Use of ventral coloration in identification of species is further confounded by sexual dimorphism in some species. Females ofE. degener and E. subsigillatus have dark reticula- tions on the belly, whereas the belly in males is uniform. Most Eleutherodactylus in western Ecuador have two or three dark labial bars, a dark canthal stripe, and a dark supratympanic stripe. In a few species (e.g., E. colonial), the entire side of the head is dark, so that canthal and supratympanic stripes are not distinguishable; facial markings are absent in E. degener. A pale labial stripe appears as a polymor- phism in E. achatinus, illotus, longirostris, and walkeri. The color pattern on the flanks (exclusive of the axilla and groin) is distinctive in several species of Eleutherodactylus in western Ecuador. The flanks are pale with large black spots in E. w-nigruni, black with white spots in E. cajamarcensis, and brown with thin diagonal black lines in E. caprifer Eleutherodactylus luteolateralis has pale dorsolat- eral stripes, and E. parvillus has a pale yellow patch on the posterior half of the flanks. A dark reticulum encloses large, pale spots on the flanks in E. phoxocephalus, subsigillatus, and females of E. degener However, striking polymorphism in flank pattern occurs in E. quinquagesimus; in some indi- viduals, the flanks are dark with diffuse pale spots, whereas bold black and white diagonal bars are present on the flanks of other individuals. The color pattern on the posterior surfaces of the thighs is highly variable among species of Eleutherodactylus in western Ecuador. The most common pattern of a dark brown or black ground color with yellow, cream, or white spots or flecks occurs in 27 species (e.g., E. achatinus, crenunguis, duellmani. labiosus, and surdus): in five species (£". 38 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 ilegener, luiiniolac. Iwctiis. suhsiiiillatus. and walkeri), the pale spots or flecks may be present or absent. In 13 species (e.g., E. appendiculatus, longirostris, ocellatus, and sobetes), the posterior surfaces of the thighs are uniform brown, whereas in eight species (e.g., E. anatipes, anoinalus, and phoxocephalus) the thighs are cream or dull yellow with brown or black mottling or reticulation, and the thighs are brown, with black reticulations in E. helonotus and cream mottling in E. pteridophilus. The thighs also are cream or yellow, but with dark flecks, in E. actites and E. relator. The thighs are unpigmented in E. chalceiis and uniform cream in E. dissimidatus. In E. w-nignnn, the thighs (and flanks) may be yellow or white with large black spots or black with large yellow or white spots. Living individuals of a few species have distinctive coloration on the posterior surfaces of the thighs; also, the same color usually is on the underside of the shank and in the groin and axilla. These ex- amples are: deep blue in E. crucifer, green with pale spots in E. ornatissimus, yellow in E. eremitiis and E. parvillus, orange in E. dissimulatus and E. scolodiscus. and red in E. leoni and E. pyrrhomerus. In living individuals of most species of Eleutherodactyliis in western Ecuador, the iris var- ies from bronze to reddish-copper with varying amounts of black flecking and a variably distinct red or reddish-brown median, horizontal streak. We are convinced that the intensity of the color in the iris depends on the brightness of ambient light when observations are made. In some of the large, streamside species (E. anatipes, anonuilus, and necerus), the iris is pale bronze with brown or gray triangles (apex medial) anteriorly and posteriorly. However, distinctive iris colors are present in liv- ing individuals of several species — green in E. cerastes, blue in E. calcandatus and E. scolodiscus, bluish-gray in E. hectus and E. leoni, orange in E. degener and E. sobetes, red in E. vertebral is, choco- late-brown inf. duellmani, and black in E. chalceus. Adult sizes. — Body size is useful to distinguish species of Eleutherodactylus, but it must be used with care (Table 5). Because frogs of the genus Eleutherodactylus have direct development and because adults and juveniles occur in the same habitat, for at least some parts of the year there seems to be a continuum in sizes from a few to as much as 8()-l()() mm SVL. At other times, if repro- duction is seasonal, two clear modes of sizes are evident in a single sample of a species. Adulthood in males can be determined exter- nally if secondary sex characters (vocal slits and/or vocal sac, nuptial pads) are present, but adulthood is best verified by inspection of the testes. Repro- ductive males have larger, swollen testes than do nonreproductive males and immature individuals. In a few species, mesorchial pigmentation seems to be age-dependent (e.g., E. curtipes; Lynch, 1 98 1 a), but this is of limited value in western Ecuador where most species have white mesorchia. Unless ovarian eggs are visible through the body wall, adulthood in females can be determined only by inspecting the reproductive system. For nearly all eleutherodactyline frogs, convolution of the oviducts precedes development of large ova- rian eggs, and once the oviducts are convoluted, they do not regress during the nonreproductive season; however, this may not be true in E. labiosus (Lynch et al., 1994). Thus, females can be sorted into three age/size classes: ( 1 ) juveniles having thin, straight oviducts containing no enlarged ova- rian eggs; (2) subadults having thin oviducts with some convolutions containing only small or mod- erate-sized eggs; and (3) adults having strongly convoluted oviducts with large lumina, and ova- rian eggs enlarged or not. If there is regularity to sexual maturity (i.e., no regression after reproduc- tion), these three size classes seem to be largely sequential with only modest overlap between classes. Frogs of the genus Eleutherodactylus are mark- edly sexually dimoiphic in size. This is most obvi- ous in amplectant pairs, because males typically are only about two-thirds the size of the females. Some caution must be used when referring to amplectant pairs taken from collecting bags (and possibly field observations as well), because am- plexus only assures the observer of the male's maturity. We have found males in amplexus with other males, juvenile females, and adult females. Males of one species have been observed to amplex individuals of another species in the collecting bag, and taxonomic decisions should not be made on the assumption that the male "knew" what he was doing. Because of the sexual dimorphism in the ELEUTHERODACTYLUS IN WESTERN ECUADOR 39 Table 5. Body sizes (SVL in mm) of Eleuthcrochictylus in western Ecuador. Values include range (mean ± 1 SE. sample size). SD = sexual dimorphism. Species Males Females SD E. achatinus^ E. achatinus^ E. actites E. anatipes E. aiwmaliis E. apicidatiis E. appendiculatiis E. babax^ E. cajamarcensis E. calcarulatus E. caprifer^ E. celator E. cerastes E. chalceus E. colomai E. crenunguis E. crucifer E. degener E. dissimidatus E. diiellmani E. eremitus E. eugeniae E. floridus E. gentryi E. gularis E. hamiotae E. hectus E. helonotus E. illotus E. labiosus E. laticlavius E. latidiscus E. leoni E. longirostris E. loustes E. luteolatendis E. lymani E. muricatus^ E. muricatus^ E. necerus E. nyctophylax E. ocellatus E. omatissimus E. pannllus E. phoxocephalus E. pteridophilus E. pyrrhomerus E. quinquagesimus 23.0-35.1 (28.7 + 0.4,46 24.4-34.2 (28.8 ± 0.3, 42 30.0-40.0(35.0 + 0.4.32 42.9-64.3 (53.9. 4 31.5-61.0(47.0+ 1.4,46 17.8-21.8(20.3 + 0.2,24 18.8-21.0(19.7,3 42.4(1 19.2-24.1 (22.4 + 0.3,21 17.8-24.6 (22.2 ±0.3, 21 21.0-30.4 (25.8 ± 1.1.9 19.6-21.4(20.5 10.2,6 28.3-33.4 (30.8, 2 17.5-26.9 (23.5 ±0.4, 26 16.5-17.8(17.2,2 32.4-^9.2 (40.3 ± 1.9,9 18.3-20.6 (19.4 ±0.2, 9 22.2(1 24.9-36.0 (31.2 ± 1.2,32 17.2-21.8 (19.3 ±0.6. 7 26.0(1 17.4-18.3(17.8,2 23.0-28.5 (25.9 ±0.5, 13 20.2-21.6 (21.0 ±0.2, 9 28.2-30.9(29.6.2 13.6-16.8 (15.2 ±0.2. 13 25.9-29.3 (27.6, 2 35.4-50.8 (44.3 ± 1.6, 12 22.5-26.3(25.1,5 21.9-25.9 (23.8 ±0.3, 16 14.8-18.3 (16.7 ±0.5, 7 28.8-34.4 (31.9 ±0.5. 14 31.2-37.1 (34.8,6 16.6-23.6 (22.0 ±0.4, 10 25.7^3.6 (33.2 ± 1.2,23 18.7(1 31.8-40.7(36.0,4 44.9-68.4 (54.2, 4 21.9-31.4 (26.8 ±0.5. 21 21.6-27.0 (24.1 ±0.6, 9 15.5-19.6 (18.0 ±0.2, 27 22.3-29.9 (27.2 ± 0.4, 29 17.6-25.1 (22.4 ±0.3, 32 16.2-18.9(17.5,3 27.8-30.8 (29.0 ± 0.2, 12 33.6^5.4 (39.9 ± 0.6, 22) 37.0-46.1 (42.1 ±0.4,33) 48.2-64.2 (54.9 ± 1.0,21) 98.6(1) 76.5-92.4 (85.4 ± 1.5, 10) 21.6-26.3 (23.3 ±0.3, 17) 30.0-35.0 (32.2, 5) 45.6^8.7 (47.0, 6) 27.1-33.8 (29.8 ±0.5, 17) 25.3-28.9 (26.8 ± 0.2, 20) 40.5^3.8 (42.7, 3) 22.0-24.5 (23.5 ±0.3, 7) 44.4-55.8 (47.0 ± 1.2.9) 27.7-31.2 (29.7 ±0.3, 13) 59.1-64.5(61.9,3) 22.8-34.5 (26.8 ± 1.3,8) 31.0-31.9(31.4,2) 27.4-32.7 (29.9, 5) 36.6^5.8 (41.8 ± 1.3, 17) 27.1-27.6(27.4,2) 30.5-37.6 (33.6 ±0.9, 7) 23.3-26.7 (25.5 ±0.2, 17) 29.5-35.8 (33.4 ±0.8, 7) 21.7-24.8(23.5,5) 35.7-36.2 (36.0, 2) 19.4-22.5 (20.7 ±0.2, 20) 69.6(1) 38.3-44.6 (41.0 ±0.8, 7) 48.5-52.3 (50.4, 4) 37.2-^2.9 (39.9, 5) 35.2-53.4 (42.3 ± 1.3, 14) 19.7-25.0 (21.9 ±0.3. 22) 43.1-59.6 (48.2 ± 1.1. 15) 46.7(1) 25.6-29.5 (27.9, 5) 52.9-69.3 (60.2 ± 1.8, 10) 33.8-36.0 (34.9, 2) 83.8-93.3(88.3.4) 32. 1-37.8 (33.8 ±0.4, 22) 45.7(1) 37.8-38.1 (38.0,2) 18.4-25.9 (23.4 ±0.6, 11) 29.6-38.4(35.1 ±0.6,22) 31.9-33.9(33.1,3) 19.8-24.0 (21.6 ±0.3, 6) 33.6-40.1 (36.5 ±0.5, 14) 1.39 1.46 1.57 1.83 1.82 1.15 1.63 I. II 1 .33 1.12 1.66 1.15 1.52 1.26 1.54 1.38 1.41 1.34 1.42 1.29 1.43 1.29 1.36 1.48 1.14 1.59 1.78 1.31 1.51 1.34 1.27 1.81 1.89 1.63 1.26 1.58 1.30 1.29 1.48 1.23 1.26 40 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Table 5. Continued. Species Males Females SD E. rosadoi E. niidus E. scolodiscits E. simonboUvuri E. siopleus E. sobetes E. subsigillatus E. sidciilus E. siirdus E. tencbrionis E. thymalopsoides E. truebae E. itnistrigatus E. verecimdus E. vertebralis E. w-nigrum^ E. walkeri 16.5-16.7(16.6,2) 25.8-31.1 (28.9 ±0.7, 7) 17.6— 20.4 (1 9.2 ±0.3, 10) 16.0-19.2(17.2,4) 18.6-27.3 (23.7 ±0.7. 14) 19.3-28.5 (24.4 ±0.7. 17) 26.3 ( 1 ) 24.5-36.9(31.1 ±0.6.34) 20.8-26.8 (23.8 ±0.4, 16) 28.0-34.4(31.1.5) 25.5-39.0 (32.2 ± 1.2. 12) 14.5-24.0 (19.1 ±0.4, 46) 18.0-21.9 (19.5 ±0.6, 7) 21.1-28.0 (24.8 ±0.6. 15) 25.0^6.1 (33.0 ±0.6. 58) 13.8-18.5 (16.2 ±0.2, 24) 23.4-25.7 (24.4. 3) 1.47 37.1-39.8 (37.9 ±0.4. 7) 1.31 18.4-22.3(20.7.6) 1.08 18.5-22.0 (20.6 ±1.2. 22) 1.20 34.3^0.0(37.4.4) 1.58 41.3(1) — 30.0-33.4 (31.6 ±0.4. 8) 1.30 43.0 ( 1 ) 1.63 40.4-54.6 (45.8 ± 0.7, 14) 1.47 30.6-36.9 (33.8 ±0.8. 7) 1.42 46.9-55.4 (52.2 ±09, 9) 1.68 39.8^5.8(42.6.5) 1.32 23.0-31.0 (26.8 ±0.4. 22) 1.40 20.7-22.5(21.8.5) 1.12 35.0^3.9 (37.7 ±0.6, 20) 1.52 50.2-71.5 (59.8 ± 1.2,21) 1.81 19.4-25.3 (21.6 ±0.3, 23) 1.33 'Specimens from lowlands below 700 m. -Specimens from Pacific slopes (Balsapamba. Mindo, Tandapi). -^Specimens from southern Colombia and Ecuador. ■^Specimens from Ecuador only. ^Specimens from Rio Palenque and Santo Domingo de los Colorados (220-600 m) ^Specimens from Rio Faisanes ( 1380 m). ^Specimens from Pilalo. Provincia Cotopaxi. '^Specimens from Provincia Pichincha. sizes of adults, it is necessary to determine the sex and stage of matuiity to avoid the apparent continu- ous variation in size. Juvenile and young females are about the same size, or only slightly larger than, adult males and bridge the size gap that typically occurs between adult males and adult females. The last exacerbating factor relative to size is the tendency for frogs from higher elevations to be larger that conspecifics of the same age from lower elevations. In some species, the spotty nature of sampling demonstrates sharp differences in sizes between frogs that otherwise are alike (Heyer. 1970. for Leptodacty'lus: Lynch. 1980a. for E. nigrovittatus). Given these caveats, size data are extremely useful in sorting and distinguishing pre- served specimens of most, if not all. species of Eleutherodactyliis. Proportions. — We provide proportional data (ranges, means ± one standard error of the mean) in our descriptions, because proportions give preci- sion to such ambiguous modifiers as "wide," "long," or "short." While we agree that proportions are of little or no value in the identification of most species, we report the data at our disposal in order to permit more accurate comparisons than other- wise would be possible. Some proportional data are of limited use. For example, some species of the subgenus Craugastor and those of the E. sidcatus group are designated as "broad-headed," and the adjective can be expressed as a proportion. HW/SVL > 46%. Species of the E. diastema group have short hind limbs; this can be expressed most accurately as Tibia/SVL, but spe- cies not included in that group have equally short hind limbs. Many species in the E. conspicillatus and E. fitzingeri groups have long legs, and an accurate determination of relative leg length can be obtained from the ratio, Tibia/SVL. The concern of those investigators who eschew using proportions is that some users will make too much of a depar- ELEUTHERODACTYLUS IN WESTERN ECUADOR 41 ture of a tympanum/eye ratio relative to reported ranges when most of the difference possibly is owing to differences in technique of measurement and/or shrinkage. All of the measurements reported herein were taken by JDL, who, from time to time because of memory lapses or misplaced files, has measured some individuals repeatedly over a span of 20 years and finds only modest variation in values. Out of habit and tradition (which may be rooted in some forgotten reasoning), JDL measures snout- vent length (S VL), tibia length, greatest head width (HW), an estimate of head length (straight line distance parallel to body axis from angle of jaw to point equal to tip of snout) (HL), chord of head length (tip of snout to posterior angle of jaws), upper eyelid width (perpendicular to outer edge of eyelid), interorbital distance (lOD), horizontal di- ameter (length) of tympanum, length of orbit (= visible eye), and distance between eye and nostril. Five proportions are calculated from these 1 0 mea- surements—Tibia/SVL, HW/SVL, eyelid/IOD, tympanum/eye, and E-N/eye. Subgenera and Species Groups For the past quarter century, the genus Eleuthewdactyhis has grown in size to the point that most of our colleagues think that the genus should be divided into several smaller genera. As the number of known species has grown, so too has developed the necessity for some book-keeping devices (species groups) to sort the many species into at least phenetic groups. Lynch (1976b) pro- posed a series of species groups for the many species then known from South America and at- tempted to review the species group arrangement for the entire genus. Among the least useful conse- quences of that review was the recognition of an Eleuthewdactyhis unisthgatus group for more than 100 species distributed primarily within South America; most colleagues thought this group too large to be useful. Subsequently, Lynch and Duellman (1980) proposed using assemblies as a category within species groups, especially for the E. unistrigatus group. They assigned all species from the Amazonian slopes of the Ecuadorian Andes to assemblies and allocated many other extralimital species to those assemblies. Those proposals of species groups and assemblies were admittedly phenetic but based on the hope that the monophyly of at least some of the units could be established. The use of such units was driven pri- marily by pragmatism. During the past decade Eleuthemdactylus sys- tematics has undergone something of a revolution as the operational taxonomic work (including the recognition of species groups defined only by con- tent) was joined with explicit searches for synapomorphies and the adoption of a convention that only monophyletic units be recognized. Lynch (1986a) suggested that the subgenus Craugastor might be used for the Middle American clade of Eleuthemdactylus. In Craugastor, the mandibular ramus of the trigeminal nerve lies medial to the m. levator posterior mcmdibulae subexternus (= "e- condition" of adductor muscle), whereas in all other Eleutherodactylus (and all other lepto- dactylids), the mandibular ramus of the trigeminal nerve is lateral to the m. levator posterior mcmdibulae subexternus (= "s-condition" of the adductor muscle). Obviously, only the "e-condi- tion" is phylogenetically informative. Hedges (1989) proposed the recognition of a total of five subgenera, one of which (Eleuthero- dactylus) was viewed by him as paraphyletic be- cause no synapomorphies were known. Hedges (1989) and Savage (1987) also employed series, a category between subgenus (or genus) and species group. Savage (1987) suggested that Lynch and Duellman's (1980) assemblies should be treated simply as species groups on the grounds that the assemblies appeared to be coordinate categories to his species groups. We agree with Savage's sugges- tion concerning assemblies because we think they have outlived their usefulness. The 63 species we recognize or anticipate to occur in western Ecuador are placed by us in 15 species groups in two subgenera. The species groups are defined using the format employed by Savage (1987) with only minor modifications. We also employ "series," a category between subgenera and species groups, to reflect our perception of systematic arrangement and as a "temporary de- vice" until subgenera can be diagnosed effectively. We restrict usage of Series to the subgenus 42 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 E. abbotti Group; Pelohus ^•"^ ''^'C^KT y C3 «s=i£5ii^^ -a*. E. alfredi Group — --v J E. wightmanae^ E. conspicillatus Group E. ramagii Group m E. lacteus m^ Group ...^ Fig. 18. Distribution of the genus Eleutherodoctylus (stippled and hatched areas plus Lesser Antilles). Stippled area is inhabited by species having plesiomorphic state of relative lengths of toes ( Toe V shorter than Toe III). Hatched areas are inhabited by species having Toe V longer than Toe III but not extending to distal subarticular tubercle of Toe IV. (See Appendix III.) Eleutherodactylus sensu Hedges ( 1 989). When diag- nostic synapomorphies are known to us, they are identified as such by asterisks (*). All named species oi Eleutherodactylus are listed with respect to subge- nus, series, and species group in Appendix 111. Variation in toe lengths as a character of sys- tematic importance. — The 15 species groups can be assembled into three clusters based on variations in the lengths of the third and fifth toes (Fig. 16). In thef. anomalus, bufoniformis,fitzingeri, and sulcatus groups. Toe V is shorter than Toe III, whereas in the E. diastema and E. unistrigatus groups. Toe V is much longer than Toe III. (The disc on Toe V extends to level of distal subarticular tubercle on Toe IV.) In the other nine species groups. Toe V is longer than Toe III. Each toe reaches about to the level of the penultimate subarticular tubercle on Toe IV in the frogs of the E. conspicillatus and E. myersi groups. In the other seven groups, the Toe V reaches about half way between the distal and penultimate subarticular tubercles on Toe IV. This variation seems to be significant. If all ELEUTHERODACTYLUS IN WESTERN ECUADOR 43 Fig. 19. Hatching defines the distribution of species of Eleutherodactylus having Toe V much longer than Toe III. (See Appendix III.) The distribution of the genus is stippled. leptodactylids except Eleutherodactylus are used as an outgroup, a shoil fifth toe (Condition A in Appen- dix III) is plesiomorphic (found in all other leptodactylids) and is seen in some species of all subgenera of Eleutherodactylus recognized by Hedges ( 1989) except Pelorius, whereas the other conditions are not seen in leptodactylids other than in the genus Eleutherodactylus. The fifth toe being longer than the third but not extending to the distal subarticular tubercle on the fourth toe (Condition B in Appendix III) is sporadic in distribution (the E. alfredi group of Craugastor, one Cuban species of Euhyas, all Pelorius. and many South Ainerican species of the subgenus Eleutherodactylus) (Fig. 18). The spotty distribution of this condition and its incompatibility with the "e- condition'ofthe adductor mandibulae( Lynch, 1986a) suggest that the third condition has evolved repeat- edly. The fifth toe being much longer than the third (Condition C in Appendix III) is confined to a subset (186 named taxa) of the species assigned to the subgenus Eleutherodactylus by Hedges ( 1989) and Duellman ( 1993), primarily those assigned to the E. auriculatus and E. unistrigatus species groups. In contrast, the condition of the fifth toe being much longer than the third seems to be a uniquely derived feature (Fig. 19). The classificatory consequence of 44 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 such a conclusion is that the 186 named species, including E. martinicensis, the type species of Eleuthewdactyhis. having the liflh toe much longer than the third would constitute a subgenus Eleiithewdactyhis (sensu stricto) and some 150 other species placed in that subgenus by Hedges (1989) and Duellman (1993) would need to be assigned to some other subgenus or subgenera. Subgeneric names are available for some of these clusters. In addition to a subgenus Eleuthewdactyhis (cluster for the E. diastema and E. unisthgatus groups), Liiiinophys could be ap- plied as a subgenus for the E. sidcatus group. The third cluster (9 groups) is diagnosed by a toe-length character that we suspect to be convergent and, thus, are reluctant to apply any of the available subgeneric names (e.g., Phstimantis, Trachy- phrymis). Therefore, we continue Hedges' (1989) practice of recognizing a subgenus Eleuthew- dactyhis as a catchall for species and groups not included in the subgenera Craugastor, Eiihyas, PeU^hus, and Syrrhophus. However, to identify the apparent systematic structure, we employ the in- formal category series for apparent subdivisions within this subgenus. Subgenera Of the five subgenera oi Eleutherodactyhts rec- ognized by Hedges (1989), only two {Craugastor and Eleutherodactyhts) occur \n Ecuador. The other three subgenera include one (Syrrhophus) that is essentially Mexican and two (Euhyas and Pelorius) that are restricted to the Greater Antilles. Our use of Hedges' (1989) classification is not an unqualified endorsement of his analysis of his dataset. Rather, we find his proposed classification to be a useful point of departure for discussion of relationships within this large genus. We are skeptical of the monophyly of the subgenera Euhyas (as defined by Hedges, 1989) and of /'Wom/.v (Lynch, 1996), and we view the subgenus Eleutherodactylus as a nonmonophyletic assemblage of species, as evi- denced by the disparity between Hedges's and our treatments of series and species groups included therein. The basis of an alternate informal classifi- cation proposed by Savage ( 1 987 ) overemphasized the importance of variation in the ///. depressor mandihulae. For reasons given by Lynch (1993), we doubt the validity of the six lineages identified by Savage (1987). In order to provide some measure of compara- bility with the groups found in western Ecuador, we provide brief definitions of the three extralimital subgenera before addressing the subgenera and species groups that occur in western Ecuador. Subgenus Syrrhophus Cope, 1 878 Definition. — Mostly small frogs (SVL in fe- males 21—40 mm) having the "s-condition" of adductor musculature; head narrow; cranial crests absent; frontoparietals fused with prootics; vomer- ine odontophores absent; skin on venter areolate or smooth; Finger I about equal in length to Finger II; vocal slits present (except in E. longipes); nuptial pads absent; tympanic membrane and annulus prominent, not sexually dimorphic in size; super- numerary tubercles prominent, numerous; Toe V shorter than Toe III; toe webbing absent. Content and distribution. — Twenty-four spe- cies (Duellman, 1993) distributed from Texas to Guatemala. Remarks. — Hedges (1989) suggested that Syrrhophus and Euhyas are sister groups. This was based in part on their common possession of frontoparietal-prootic fusion. However, they are unique in having this fusion (Lynch 1971b; Savage, 1987). Subgenus Euhyas Fitzinger, 1 843 Definition. — Minute to large, mostly small to medium-sized frogs (SVL in females 12-83 mm) having the "s-condition" of adductor musculature; head narrow; cranial crests absent; frontoparietals fused with prootics; vomerine odontophores present, broad and arched in about half the known species; skin on venter smooth; Finger I shorter than Finger II; vocal slits usually absent (present in about 25% of species); nuptial pads absent; tympanic mem- brane and annulus prominent, not sexually dimor- phic in size; supernumerary tubercles prominent, numerous; Toe V shorter than Toe III; toe webbing absent, except in E. orcutti. Content and distribution. — Approximately 80 ELEUTHERODACTYLUS IN WESTERN ECUADOR 45 recognized species (Dueilman. 1993) primarily distributed in Cuba. Hispaniola. and Jamaica, with two species on Puerto Rico {E. lentiis and E. ricluiioiuli) and one on Mona Island (E. mouensis). Remarks. — Hedges (1989) inexplicably as- signed E. lentus and E. richmondi to the subgenus Eleuthemdactylus, but Joglar ( 1 989) included both of these species in his E. ricorclii group, which we recognize as belonging to the subgenus Euhyas. Each species shares the distinctive arched vomer- ine odontophores of many other Euhyas. Subgenus Pe/orm^ Hedges, 1989 Definition. — Medium-sized to large frogs (S VL in males 41-73 mm) having the "s-condition" of adductor musculature; head narrow; cranial crests present in E. inoptatus group, absent in E. ruthae group; frontoparietals not fused with prootics; vomerine odontophores present, prominent, short to moderately broad and arched; skin on venter smooth; Finger I usually longer than Finger II; vocal slits present; nuptial pads absent; tympanic membrane and annulus prominent, not sexually dimorphic in size; supernumerary tubercles promi- nent, numerous; Toe V longer than Toe III; toe webbing absent. Content and distribution. — Six species in two groups on Hispaniola (Dueilman, 1993). Remarks. — Unlike other Eleutherodactylus, males of the subgenus Pelorius are not markedly smaller than conspecific females ( Hedges and Tho- mas, 1987). Despite this putative synapomorphy, the monophyly of Pelorius is questionable (Lynch, 1996). Subgenus Craugastor Cope, 1862 Definition. — Minute to huge frogs (SVL in females 14-120 mm) having the "e-condition" of adductor musculature*; head narrow, except in E. anomalus, biporcatus, and bufonifonnis groups; cranial crests absent, except in E. biporcatus and E. bufonifonnis groups; frontoparietals not fused with prootics; vomerine odontophores large, triangular in outline, and nanowly separated in most taxa (absent in E. hobartsniitlii and E. pygmaeus: slanted and widely separated in E. sartori): skin on venter smooth, except areolate in E. omiltemanus group and some members of E. rhodopis group; Finger I longer than Finger II in most taxa (shorter in some members of E. omiltemanus and E. rhodopis groups); vocal slits usually present; nuptial pads usually present; tympanic membrane and annulus prominent in most species, sexually dimorphic in size, except E. alfredi, anomalus, and bufonifonnis groups, and parts of E. augusti and E. fitzingeri groups; supernumerary tubercles usually absent (present in E. augusti and E. rhodopis groups); Toe V shorter than Toe III (longer in E. alfredi group); toe webbing obvious in E. anomalus, fitzingeri, gollmeri, matudai, and rugulosus groups. Content and distribution. — At present, the subgenus is divided into 1 1 species groups (the E. alfredi, anomalus, augusti, biporcatus, bufoni- fonnis, fitzingeri, gollmeri, matudai, omiltemanus, rhodopis, and rugulosus groups) to accommodate 74 species. In addition, E. uno is not assigned to a species group. Two groups, the E. anomalus group and E. bufonifonnis group, are proposed here. Frogs of subgenus Craugastor are distributed pri- marily in Middle America, from Arizona and Texas through Panama, into western Colombia and Ecua- dor. In addition, there is a single species, E. maussi, in Venezuela. Craugastor is the most diverse sub- genus of eleutherodactyline frogs in Middle America, which also is inhabited by 24 species of the subgenus Syrrhophus (Texas to Guatemala) and 10 species of the subgenus Eleutherodactylus (Honduras to Panama). Remarks. — The subgenus Craugastor, as rec- ognized by Lynch (1986a) and Hedges (1989), includes the nominal genus Hylactophryne. How- ever, Savage (1987) sometimes recognized Hylactophryne in his arrangement of "Section I" of the genus Eleutherodactylus, and at other times, treated the three species of Hylactophryne as one of seven species groups within the ''Eleutherodactylus fitzingeri series." For example, in Savage's ( 1987:47) cladogram, Hylactophryne and two "lin- eages" are grouped in Section I, whereas elsewhere (Savage, 1987:44) one lineage (presumably Lin- eage 1 ) is stated to contain "£. alfredi, E. biporcatus, E. fitzingeri series, and E. omiltimanus [sic] group." Reanalysis of Savage's data suggests that some plesiomorphic similarities were used to group pre- 46 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 sumed entities (e.g.. Hykictophryne + Lineage 1 ). Each of Savage's arrangements has merit, but here we opt for a third arrangement in which there are no series recognized within Craugastor. Thus, tenta- tively we accept Savage's ( 1987:48) arrangement but with two modihcations: ( 1 ) recognition of the E. biporccitus and E. hufoniformis groups within what Savage appears to have viewed as the E. /^Z/wrta/Hi series (following Lynch. 1976b. 1986a). although we do not view these groups to be closely related to one another; and (2) fragmentation of Savage's E. rugidosiis group into two groups — the E. anomalus group and the E. nigidosus group. A full critique of Savage's arrangement will be pre- sented elsewhere (JDL. in prep.). Eleutherodactyhis {Craugastor) anomalus Group Definition. — Large frogs (SVL in females 76- 106 mm) with narrow to moderately wide heads (HW 37^8% SVL) and short snouts; cranial crests absent; upper eyelids much broader than lOD; body stout; limbs moderately long; skin on venter smooth; vomerine odontophores large, triangular in outline, narrowly separated medially; vocal slits present (except in E. anomalus), nuptial pads present; tympanic membrane and annulus rela- tively small, obscure, lacking sexual dimorphism in size. Discs on fingers and toes narrow (except in E. anomalus); Finger I longer than Finger H; subarticular tubercles not projecting; supernumer- ary tubercles absent; tarsus bearing fold on inner edge in E. anomalus and E. zygodactyl us (absent in other species); Toe V shorter than Toe IH; toes moderately webbed to well webbed. Content and distribution. — Four species (E. anatipes, anomalus, cheiroplethus, and zygo- dactylus) distributed in western Colombia and Ec- uador; two species {E. anatipes and E. anomalus) in western Ecuador. Remarks. — The E. anomalus group has not been recognized previously; formerly its members were assigned to the E. rugulosus group (Savage et al.. 1988; Lynch. 1990). The species in the E. rugulosus group (sensu stricto) have pronounced sexual dimorphism in tympanum size. Eleutherodactyhis ( Craugastor) hufoniformis Group Definition. — Large frogs (SVL in females 52- 94 mm) with broad heads (HW 43-58% SVL) and short snouts; cranial crests relatively low, parasagittal; uppereyelids much broader than lOD; body stout; limbs moderately long; skin on venter smooth; vomerine odontophores large, triangular in outline, narrowly separated medially; vocal slits present (E. necerus) or absent (E. hufoniformis); nuptial pads present; tympanic membrane and an- nulus relatively small, obscure, lacking sexual di- morphism in size. Discs on fingers and toes narrow; Finger I longer than Finger II; subarticular tu- bercles not projecting; supernumerary tubercles absent; tarsus lacking fold; Toe V shorter than Toe III; inner metatarsal tubercle laterally compressed; toes webbed basally (E. hufoniformis) or unwebhed (E. necerus). Content and distribution. — Two species — E. hufoniformis in western Colombia, Panama, and eastern Costa Rica, and E. necerus in western Ecuador. Remarks. — Previously, the E. hufoniformis has not been recognized formally; formerly its mem- bers were assigned to the E. hiporcatus group or series (Lynch, 1975a, 1986a; Savage, 1987). These two species do not have the strong sexual dimor- phism in tympanum size typical of species in the E. hiporcatus group (E. aphanus, hiporcatus, and maussi) and most other species in the subgenus Craugastor Eleutherodactyhis ( Craugastor) fitzingeri Group Definition. — Medium-sized frogs (SVL in fe- males 34-74 mm) with narrow heads (HW 33- 41% SVL) and long snouts; cranial crests absent; uppereyelids not broader than lOD; body slender; limbs long; skin on venter smooth; vomerine odontophores large, triangular in outline, narrowly separated medially; vocal slits and nuptial pads present; tympanic membrane and annulus promi- nent, strongly sexually dimorphic in size in E. crassidigitatus, fitzingeri, longirostris, melano- stictus, monnichorum, raniformis, rayo, and ELEUTHERODACTYLUS IN WESTERN ECUADOR 47 talamancae (no dimorphism in E. audi, cauquero, or emcelae). Discs on all fingers and toes, those on outer fingers emarginate in many species; Finger I longer than Finger II; subarticular tubercles not projecting; supernumerary tubercles absent; tarsus bearing fold on inner surface; Toe V shorter than Toe III; toes webbed basal ly except in E. talamancae (no webbing). Content and distribution. — Eight species in Central America (Honduras southward) and north- western South America. Although three species (£. fitzingeri, longirosths, and ranifonuis) are com- mon and widespread in western Colombia, only E. longiwstris extends into western Ecuador. Remarks. — Savage (1987) defined the E. fitzingeri group and included 12 species, but we include only the eight species exhibiting sexual dimorphism in the size of the tympanic annulus; JDL thinks that E. bocourti should not be included in the group, and we have doubts about the inclu- sion of three Talamancan species {E. andi, cauquero, and emcelae). Subgenus Eleutherodactylus Dumeril and Bibron, 1841 Definition. — Small to large frogs (maximum SVL in females 15-75 mm) having the "s-condi- tion" of adductor musculature; head narrow, except in E. sulcatus group; cranial crests absent, except in E. curtipes, devillei, sulcatus, and surdus groups, and a few members of other groups; frontoparietals not fused with prootics; vomerine odontophores usually present, slanted and widely separated to large, triangular in outline, and narrowly separated; skin on venter areolate; Finger I shorter than Finger II (longer in E. binotatus, conspicillatus, discoidalis, dolops, guentheri, nigrovittatus, and sulcatus groups); vocal slits usually present; nuptial pads present in most species in northwestern South America, absent in all species in southeastern Bra- zil and the West Indies; outer and middle ear structures absent in about two dozen taxa (e.g., E. anotis, baryecuus, ruidus, and surdus): otherwise, tympanic membrane present or absent and annulus present, not sexually diinorphic in size; supernu- merary tubercles absent to prominent and numer- ous; Toe V shorter than Toe III in some groups (e.g.. E. discoidalis and E. sulcatus groups), longer than Toe III in several groups (e.g., E. conspicillatus, devillei, and myersi groups), and much longer than Toe III in the E. martinicensis series*; toe webbing absent, except in E. diaphonus and E. pugnax. Content and distribution. — Approximately 336 named species (number updated from Duellman, 1993) occurring primarily in South America, with 10 species in Central America and 35 species in the West Indies (absent in Jamaica). Remarks. — At present, we are not willing to disrupt the subgeneric classification proposed by Hedges (1989), although we think that we have detected a large (186 species) monophyletic unit within the subgenus; herein, we refer to that unit as the E. martinicensis series. Little is to be gained by recognizing additional subgenera when most taxa within the subgenus Eleutherodactylus remain to be studied in detail. To restrict the subgenus Eleutherodactylus to some 186 species might be tempting to many of our colleagues (as it is to us) and irresistible to some, but such action would leave some 150 other species in oblivion in the taxonomic hierarchy. Thus, we employ the expedi- ent, but nomenclaturally informal, category of se- ries to express our emerging notions of relation- ships among species within the subgenus Eleutherodactylus (sensu Hedges, 1989) and rec- ognize four series to accommodate the species in western Ecuador. We are aware of no less than five such series; one series including the entire comple- ment oi Eleutherodactylus in southeastern Brazil is not discussed here (see Lynch, 1976b). The Eleutherodactylus myersi Series Eleutherodactylus ( Eleutherodactylus) myersi Group Definition. — Small frogs (SVL in females 17- 27 mm) having narrow heads (32^2% SVL) and short snouts; cranial crests absent; upper eyelid naiTower than lOD (except in some males); body robust; limbs short to moderately long; skin on venterareolate; vomerine odontophores small, oval or oblique, usually well separated medially; vocal slits present (except E. floridus), nuptial pads ab- sent (erroneously reported as present in E. myersi 48 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 by Lynch. 1980a); tympanic membrane present (except in E. leoni and E. ocrcatHs): tympanic annulus relatively large, not sexually dimorphic in size. Discs on fingers and toes narrow (widest in E. fiohdus): pads absent on inner fingers in E. ocreatus; Finger I shorter than Finger II; subarticular tu- bercles not projecting; supernumerary tubercles small, numerous; tarsus bearing low tubercle on inner edge (absent in E. ocreatus); Toe V slightly longer than Toe III. Content and distribution. — Nine species in the Andes of southern Colombia and Ecuador, of which four species (E. jioridus. hectus, leoni. and pyrrhomerus) occur in cloud forests in western Ecuador. Remarks. — The only available generic name for this group is Tnichyphrynus Cochran and Coin. 1963 (type species T. //n'£'/-5/). At present, there is no evidence that the E. myersi group is monophyletic; nor is there evidence to the contrary. The Eleuthewdactylus sulcatus Series Eleiithewdactylus {Eleuthewdactylus) sulcatus Group Definition. — Medium-sized to large frogs (SVL in females 29-70 mm) with broad heads (49-63% SVL) and short snouts; cranial crests present, most prominent in females; upper eyelid not wider than lOD; body robust; limbs relatively short; skin on venter areolate (E. helonotus, ingeri, ndzi, and sulcatus)ox?,VL\oo\\\ (E. cadenai, cerastes, cornutus, and sernai); vomerine odontophores large, triangu- lar in outline (nearly arched in some species), narrowly separated medially; vocal slits and nup- tial pads present in E. ruiz.i. absent in E. cerastes, cornutus, and .9/Y/f(7n/.v (condition unknown in other species); tympanic membrane and annulus promi- nent, some sexual dimorphism in size. Fingers lacking discs and pads; Finger I longer than Finger II; subarticular tubercles not projecting; supernu- merary tubercles numerous; tarsus bearing fold on inner edge inf. cornutus, ingeri, ruizi, and sulcatus (absent in other species); Toe V shorter than Toe in. Content and distribution. — Seven species in the Andes of Colombia and Ecuador and one spe- cies (E. sulcatus) in the upper Amazon Basin of Colombia. Ecuador, Peru, and western Brazil. Two species {E. cerastes and E. helonotus) are found in cloud forests in western Ecuador. Remarks. — Herpetologists long have been puzzled by a series of eleutherodactyline frogs having large, broad heads and massive cranial crests. Although Peters (1864) and Cope (1874) used the generic name Strahomantis, Boulenger (1882) set precedence by including the species known at that time within Hylodes (= Eleuthero- dactylus). Lynch (1975a) attempted an early sur- vey, which has undergone several revisions (Lynch, 1981a; 1986a, b), as additional species were dis- covered and as cladistic methodology was applied more rigorously. Our usage of the E. sulcatus group follows Lynch (1986a, b) wherein E. hiporcatus and E. maussi are assigned to the subgenus CraugastoK At present, no synapomoiphy is known for frogs of the E. sulcatus group, but it seems inappropriate to treat these eight species merely as part of an amorphous subgenus Eleutherodactylus. Pending publication of a revision in progress by JDL. we defer usage of the earliest available subgeneric name, Limnophys iimenez de la Espada, 1870. The Eleutherodactylus conspicillatus Series Eleutherodactylus (Eleutherodactylus) cerasinus Group Definition. — Small to medium-sized frogs (SVL in females 25-64 mm ) with naiTOw heads (37-42% SVL) and short snouts; upper lips flared in large females producing a less slender head shape; cra- nial crests absent; upper eyelid usually wider than lOD; body slender; limbs relatively long; skin on venter areolate (often appears smooth in large females); vomerine odontophores large, triangular in outline, nairowly separated medially; vocal slits usually present (absent in E. orpacobates and E. ruhicundus); nuptial pads usually absent (present in E. cerasinus and E. orpacobates); tympanic membrane and annulus present, relatively large, not sexually dimorphic in size. Discs on fingers and toes expanded; Finger I shorter than, or subequal to. Finger II; subarticular tubercles not projecting, except for basal subarticular tubercles; supernu- ELEUTHERODACTYLUS IN WESTERN ECUADOR 49 merary tubercles few or absent; tarsus lacking fold on inner edge; Toe V longer than Toe III. Content and distribution. — Six species occur in lower Central America and western Colombia and Ecuador; a seventh (E. nibicundiis) inhabits cloud forests on the Amazonian slopes of the Andes in Ecuador. Four species {E. crenunguis. lahiosus, ocellatus, and tenehrionis) occur in western Ecua- dor. Remarks. — Frogs of the E. cerasinus group have long, slender digits with large digital discs. The slenderness of the digits is accentuated be- cause the digits lack lateral fringes or keels that in other frogs make the digits appear to be thicker. Lynch et al. ( 1 994) proposed this species group, for which no synapomorphy is known. Eleutherodactylus {Eleutherodactylus) conspicillatus Group Definition. — Small to large frogs (SVL in fe- males 25-75 mm) with narrow heads (30^5% SVL) and long snouts; cranial crests absent; upper eyelid usually narrower than lOD, rarely as wide as, or wider than, lOD; body slender; limbs rela- tively long (short in E. vilarsi); skin on venter smooth (areolate in E. caprifer); vomerine odontophores large, triangular in outline, narrowly separated medially; vocal slits usually present (ab- sent in E. carmelitae, gaigei, illotus, insignitus, johannesdei, lymani, thectopTemus, and viridicaus); nuptial pads usually present (absent in E. caprifer, gaigei, illotus, insignitus, and viridicans); tym- panic membrane and annulus present, relatively large, not sexually dimorphic in size. Discs on fingers and toes expanded, those on inner fingers narrow; Finger I usually longer than Finger II; subarticular tubercles not projecting; supernumer- ary tubercles few or absent; tarsus bearing short fold on inner edge in many species; Toe V slightly longer than Toe III; toes unwebbed (basal webbing in E. malkini). Content and distribution. — Twenty-nine spe- cies in the lowlands and cloud forests from Costa Rica to Suriname and Bolivia. Several species inhabit the Andean slopes in Colombia and Ecua- dor, and six species [E. achatinus, actites, caprifer, illotus, lymani, and w-nigrum) occur in western Ecuador. Remarks. — The E. conspicillatus group was recognized by Lynch (1986a; to accommodate those species once assigned to the E. fitzingeri group (Lynch. 1976a, Lynch and Myers. 1983) that are not members of the subgenus Craugaston Ex- ternally, frogs of the E. conspicillatus group closely resemble many of those in the subgenus Craugastor, but even juveniles can be separated by the relative lengths of Toes III and IV (Lynch, 1994). Most species e^ccur in the lowlands from Costa Rica to southeastern Brazil and Bolivia, but mem- bers of this group also inhabit the highlands of the Sierra Nevada de Santa Marta (E. carmelitae and E. insignitus) and the Andes in Colombia, Ecuador, and northern Peru. The most widely distributed species (E.fenestratus, malkini, peruvianas, vilarsi, and zeuctotylus) occur east of the Andes. At present, there is no evidence that the E. conspicillatus group is monophyletic. Eleuthero- dactylus caprifer is assigned here (following Lynch and Myers, 1983) although it has areolate skin on the venter and Finger I is shorter than Finger II. Eleutherodact}'lus gaigei (Dunn) is assigned here despite its bearing little resemblance to any other species in the group (Lynch, 1980b). No generic name is available for frogs of the E. conspicillatus group. Eleutherodactylus {Eleutherodactylus) curtipes Group Definition. — Small to medium-sized frogs (SVL in females 26-50 mm) with narrow heads (29^2% SVL) and short snouts; cranial crests present; up- per eyelid narrower than lOD; body robust; limbs short; skin on venter areolate; vomerine odonto- phores small, slanted, widely separated medially; vocal slits absent; nuptial pads absent (present in E. huckleyiy, tympanic membrane and annulus absent (present in E. buckleyi). Discs on fingers and toes narrow; Finger I shorter than Finger II; subarticular tubercles not projecting, except for basal subarticular tubercles; supernumerary tubercles numerous; tarsus lacking fold on inner edge but indistinct tubercle present in some; Toe V longer than Toe III but not extending to distal subarticular tubercle on Toe IV. 50 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Content and distribution. — Six species occur at high elevations (usually above 3000 m) in the western Andes of Colombia and Ecuador south to the Nudo de Azuay; only E. gentryi is treated as part of the fauna of western Ecuador. Remarks. — Lynch (1995) proposed that the cranial crests of these frogs represent a synapo- morphy for the group. Many other highland Eleuthewdactyliis are known from the Cordillera Central of Ecuador and from Ecuador, but these lack cranial crests and have Toe V much longer than Toe III, as is characteristic of the subgenus Eleutherodactylus (sensu stricto). Eleuthewdactyliis ( Eleutherodactylus) dolops Group Definition. — Medium-sized frogs (SVL in fe- males 42-58 mm) with narrow heads (38^2% SVL) and long snouts; cranial crests absent; upper eyelid broader than lOD; body slender; limbs rela- tively long; skin on venter smooth; vomerine odontophores large, triangular in outline, narrowly separated medially; vocal slits present; nuptial pads absent; tympanic membrane and annulus present, relatively large, not sexually dimorphic in size. Discs on fingers and toes narrow, larger on toes than on fingers; Finger I much longer than Finger II; subarticular tubercles not projecting; supernumer- ary tubercles absent; tarsus lacking folds and tu- bercles; Toe V slightly longer than Toe III. Content and distribution. — Two species {E. babax and E. dolops) have been described; the former is distributed in cloud forests in western Colombia and Ecuador, the latter in cloud forests on the Amazonian slopes of the Andes in Colombia and Ecuador. There is an undescribed species on the western flanks of the Cordillera Central in Departamento de Caldas, Colombia. Remarks. — Lynch (1989) proposed this spe- cies group when he dismantled what he had earlier (Lynch, 1976b) termed the E. discoidalis group. The "synapomorphy'" cited by Lynch (1989) re- mains to be studied in a larger suite of eleuthero- dactylines; at present, it seems unlikely that the cited character (large nasal bones in median con- tact) is uniquely derived. Eleutherodactylus ( Eleutherodactylus) devillei Group Definition. — Medium-sized frogs (SVL in fe- males 28-52 mm) having narrow heads (35-43% SVL) and short snouts; cranial crests present (ab- sent in E. acatallelus and E. appendiculatus); up- per eyelid not wider than lOD; body slender to robust; limbs moderately short to relatively long; skin on venter areolate; vomerine odontophores prominent, triangular in outline, narrowly sepa- rated medially; vocal slits absent (present in E. acatallelus and E. appendiculatus); nuptial pads absent (present in E. vertebralis); tympanic mem- brane and annulus present (absent in E. siopelus), not sexually dimorphic in size. Discs on fingers and toes expanded; Finger I shorter than Finger II; subarticular tubercles not projecting; supernumer- ary tubercles present; tarsus lacking or having ill- defined fold along inner edge; Toe V longer than Toe III. but not extending to distal subarticular tubercle on Toe IV. Content and distribution. — Eight species oc- cur in cloud forests in western Colombia and Ecua- dor, and one species {E. devillei) inhabits the Ama- zonian slopes of the Andes in northern Ecuador. Of these, E. appendiculatus, quinquagesimus, truebae, and vertebralis) are in western Ecuador. Remarks. — We are not confident in associating these species but are convinced that within the group, E. devillei, truebae, and vertebralis form a subgroup, and that E. cacao and E. sulculus are sister species. In South America, there is a broad array of species having the intermediate fifth toe length and the general pattern of structure indicated in the definition. At present, too little is known of most species to be confident in delimiting this species group. Eleutherodactylus {Eleutherodactylus) loustes Group Definition. — Medium-sized frogs (SVL in fe- males 37-56 mm) with narrow heads (38-43% SVL) and long snouts; cranial crests present in females of E. jaimei and E. loustes; upper eyelid broader than lOD; body slender; limbs relatively long; skin on venter areolate; vomerine odonto- ELEUTHERODACTYLUS IN WESTERN ECUADOR 51 phores large, oval to triangular in outline, narrowly separated medially; vocal slits present: nuptial pads absent; tympanic membrane and annulus present, small in E. hyhotnigus and E. jaimei (membrane absent in E. loustes), not sexually dimorphic in size. Discs on fingers and toes expanded; Finger 1 usually slightly shorter than Finger II (except in E. loustes); subarticular tubercles not projecting; su- pernumerary tubercles few or absent; tarsus bear- ing fold along inner edge; Toe V longer than Toe III. but not extending to distal subarticular tubercle on Toe IV; toes unwebbed {E. hyhotragus) to about 1/ 3 webbed (E. loustes). Ventral edge of zygomatic ramus of squamosal expanded, evident externally as a knob immediately anterior to tympanic annu- lus*. Content and distribution. — Three species are known from the lowlands and cloud forests of western Colombia and northwestern Ecuador; only E. loustes is found in Ecuador. Remarks. — Lynch (1992a) proposed the E. loustes group based on the synapomorphy of the expanded zygomatic ramus of the squamosal. Prior to the association of these three species, E. loustes was suspected to be related to certain species in the E. cerasinus group; this was based on an over- emphasis by JDL of head shape; the upper lips of females of several species in the two groups are flared. Curiously, the toe length associates the two groups as well, but the condition of the moderately long fifth toe seems to have evolved repeatedly within Eleutherodactylus. Eleutherodactylus {Eleutherodactylus) orestes Group Definition. — Small frogs (SVL in females 18- 27 mm) with narrow heads (34-40% SVL) and short snouts; cranial crests absent; upper eyelid narrower than lOD; body robust; limbs relatively short; skin on venter areolate; vomerine odonto- phores oval, well separated medially; vocal slits present; nuptial pads absent; tympanic membrane ill-defined, annulus present; discs on fingers and toes narrow (only slightly wider than digit); Finger I shorter than Finger II; subarticular tubercles not projecting; supernumerary plantar tubercles nu- merous, low; ill-defined tubercle along inner edge of tarsus; Toe V longer than Toe III, extending to proximal border of distal subarticular tubercle of Toe IV; toes unwebbed. Content and distribution. — Three species (£. orestes, simonholivari, and vidua) inhabit paramos in southern Ecuador and high-altitude cloud forests in Provincia Bolivar, Ecuador. Remarks. — Because the fifth toe is consider- ably longer than the third, we initially considered these frogs to be associated with the E. martinicensis series. However, because there are several terres- trial paramo inhabitants (e.g., E. obmutescens, orcesi.philipi, racemus, simoterus, andthymelensis) in that series having a long fifth toe (extending to distal border of distal subarticular tubercle of Toe IV), we decided that these three small species are best disassociated systematically. The three spe- cies are structurally similar and have been con- fused (Wiens and Coloma, 1992) with frogs of the E. myersi group (frogs having much shorter fifth toes). The available osteological data are inad- equate for us to argue against several possible hypotheses of relationships, but we are inclined to continue in the spirit of Savage's (1987) approach in recognizing species groups so long as the data- base remains narrow. All three of these species have the frontoparietals fused with the prootics, in contrast to the plesiomorphic condition of unfused elements in most species of the E.unistrigatus group. Eleutherodactylus {Eleutherodactylus) surdus Group Definition. — Medium-sized frogs (SVL in fe- males 36-55 mm) with narrow heads (35^1% SVL) and short snouts; cranial crests prominent in E. surdus, less prominent in other species; upper eyelid about as wide as lOD; body robust; limbs relatively long; skin on venter areolate; vomerine odontophores prominent, triangular in outline, nar- rowly separated medially; vocal slits absent; nup- tial pads present {E. duellnumi and E. hamlotae) or absent; tympanic membrane and annulus absent. Discs on fingers and toes expanded; Finger I shorter than Finger II; subarticular tubercles not project- ing; supernumerary tubercles few; tarsus lacking fold along inner edge; Toe V longer than Toe III, but 52 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 not extending to distal subarliciilar lubeicic on Toe IV; toes unwebbed ( basal webbing in E. diiellmuni). Content and distribution. — Four speeies (E. (lui'llnuiiii, hamiotac, sobetes, and siirdus) inhabit cloud forests in western Ecuador. Remarks. — Since having been proposed by Lynch ( I98()c). the E. siirdus group has continued to undergo revision. Initially, the group was diag- nosed in part by the absence of tympana and tym- panic annuli, but a variety of other species lack these structures and are not thought to be related to this cluster of species in western Ecuador. Several of the species included by Lynch (1980c) in the group {E. baryeciius, piignax, and ruidus) have the fifth toe much longer than the third and are now assigned to the Eleiitliewdactyliis nuirtiniceiisis series. Osteological material is available for only two species in the E. surdits group, and, as yet, no osteological features are apparent that might be restricted to this group of frogs. The Eleutherodactylus martinicensis Series In frogs of this series. Toe V is much longer than Toe III; the tip of the fifth toe extends to the distal subarticular tubercle of Toe IV*. This feature is derived and provides evidence for a monophyletic grouping of some 1 86 species distributed through- out the West Indies (except Jamaica) and north- western South America. The type species of Eleutherodactylus is included in this series, which equals the subgenus Eleutherodactylus (sensu stricto). Eight Central American species (£". altae, caryophyllaceus, cruentus, diastema, hylaeformis, pardalis, ridens, and vocator) join with more than 140 species found primarily in Colombia, Ecuador, and northern Peru in composing the mainland com- ponent of the series. In South America, the series is absent from the Brazilian and Guianan highlands and from coastal Brazil (Fig. 19). Species are present along the Amazon and in the Guianas (£. inguinalis and E. marmoratus) and extend as far south as southern Peru {E. ockendeni and E. platydactylus). The only purported Mexican spe- cies (£. batrachylus) is thought by JDL to be a member of the West Indian fauna that became mislabeled as originating from Tamaulipas. Eleutherodactylus (Eleutherodactylus) diastema Group Definition. — Minute to small frogs (SVL in females 13-31 mm) with narrow heads (34^1% SVL) and short snouts; cranial crests absent; upper eyelid narrower than lOD; body robust; limbs short; skin on venter areolate; vomerine odonto- phores slanted and widely separated medially in E. chalceiis and E. scolodiscus, large, oval, and nar- rowly separated in other species; vocal slits present; nuptial pads absent; tympanic membrane present or absent, annulus present, not sexually dimorphic in size. Discs on fingers and toes large, usually cuspidate or bearing terminal papillae; Finger I shorter than Finger II; subarticular tubercles not projecting, bifid in some species; supernumerary tubercles numerous, ill defined; tarsus lacking fold along inner edge; toes unwebbed (basal webbing in E. gularis). Content and distribution. — Six species rec- ognized at present are distributed from Honduras to Ecuador; three of these (E. chalceus, gularis, and scolodiscus) occur in western Ecuador. Remarks. — At present, the E. diastema group is undiagnosed and may be an illusion, but these six species are placed in it pending further resolution. Eleutherodactylus chalceus and E. scolodiscus are distinctive among members of the group (and spe- cies of the genus in western Ecuador) in lacking vomerine teeth or having greatly reduced vomerine dentition. Cochran and Goin ( 1970) attempted to define the group based on the possession of longitudinal folds (of the vocal sacs) on the throat. This trait is of uncertain polarity or importance and was men- tioned first by Dunn (1926) when discussing rela- tionships among Cuban Eleutherodactylus. Dunn noted that several Cuban Eleutherodactylus were like E. diastema in having the longitudinal folds on the vocal sacs. Such folds may reflect some particu- lar character, but until throat musculature is inves- tigated in several groups of Eleutherodactylus (in- cluding members of the E. diastema group as well as the peculiar taxa in Cuba and Hispaniola), we defer to use this trait as diagnostic. Our reluctance is heightened because we do not see such distinc- tive folds in either E. chalceus or E. scolodiscus. ELEUTHERODACTYLUS IN WESTERN ECUADOR 53 Eleutherodactylus {Eleiithcwdactylus) unistrigatus Group Definition. — Small to medium-sized frogs (SVL in females 1 8-55 mm ) with narrow heads (3 1^3% SVL) and short snouts; cranial crests usually ab- sent (present in a few taxa, e.g., E. niidiis and E. thymalopsoides): upper eyelid usually as wide as, or wider than, lOD; body slender to robust: limbs moderately long (shorter in many highland spe- cies); skin on venter areolate; vomerine odonto- phores slanted and widely separated medially to large, triangular in outline, and narrowly separated; vocal slits usually present; nuptial pads usually present; tympanic membrane usually prominent, not sexually dimorphic in size. Discs on fingers and toes broad; Finger I shorter than Finger II; subarticular tubercles not projecting; supernumer- ary tubercles few in number to numerous and ill defined; tarsus usually lacking fold along inner edge (tubercle or foldlike tubercle in some spe- cies); toes usually unwebbed (basal webbing in E. diaphonus and E. pugnax). Content and distribution. — Approximately 150 species can be assigned to this group, but we are uncertain about the limits of the group. We confidently place five Central American species {E. altae, caryophyllaceus, cruentus, pardolis, and ridens) in this group, as well as one species (E. taywmi) from the Sierra Nevada de Santa Marta. This group is well represented in the Andes of Colombia and Ecuador and the upper Amazon Basin, but remains unknown from Venezuela. The species in western Ecuador are E. apiculatus, cajamarcensis, calcarulatus, celator, colomai, cru- cifer, degenei; dissimulatus, eremitus, eiigeniae, laticlavius, latidiscus, luteolateralis, muricatus, nyctophylax, ornatissimus, parvilhis, phoxo- cephalus, pteridophilus, rosadoi, ruidus, subsi- gillatus, thymalopsoides, unistrigatus, verecundus, and walkeri. Remarks. — One of us (JDL) was once confi- dent that the superficially similar Antillean group {E. martinicensis) would prove to be separable from the South American frogs placed by Lynch (1976b) in the E. unistrigatus group based on fusion of the frontoparietals and prootics in the former. The studies of Joglar (1986, 1989) sug- gested that more variation obtains among the Antillean representatives than once thought, but few South American taxa exhibit the fusion. Using the relative lengths of the toes to sort mainland Eleutherodactylus results in the discov- ery that the long fifth toe is curiously distributed. Most species with a long fifth toe occur in Colom- bia and Ecuador; only a few are distributed along the Amazon into northeastern South America (the Guianas). Especially curious is the near absence of such species from Venezuela and their absence from the Guiana Highlands. A number of species are known from Peru (Amazonian slopes as well as Amazonian lowlands) but only one (E. platy- dactylus) extends into Bolivia. No species having this toe condition occurs in association with the Brasilian Shield or along the once-forested Atlantic coast of Brasil. These observations do not assure that the E. unistrigatus group is monophyletic but suggest that it may be so. The long fifth toe condition is not an ecological association because although most species having the condition are arboreal, several are not. The terrestrial representatives are found in the high Andean grasslands of Colombia, Ecuador, and northern Peru. Key to the Species This key emphasizes external features mostly independent of maturity and sex. However, in at- tempting to identify Eleutherodactylus, there is no substitute for knowing the sex of the individual frog or for having an estimate as to whether or not it is mature. Juveniles are always difficult to iden- tify because the distinctness of the tympanic mem- brane and annulus increases with age, and many features of color patterns on the venter and con- cealed surfaces of the limbs develop with age. Furthermore, juveniles commonly are more warty dorsally than are adults. So long as the specimen is an adult, we think the key will enable quick and correct identifications. If the specimen is poorly preserved, we doubt that most users can identify the frog correctly using this key. Given these caveats, we agree with Stuart's (1955:10) qualification re- garding the use of keys: "... the worker who knows what species he has before him should experience few difficulties in its use." 54 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 This key docs not take into account the bright colors that disappear after extended storage in alcohol. If a living individual is at hand or colors in life are known, several species can be identified readily because the dorsum is green or because flash colors are evident in the axilla, groin, or concealed surfaces of the thighs. Refer to the plates and the descriptions of colors in life in the Accounts of Species. 1. Toe III longer than Toe V (Fig. 16B-D) 2 Toe 111 shorter than Toe V (Fig. 16A) 9 2. Toes webbed, webbing enclosing at least basal subarticular tubercles on all toes 3 Toes lacking webbing 5 3. Toe webbing extending to discs on some toes; skin of dorsum spiculate, tuberculate, or bear- ing numerous folds 4 Toe webbing not reaching disc on any toe; skin of dorsum smooth E. longirostris 4. Inner tarsal fold present; fingers lacking discs E. onomahis Inner tarsal fold lacking; fingers bearing nar- row discs E. cmatipes 5 Skin on venter areolate (granular) E. helonotus Skin on venter smooth 6 6. Inner edge of tarsus bearing tubercle; HW/ SVL<0.35 7 Inner edge of tarsus lacking tubercles; HW/ SVL>0.45 8 7. Tympanic membrane prominent; digit tips rounded; Finger I longer than Finger II Barycholos pulcher Tympanic membrane absent; digit tips swol- len; Finger I shorter than Finger II Phyllonastes sp. 8. One elongate tubercle evident on upper eyelid E. cerastes Upper eyelid tuberculate. none elongate E. necerus 9. Skin on venter smooth 10 Skin on venter areolate 18 10. Tip of Toe V extending about half way be- tween penultimate and distal subarticular tu- bercles of Toe IV (Fig. 16C); skin of dorsum shagreen or tuberculate 1 1 Tip of Toe V reaching no more than one fourth of the way between penultimate and distal subarticular tubercles of Toe IV (Fig. 16B); skin of dorsum smooth to shagreen, never tuberculate 13 1 1 . Enlarged conical tubercles present on upper eyelids E. labiosus Enlarged tubercles absent on upper eyelids . 12 12. Brown chevrons across throat; posterior sur- faces of thighs brown E. ocellatus Throat unicolor or reticulated; posterior sur- faces of thighs black with cream flecks E. crenunguis 13. Discs narrow, less than twice width of digit (Fig. 15A) E. babax Some discs broad (twice or more width of digit), especially on outer fingers 14 1 4. Inner surface of tarsus bearing fold extending at least one half length of tarsus 15 Inner edge of tarsus bearing tubercle or not, never a fold 16 15. Posterior surfaces of thighs and flanks densely stippled with black E. actites Posterior surfaces of thighs and flanks marbled black and cream E. lymani 1 6. Body lacking dorsolateral folds; usually black spots in groin E. w-nigrum Body with dorsolateral folds; groin lacking black spots 17 ELEUTHERODACTYLUS IN WESTERN ECUADOR 55 17. Small tubercle on heel; throat and chest spot- ted with black E. illotus Heel lacking tubercles; throat and chest im- maculate to reticulate E. achatinus 18. Tip of Toe V not reaching distal subarticular tubercle of Toe IV 19 Tip of Toe V reaching distal border of distal subarticular tubercle of Toe IV 40 19. Tympanic annulus and membrane absent (no external evidence of ear) 20 Tympanic annulus and membrane visible ex- ternally 28 20. Tip of Toe V extending slightly (or not at all) beyond distal border of penultimate subar- ticular tubercle of Toe IV (Fig. 16B) 21 Tip of Toe V extending beyond (by length of disc) distal border of penultimate subarticular tubercle of Toe IV (Fig. 16C) 23 21. At least some subarticular tubercles of foot enclosed by basal webbing; posterior surfaces of thighs brown with cream flecks 22 No basal webbing on foot; posterior surfaces of thighs brown with white spots; groin marbled brown and white E. surdiis 22. Snout truncate in profile; tubercles on heel and outer edge of tarsus E. duellmani Snout sloping in profile; heel and outer edge of tarsus lacking tubercles E. hamiotae 23 . Toes webbed basally (webbing enclosing basal subarticular tubercles) E. loustes Toes lacking webbing 24 24. Elongate tubercle (calcar) on heel; dorsolat- eral folds on anterior half of body E. siopelus No elongate tubercle on heel; dorsolateral folds extending length of body (if evident) 25 25. Axilla and groin bearing small white spots; adults less than 23 mm SVL E. simonbolivari Axilla and groin brown or pigmentless; adults more than 23 mm SVL 26 26. Posterior surfaces of thighs brown with cream spots E. truehae Posterior surfaces of thighs uniformly brown 27 27. Upper eyelid narrower than lOD; finger discs narrow; skin on dorsum bearing low warts .. E. gentryi Upper eyelid as wide as lOD; finger discs broad; skin of dorsum smooth with scattered pustules E. sobetes 28. Tip of Toe V extending slightly (or not at all) beyond distal border of penultimate subar- ticular tubercle of Toe IV 29 Tip of Toe V extending beyond (by length of disc) distal border of penultimate subarticular tubercle of Toe IV 32 29. Discs on fingers twice as wide as digits; no inner tarsal tubercle E.floridus Discs on fingers only slightly wider than fingers; tubercle or ridge on inner edge of tarsus 30 30. Dorsolateral folds present; small tubercle on heel E. hectiis Dorsolateral folds absent; heel lacking tu- bercles 31 3 1 . One conical tubercle on upper eyelid E. pyrrhomerus Several conical tubercles on upper eyelid .... E. leoni 32. Fleshy proboscis at tip of snout; elongate tu- bercle on upper eyelid E. appendicidatus Fleshy proboscis and conical eyelid tubercles absent 33 33. Elongate tubercle (calcar) on heel; interocular fold present E. quinquagesimus 56 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Heel lacking tubercle.s (or only a conical tu- bercle present); interocular fold absent .... 34 34. Finger discs scarcely wider than digits; white spots in a.xilla and groin E. sinionholivari Some finger discs 1.5-2.5 times digit width; lacking white spots in axilla and groin 35 35. Dorsolateral folds extending from supratym- panic fold to above groin 36 Dorsolateral folds absent 37 36. Tympanic membrane and annulus distinct; tops of digits black E. vertehndis Tympanic membrane and annulus often indis- tinct; tops of digits not black E. truebae 37. Small conical tubercles on upper eyelid and heel 38 Upper eyelid and heel lacking conical tubercles 39 38. Skin on dorsum finely granular ...£. kibiosiis Skin on dorsum smooth but bearing scattered tubercles E. tenebrionis 39. Skin on dorsum smooth; brown streaks on throat E. caprifer Skin on dorsum bearing many small tubercles; throat mottled with brown E. sulculus 43. Conical tubercle on heel; pale spots in groin and on anterior surface of thigh irregular in shape E. luteolateralis Heel lacking tubercles; pale spots in groin and on anterior surfaces of thigh ovoid E. parvillus 44. Inner edge of tarsus bearing tubercle or dis- tinct fold 45 Inner edge of tarsus lacking tubercle; fold, if present, short, indistinct, and connected to inner metatarsal tubercle 46 45. Tympanic membrane and annulus absent; no conical tubercles on upper eyelid or heel E. riiidus Tympanic membrane and annulus present; conical tubercles on upper eyelid or heel 49 46. Upper eyelid bearing three or more conical tubercles 47 Upper eyelid bearing one conical tubercle or tubercles flattened 54 47. Conical tubercles along outer edge of forearm and tarsus E. cnicifer Only small tubercles along outer edge of fore- arm and tarsus 48 40. Groin dark enclosing pale spots (yellow or orange in life); no conical tubercles on upper eyelid or heel 41 Groin lacking dark field enclosing pale spots (or, if present, conical tubercle on upper eyelid or heel) 44 41. Inner edge of tarsus bearing tubercle 42 Inner edge of tarsus lacking tubercles 43 42. Thin dorsolateral folds present; cranial crests in adults; adults > 28 mm SVL E. thymalopsoides Dorsolateral folds and cranial crests absent; adults < 25 mm SVL E. walkeri 48. Partial dorsolateral folds (region of sacrum) present; posterior surfaces of thighs brown with cream flecks E. verecundus Dorsolateral folds absent; posterior surfaces of thighs brown E. muricatus 49. Posterior surfaces of thighs uniformly brown 50 Posterior surfaces of thighs bearing cream spots (or posterior thigh mostly cream).... 51 50. Three conical tubercles on upper eyelid E. muricatus One conical tubercle on upper eyelid E. ccdcandatus ELEUTHERODACTYLUS IN WESTERN ECUADOR 57 5 1 . Conical tubercle on upper eyelid larger than heel tubercle E. rosadoi Conical tubercle on upper eyelid not larger than heel tubercle 52 52. Papilla on tip of snout; males with vocal slits; flanks and concealed thighs cream with dark markings E. subsigillatus Tip of snout lacking papilla; males lacking vocal slits; flanks and concealed thighs pre- dominantly brown 53 53. Skin on dorsum smooth with tubercles only on lower back and along stripes; males lacking nuptial pads E. laticlaviiis Skin on dorsum tuberculate or having tubercles on both anterior and lower back; males bearing nuptial pads E. latidiscus 54. Snout rounded or truncate in profile; heel lack- ing enlarged tubercle 54 Snout protruding in profile (Fig. 12F); heel bearing calcar E. colomai 55. Tips of at least some toes bearing papillae; digital pads usually cuspidate 57 Tips of toes rounded, lacking papillae; digital pads rounded or truncate 58 56. Skin on dorsum areolate; outer fingers bearing papillae (Fig. 15B) E. chalceus Skin on dorsum shagreen; fingers lacking pa- pillae E. scolodiscus 57. Small conical tubercle on upper eyelid .... 58 Tubercles on upper eyelid flattened (if present ), none conical 60 58. Posterior surfaces of thighs uniformly brown E. apicidatus Posterior surface of thighs not uniformly brown 59 59. Posterior surfaces of thighs colorless E. eremitiis Posterior surfaces of thighs (and flanks) cream with black lines E. dissimulatus 60. Tympanic annulus concealed beneath skin; toes short and broad with basal webbing E. gularis Tympanic annulus visible externally ; toes usu- ally long and slender, never basally webbed 61 6 1 . Skin on dorsum finely areolate; posterior sur- faces of thighs cream with brown flecks; digits short E. celator Skin on dorsum smooth or shagreen; posterior surfaces of thighs variable; digits long and slender 62 62. Digital pads less than twice width of digit proximal to pad 63 Digital pads large, more than twice width of digit proximal to pad 65 63. Small conical tubercles along outer edges of forearm and tarsus E. pteridophdus Forearm and tarsus lacking tubercles on outer edges 64 65. White (red in life) spots on concealed surfaces of thighs; venter with brown or gray spots ... E. cajamarcensis Concealed surfaces of thighs lacking white spots (not red in life); venter lacking distinct spots E. unistrigatiis 65. Skin on dorsum smooth E. degener Skin on dorsum shagreen 66 66. Dorsum bearing black spots and lines, includ- ing a curved stripe along flank; snout long (E- N greater than eye length) ... E. ornatissimus Dorsum lacking distinct pattern, never a pat- tern of black lines; snout short (E-N less than eye length) 67 67 . Posterior surfaces of thighs reticulate, bearing large cream spots; vertical keel at tip of snout E. phoxocephahis 58 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Posterior surfaces of thighs not bearing large 2. cream spots; snout lacking vertical keel at tip 68 68. Posterior surfaces of thighs brown with cream flecks E. nyctophylax Posterior surfaces of thighs cream with brown -\ reticulation E. eugeniae Clave de las Especies Esta clave pone enfasis en los rasgos externos independientemenle de la madurez y sexo de los especimenes. Sin embargo, para intentaridentificar Eleutherodact\lus no hay nada como conocer el 4 sexo del individuo o tener una estima de si es maduro o no. Los juveniles son siempre dificiles de identificar debido a que la membrana y el anillo timpanicos se hacen mas visibles con la edad, y muchos rasgos del patron de coloracion del vientre y superficies ocultas de los miembros se desarrollan 5 . tambien con la edad. Ademas los juveniles suelen tener la piel dorsal mas verrugosa que los adultos. Siempre que el especimen sea un adulto, creemos que la clave posibilitara una rapida y correcta 5 identificacion.Sisetratadeunejemplarpobremente preservado, dudamos que la mayoria de los usuarios puedan identificarlo correctamente usando esta clave. Dadas estas advertencias, estamos de acuerdo con la opinion de Stuart (1955:10) en relacion al uso de claves: " ... el investigador que sabe que especie tiene delante deberia tener pocas dificultades en su uso." La clave no tiene en cuenta los colores vistosos que desaparecen despues de una prolongada preservacion en alcohol. Si se tiene el individuo vivo en la mano o se conocen los colores en vida, algunas especies pueden ser identificadas en seguida 8 . porque el dorso es verde o porque hay colores vistosos en la axila, ingle o superficies ocultas de los muslos. Recomendamos fijarse en las laminas y las descripciones de coloracion en vida de la seccion "Account of Species." I . Dedo Pedial III mas largo que el Dedo V (Fig. 16B-D) 2 Dedo Pedial III mas corto que el Dedo V (Fig. 16A) 9 Dedos posteriores con palmeadura, las palmeaduras incluyen por lo menos los tuberculos subarticulares basales en todos los dedos 3 Dedos posteriores sin palmeaduras 5 La palmeadura pedial se extiende hasta los discos en algunos dedos; piel del dorso con espinulas, tuberculos o numerosos pliegues 4 Palmeadura pedial no llega a los discos en ningiin dedo; piel del dorso lisa E. longirostris Pliegue tarsal interno presente; dedos sin dis- cos E. anomalus Pliegue tarsal interno ausente; dedos con dis- cos no expandidos E. anatipes Piel del vientre areolada (granular) E. helonotus Piel del vientre lisa 6 Borde interno del tarso con tuberculo; AC/ LRC (HW/SVL) < 0.35 7 Borde interno del tarso sin tuberculos; AC/ LRC (HW/SVL) > 0.45 8 Membrana timpanica prominente; extremos digitales redondeados; Dedo Manual I mas largo que el Dedo II Barycholos piilcher Membrana timpanica ausente; extremos digitales hinchados; Dedo Manual I mas corto que el Dedo II Phyllonastes sp. Un tuberculo grande evidente en el parpado superior E. cerastes Parpado superior con tuberculos, ninguno de ellos grande E. necerus ). Piel del vientre lisa 10 Piel del vientre areolada 18 10. Extremo del Dedo Pedial V se extiende hasta la mitad de la distancia entre el ultimo (distal) ELEUTHERODACTYLUS IN WESTERN ECUADOR 59 y penultimo tuberculos subarticulares del Dedo Pedial IV (Fig. 16C); piel del dorso con tuberculos o aspera 1 1 Extremo del Dedo Pedial V no se extiende mas alia de la cuarta parte de la distancia entre el ultimo (distal) y penultimo tuberculos subarticulares del Dedo Pedial IV (Fig. 16B); piel del dorso desde lisa hasta aspera, nunca con tuberculos 13 11. Tuberculos conicos grandes en parpados superiores E. labiosus Sin uberculos grandes en parpados superiores 12 12. Marcas en foma de V invertida cafes a traves de la garganta, superficie posterior de los muslos cafe E. ocellatus Garganta de un solo color o reticulada; superficie posterior de los muslos negra con pintas crema E. cremmguis 13. Discos no expandidos, menos del doble del ancho del digito (Fig. 15 A) E.habax Algunos discos expandidos (el doble o mas del ancho del digito), especialmente en dedos externos 14 14. Superficie tarsal interna con pliegue que se extiende por lo menos hasta la mitad de la longitud del tarso 15 Borde interno del tarso puede o no tener un tuberculo, nunca un pliegue 16 15. Superficie posterior de muslos y flancos densamente punteada de negro E. actites Superficie posterior de muslos y flancos marmorea con negro y crema E. lymani 16. Cuerpo sin pliegues dorsolaterals; usualmente manchas negras en la ingle E. w-nigrum Cuerpo con pliegues dorsolaterals; ingle sin manchas negras 17 17. Tuberculo pequeiio en el talon; garganta y pecho moteado de negro E. illotus Talon sin tuberculos; garganta y pecho de inmaculados a reticulados E. achatimis 18. Extremo del Dedo Pedial V no alcanza el tuberculo subarticular distal del Dedo IV .... 19 Extremo del Dedo Pedial V alcanza el tuberculo subarticular distal del Dedo IV 40 1 9. Anillo y membrana timpanica ausente (no hay evidencia externa del oido) 20 Anillo y membrana timpanica visible externamente 28 20. Extremo del Dedo Pedial V se extiende apenas (o nada) sobre el borde distal del penultimo tuberculo subarticular del Dedo IV (Fig. 168) 21 Extremo del Dedo Pedial V se extiende mas alia (toda la longitud del disco) del borde distal del penijltimo tuberculo subarticular del Dedo IV (Fig. 16C) 23 2 1 . Por lo menos algunos tuberculos sub-articulares del pie incluidos por la palmea-dura basal; superficie posterior de los muslos cafe con puntos crema 22 Sin palmeadura basal pedial; superficie poste- rior de los muslos cafe con manchas blancas; ingle marmorea con cafe y bianco E. siirdus 22. Perfil del rostro truncado; tuberculos en el talon y borde externo del tarso E. duellmani Perfil del rostro inclinado; talon y borde externo del tarso sin tuberculos E. hamiotae 23. Dedos pediales con palmeaduras basales (la palmeadura incluye los tuberculos subarti- culares basales) E. loustes Dedos pediales sin palmeaduras 24 24. Tuberculo grande (calcar) sobre el talon; pliegues dorsolaterals en la mitad anterior del cuerpo E. siopelus 60 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Sin tuberculo elongado en el talon; pliegues dorsolaterales se extienden a lo largo del eucrpo (si son evidentes) 25 25. Axila e ingle con pequeiias manchas blancas; en adultos LRC (SVL) < 23mm E. simoiihollviiri Axila e ingle cafes o sin pigmento; en adultos LRC (SVL) > 23mm ^. 26 26. Superncie posterior de los muslos cafe con manchas crema E. truehae Superficie posterior de los muslos uniforme- mente cafe 27 27. Parpado superior mas corto que la distancia interorbital; discos digitales no expandidos; piel del dorso con verrugas poco elevadas ... E. gentryi Parpado superior tan ancho como la distancia interorbital; discos digitales expandidos; piel del dorso lisa, con espinulas dispersas E. sobetes 28. Extremo del Dedo Pedial V se extiende apenas (o nada) sobre el borde distal del peniiltimo tuberculo subarticular del Dedo IV 29 Extremo del Dedo Pedial V se extiende mas alia (toda la longitud del disco ) del borde distal del peniiltimo tuberculo subarticular del Dedo IV 32 29. Discos digitales el doble de ancho que los digitos; sin tuberculo tarsal interno E. fioridus Discos digitales solo un poco mas anchos que los digitos; tuberculo o reborde en el borde interno del tarso 30 30. Pliegue dorsolateral presente; pequeiio tuberculo sobre el talon E. hectus Pliegue dorsolateral ausente; sin tuberculos sobre el talon 31 31. Un tuberculo conico en el paipado superior E. pyrrhomerus Varios tuberculos conicos en el parpado supe- rior E. leoni 32. Proboscis carnosa en la punta del rostro; tuberculo grande en el parpado superior E. appeiullculatus Sin proboscis carnosa ni tuberculos conicos en el parpado ( ni cercanamente tan bien formados) 33 33. Tuberculo grande (calcar) sobre el takSn; pliegue interocular presente E. quinquagesimus Sin tuberculos sobre el talon (o solo un tuberculoconico presente); pliegue interocular ausente 34 34. Discos digitales escasamente mas anchos que los digitos; manchas blancas en la axila e ingle E. siimmhoUvari Algunos discos digitales de 1.5 a 2.5 veces el ancho del digito; sin manchas blancas en la axila e ingle 35 35. Pliegues dorsolaterales se extienden desde el pliegue supratimpanico hasta sobre la ingle 36 Sin pliegues dorsolaterales 37 36. Membrana y anillo timpanicos diferenciados; superficies dorsales de los digitos negras E. vertebralis Membranay anillo timpanicos frecuentemente indiferenciados; superficies dorsales de los digitos no negras E. truehae 37. Pequeiios tuberculos conicos en el parpado superior y talon 38 Parpado superior y talon sin tuberculos conicos 39 38. Piel del dorso finamente granular E. labiosus Piel del dorso lisa pero con tuberculos disperses E. tenebrionis ELEUTHERODACTYLUS IN WESTERN ECUADOR 61 39. Piel del dorso lisa; lineas cafes en la garganta E. caprifer Piel del dorso con muchos tuberculos pequenos; garganta moteada con cafe E. sidciilus 40. Ingle oscura encerrando manchas palidas (amarillas o naranjas cuando vivos); sin tuberculos conicos en el parpado superior o talon 41 Ingle sin fondo oscuro encerrando manchas palidas (o, si presente, tuberculos conicos en el parpado superior o talon) 44 41. Borde interno del tarso con tuberculo 42 Borde interno del tarso sin tuberculos 43 42. Pliegues dorsolaterales delgados presentes; crestas craneales en adultos; en adultos LRC (SVL) > 28mm E. thymalopsoides Pliegues dorsolaterales y crestas craneales ausentes; en adultos LRC (SVL) < 25 mm E. walkeri 43. Tuberculo conico sobre el talon; manchas palidas en la ingle y en la superficie anterior del muslo de forma inegular de dos E. luteolateralis Talon sin tuberculos; manchas palidas en la ingle y en la superficie anterior de los muslos ovoide E. pannllus 44. Borde interno del tarso con tuberculo o pliegue fuerte 45 Borde interno del tarso sin tuberculo; carece pliegue tarsal o, si presente, corto, indistinto, y tocando tuberculo metatarsal interno 46 45. Membrana y anillo timpanicos ausentes; sin tuberculos conicos en el parpado superior o talon E. ruidiis Membrana y anillo timpanicos presentes; con tuberculos conicos en el parpado superior o talon 49 46. Parpado superior con tres o mas tuberculos conicos 47 Parpado superior con un tuberculo conico u tuberculos aplanados 54 47. Tuberculos conicos a lo largo de los hordes externos del antebrazo y tarso E. crucifer Solo pequefios tuberculos a lo largo de los hordes externos del antebrazo y tarso 48 48. Pliegues dorsolaterales parciales (en la region del sacro); superficie posterior de los muslos cafes con puntos color crema E. verecumius Sin pliegues dorsolaterales; superficie poste- rior de los muslos cafe E. muricatus 49. Superficie posterior de los muslos uniforme- mente cafe 50 Superficie posterior de los muslos con manchas crema (o muslo posterior mayormente crema) 51 50. Parpado superior con tres tuberculos conicos E. muricatus Parpado superior con un tuberculo conico ... E. calcarulatus 5 1 . Tuberculo conico en el parpado superior mas grande que el tuberculo del talon E. rosadoi Tuberculo conico en el parpado superior no mas grande que el tuberculo del talon 52 52. Papila en la punta del rostro; machos con hendiduras vocales; flancos y partes ocultas de los muslos crema con marcas oscuras E. subsigillatus Punta del rostro sin papila; machos sin hendiduras vocales; flancos y partes ocultas de los muslos predominantemente cafes 53 53. Piel del dorso lisa con tuberculos solo en la parte baja y a lo largo de las bandas; machos sin cojinetes nupciales E. laticlavius Piel del dorso tuberculada o con tuberculos en la espalda alta y baja; machos con cojinetes nupciales E. latidiscus 62 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 54. Pertil del ro.stro redondeado o iruncado; talon sin tuberculo agrandado 55 Perfil del rostro hacia afuera (Fig. 12F); talon sin calcar E. colonial 55. Puntas de por lo menos algunos dedos posteriores con papilas; cojinetes digitales usualmente en cuspid 56 Puntas de los dedos posteriores redondeadas, sin papilas; cojinetes digitales redondeados o truncados 57 56. Piel del dorso areolada; dedos anteriores externos con papilas (Fig. 15B) E. chalceus Piel del dorso aspera; dedos anteriores sin papilas E. scolodiscus 57. Tuberculos en el parpado superior aplanados (si presentes), ninguno conico 58 Pequefios tuberculos conicos en el parpado superior 60 58. Superficie posterior de los muslos uniforme- mente cafe E. apiciilatus Superficie posterior de los muslos no uniformemente cafe 59 59. Superficie posterior de los muslos sin color E. eremitus Superficie posterior de los muslos (y flancos) crema con lineas negras E. dissimiilatiis 60. Anillo timpanico oculto bajo la piel; dedos pediales cortos y anchos con palmeaduras basales E. giilaris Anillo timpanico visible externamente; dedos pediales usualmente largos y delgados, nunca con palmeaduras basales 61 posterior de los muslos variable; digitos largos y delgados 62 62. Cojinetes digitales con menos del doble de ancho del digito proximo al cojinete 63 Cojinetes digitales grandes, mas del doble de anchos que el digito proximo al cojinete 65 63. Pequeiios tuberculos conicos a lo largo de los bordes externos del antebrazo y tarso E. pteridophilus Antebrazo y tarso sin tuberculos en los bordes externos 64 64. Manchas blancas (rojas cuando vivos) en las superficies ocultas de los muslos; vientre con manchas cafes o grises E. cajamarcensis Superficies ocultas de los muslos sin manchas blancas (no rojas cuando vivos); vientre sin manchas que lo diferencien ...E. unisthgatus 65. Piel del dorso lisa E. degener Piel del dorso aspera 66 66. Dorso con Ifneas y manchas negras, incluida una banda curva a lo largo del flanco; rostro largo (distancia ojo-narina mayor a la longitud del ojo) E. ornatissimus Dorso sin patron que lo diferencie, nunca lineas negras; rostro corto (distancia ojo-narina menor a la longitud del ojo) 67 67. Superficie posterior de los muslos reticulada, con grandes manchas crema; quilla vertical en el extremo del rostro E. phoxocephalus Superficie posterior de los muslos sin grandes manchas crema; rostro sin quilla vertical en el extremo 68 61. Piel del dorso finamente areolada; superficie 68. Superficie posterior de los muslos cafes con posterior de los muslos crema con puntos manchas crema E. nyctophylax cafes; digitos cortos E. celator Superficie posterior de los muslos crema con Piel del dorso desde lisa hasta aspera; superficie reticulacion cafe E. eugeniae ELEUTHERODACTYLUS IN WESTERN ECUADOR 63 Accounts of Species Each of the following accounts (except that of Eleuthewdactylus taeniatus, a name misapplied to specimens from western Ecuador) begins with a synonymy in which the following are given ( 1 ) original name(s) with reference to citation, type specimen(s), and type locality; (2) tirst usage of present name combination (if different from origi- nal combination): and (3) junior synonym(s), if any, with citation to work in which the name was placed in synonymy. In all accounts, except that of E. unistrigatus, a species that has been discussed at length elsewhere (Lynch, 1981a), the following sections are included. Diagnosis. — A statement of association with a species group is followed by a series of 14 num- bered statements (or sets of statements) that are modified slightly in content, but not numerical sequence, from those used by Lynch and Duellman (1980) and followed in most subsequent descrip- tions o{ Eleuthewdactylus. These statements sum- marize the taxonomically informative characters of the species; characters are defined and discussed in a foregoing section (Description of Characters). The same format is used in all accounts to facilitate comparison. A final paragraph is devoted to com- parisons with similar species. Description. — A complete description is given for all new species. In cases of previously described taxa, references to, and where appropriate com- ments on, published descriptions are provided. Coloration in life. — In addition to general state- ments, detailed descriptions have been taken from collectors' field notes. The notes on particular specimens (noted herein by present museum code and number) are referenced to the field notes by their author. Those referenced most frequently are designated by initials only; thus, JAP = James A. Peters, JDL = John D. Lynch, LAC = Luis A. Coloma, RWM = Roy M. McDiarmid, THE = Thomas H. Fritts, and WED = William E. Duellman. For those species for which a color photograph is provided, reference to the color plate is given immediately beneath the name at the beginning of the account. Natural history. — Information on habitat, hab- its, life history, and behavior are summarized from field notes and the literature. In some cases, de- tailed notes have been taken from collectors' field notes; such cases are noted in the same manner as those on coloration in life. Distribution. — The geographic and altitudinal distribution is stated with respect to physiography and bioclimatic regimes in western Ecuador. For those species occurring beyond western Ecuador, the entire geographic range is given. Etymology. — The origin, meaning, and appli- cation of the specific name is given in accounts of new species. Remarks. — Herein we provide comments on particular specimens, synonymies, relationships with other species, and any other topics not appro- priate for other sections. Eleuthewdactylus achatinus (Boulenger) Plate 2 Hylodes achatinus Boulenger, 1898:120. — Holotype: BM 1947.2.15.69, an adult female, from Cachabe, Provincia Esmeraldas, Ecuador. Hylodes pagmae Fowler, 1913:1 62. — Holotype: ANSF 18244, juvenile female, from Pagma forest in the Chanchan River Basin, Provincia Chimborazo, Ec- uador. Synonymy fide Lynch and Myers ( 1983:509). Eleutherodactyliis hrederi Dunn, 1934:1. — Holotype: AMNH 40523, adult female, from Chalichiman's Creek, Provincia Darien, Panama. Synonymy fide Lynch and Myers, 1983:509. Eleuthewdactylus achatinus — Peters, 1955:339; Lynch and Myers, 1983:509. Eleuthewdactylus pagmae — Peters, 1 955 : 350. Diagnosis. — A member of the Eleuthew- dactylus {Eleuthewdactylus) conspicillatus group having ( 1 ) skin on dorsum shagreen, that on venter smooth; discoidal fold prominent; dorsolateral folds low, granular; (2) tympanic membrane and tym- panic annulus prominent, its length -A-V?. length of eye; ( 3 ) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, about as wide as lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males with vocal slits and nuptial pads; (7) first finger longer than second; discs on outer fingers broad; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel tubercles absent; outer tarsal tubercles absent; inner tarsal tubercle small, usually present; 64 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 (11) inner metatarsal tubercle elongate, 4-6x round outer metatarsal tubercle; supernumerary tubercles at base of Toes II-IV; (12) toes bearing lateral fringes but no webbing; fifth toe slightly longer than third; (13) dorsum tan or pale brown with brown markings; venter cream with brown flecks on throat and chest; posterior surfaces of thighs brown with cream flecks or spots; (14) SVL in males 23.0-35.1 mm, in females 33.6—46.1 mm. The absence of heel tubercles and webbing, combined with the fifth toe being slightly longer than the third, distinguishes E. achatimis from all other species in western Ecuador except E. actites and E. w-nignim. Of these three species, only E. ochatiniis has dorsolateral folds. In E. achatinus, the posterior surfaces of the thighs are brown with cream flecks, whereas they are brown with black flecks in E. actites and cream with black spots or black with cream spots in E. w-nigrum. Further- more, adults off. achatimis are smaller than adults of the other species. Description. — A detailed description and in- formation on geographic variation were provided by Lynch and Myers (1983). Coloration in life. — The dorsal ground color is tan to grayish, greenish, yellowish, reddish brown with darker brown chevrons (narrowly bordered with cream or greenish-gray in some individuals) and transverse bars on the shanks. Distinct dark brown or black canthal and postorbital stripes are present, as is usually a pale (white to creamy- bronze) labial stripe. Some individuals have a pale tan to yellow middorsal stripe, usually narrowly bordered by dark brown or black. The venter is cream to white, mottled with gray on the chest and throat in some individuals, tending to become pale yellow posteriorly in some individuals; the vocal sac is pale yellow. The posterior surfaces of the thighs are various shades of brown with cream, yellow, orange, or red flecks or small spots. The iris is grayish green to usually golden bronze with black flecks and a median, horizontal red streak. Variation in series from diverse localities is evident from the following descriptions made from living or freshly killed individuals. KU } 11278-85 from Tandapi, Provincia Pichimha: Dorsum pale reddish brown to dark greenish gray with black and brown markings; usually black dashes along dorsolateral fold; flanks colored like dorsum but with some yellow-cream invasion; side of head darker than dorsum; black canthal stripe; lip white in some; belly yellow- cream; throat heavily mottled with brown or black; concealed surface of thigh reddish brown with red spots reticulated with black; red spots in groin in some individuals; iris gray with green tint, reticu- lated with black; reddish bronze anterior and pos- terior to pupil (JDL, 15 July 1967). KU 1 1 1287-96 from Tandapi, Provincia Pichincha: Posterior surfaces of thighs pale red- dish brown or fuscus-brown, usually with red spots; iris yellowish bronze to reddish bronze with bright reddish-bronze areas in front of, and behind, pupil (JDL, 17 July 1967). KU 119465 from Santo Domingo de las Colorados, Provincia Pichincha: Dorsum pale brown with only slightly darker markings edged with pale green-gray; flanks invaded with cream (hence, more yellowish); limbs like dorsum; canthal and supratympanic stripes black, former grading through brown on side of head; yellow-bronze area below eye; iris gold above, gray below, reddish- brown horizontal streak, all finely reticulated with black; posterior surface of thigh brown with creamy- yellow spots; venter white; vocal sac pale off- yellow; chin edged with white (JDL, 2 August 1968). KU 119469-72 from Santo Domingo de los Colorados, Provincia Pichincha: Dorsum rust brown, medium brown, or yellow brown; mark- ings, if present, brown; canthal and postocular stripes black, former grading to lip; lip yellow or pale cream-bronze; one individual has two pale yellow spots on dorsum; venter white; vocal sac dirty pale yellow; iris bright gold-bronze above, reddish-brown horizontal streak grading into gray- ish lower quarter, all reticulated with black; poste- rior surfaces of thighs greenish brown with dull yellow spots (JDL. 4 August 1968). KU 130365-67 from 4 km N Quevedo, Provincia Los Rfos: Dorsum pale reddish brown, pale brown to dark brown with brown to black markings edged with cream; canthal stripe dark brown to black; throat very pale cream; venter becoming yellowish posteriorly; posterior surfaces of thigh dull brown with cream flecks to dark brown with pale orange ELEUTHERODACTYLUS IN WESTERN ECUADOR 65 flecks; labial stripe cream with bronze wash; iris bright gold, finely reticulated with black and hav- ing dull red horizontal streak (JDL. 8 July 1970). KU 13 5326-38 from 4 kmNQuevedo, Provincia Los Rios: Dorsum brown with a green cast or brown marked with darker greenish-brown to brown to black; dorsal stripe pale yellow to cream edged with black; throat and chest white weakly mottled with gray; posterior venter and undersides of thighs washed with a pale yellowish-green; posterior sur- faces of thighs dull greenish-brown to brown flecked with cream to dull orange (THF, 8 July 1970). KU 165088-89 from 18 km W Pinas, Provincia El Oro: Dorsum olive-brown with pale red tu- bercles and dorsolateral folds; posterior thighs brown with tan flecks; venter cream — in larger individual mottled with black and orange suffusion in groin; iris bronze with horizontal red streak (WED. 6 March 1975). Natural history. — This species is unusual among Eleutherodactylus in northwestern South America because it is the only species commonly found in clearings. In forested areas, E. achatimis is absent or uncommon; if present, it is confined to roadcuts. Although the species apparently does not occur in deep forest, it is common in areas of "tree" crops — e.g., banana, cacao, and coffee groves. In Ecuador, the species has been found to elevations of 1 800 m, but it may have dispersed along roads to these elevations; in areas of cloud forest where roads do not exist, E. achatimis seldom is found at elevations exceeding 800 m. At night, the frogs have been found on low (<0.5 m) bushes and herbs, grass, earth banks, rocks along streams and near waterfalls, and the ground. By day, adults have been found under rocks, logs, and banana leaves, in the axils of elephant ears. One individual (KU 111336), when uncovered from beneath a rock at the edge of a stream, leaped into the stream, swam back toward the shore and took refuge under a rock in about 7 cm of water (JDL, 23 July 1967). Juveniles commonly are seen in leaf litter during the day, but their activity may be the result of disturbance. Apparently recently hatched young collected on 6 July 1976 at Rio Palenque, Provincia Los Rios, have SVLs of 6.6-6.8 mm. Throughout the range of the species in Ecuador, males have been noted calling in March, May, July, and November. The call has been described as single note or series of notes. (See Remarks.) Distribution. — Eleutherodactylus achatimis primarily is an inhabitant of lowlands from eastern Panama to southwestern Ecuador and in the Rio Cauca Valley in Colombia (Lynch and Myers, 1 983:map 1 ). The southern limit of the range on the Pacific slopes is in the vicinity of Piiias, Provincia El Oro (03°4r S) (Fig. 20). Of 109 collecting stations, 1 1 are in the dry tropical regime on the coastal lowlands south of the Equator and on the lowlands between the Cordillera de la Costa and the Andes south of 1 °S Lat. Eight stations are in the dry subtropical regime either in the Cordillera de la Costa or on the Andean slopes south of 2°S Lat. Twenty-nine stations are in the humid tropical Fig. 20. Distribution o\' Eleutherodactylus achatinus in western Ecuador with respect to bioclimatic regimes. 66 UNIV. KANSAS NAT. HIST. MUS. SPEC. PIJBL. NO. 23 regime, and 6 1 are in the humid subtropical regime. Records for the species extend from near sea level to elevations of 1800 m, but of 77 localities for which elevation is known. 34 are below 500 m, an additional 23 are below 1000 m, and only three are above 1500 m. Remarks. — A small amount of geographic variation in body size was reported by Lynch and Myers (1983). who noted that the smallest adult females are from western Colombia and northwest- ern Ecuador; these differences are more evident in the lower limits of the ranges than in the means. Females from the western slopes of the Andes in Ecuador (samples from Balsapamba, Mindo. and Tandapi) are larger than those from the lowlands by an average of 2-3 mm (about 5% larger). Adults from the southern Ecuador (Provincia El Oro) are the smallest; S VL in 47 males is 26.5-3 1 .5 mm ( .v = 28. 1 ± 0.6) and in 8 females, 36.5-4 1 .7 mm ( ,r = 38.5 ± 0.7). In every sample that we have studied, young females with weakly convoluted oviducts are only slightly smaller than adult females; for example, in the sample from Provincia El Oro. young females have SVLs of 35.9-36.0 mm, and the upper limit of juvenile females is 32.2 mm S VL. Lynch and Myers (1983) defined two color pattern morphs in this species. The ^'brederr morph is distinctive in having pale borders to the dark markings on the dorsum, whereas the dark mark- ings lack pale borders in the '"achatinus'" morph. They reported the ''bredeiT morph to be the most common in Panama, relatively abundant in the drainages of the Rio Atrato and Rio San Juan in western Colombia, and unknown south of the Rio Cajambre in Departamento Valle de Cauca, Co- lombia. They overlooked the fact that in his field notes, JDL described the dorsal markings as edged with pale green-gray in KU 119465 from Santo Domingo de los Colorados, Provincia Pichincha, and as edged with cream in KU 1 30365-67 from 4 km N of Quevedo, Provincia Los Rios, Ecuador. (See Coloration in life.) Lynch and Myers (1983) were concerned that they might have combined distinct species that differed, at least in part, by frequencies of two pattern polymorphs. Their concern was heightened because tape recordings of frogs from Panama ("libit" calls) are notably different from a recording off. achatinus from Maldonado, Provincia Carchi, Ecuador ("kree" call). However, the calls of three males (USNM 204640^2) from 2 km NE of Rio Blanco (town). Provincia Carchi, were described as "waah-waah, [a] single note repeated 5-7 times" (JAP, 26 November 1958). Other descriptions of calls of males from Ecuador are equally variable. The call of KU 117778 from Tandapi, Provincia Pichincha, was described as a series of high "quinks" (JDL, 4 March, 1968); that of KU 141749 from Santo Domingo de los Colorados, Provincia Pichincha, was described as a long, nasal "baaa" followed by "rap-ap-ap-ap" (WED, 4 July 1971 ). Until tape recordings of calls from throughout the range of the species are analyzed, these apparent differences in calls remain an unsolved mystery. Perhaps E. achatinus is a complex of cryptic spe- cies that have different calls, but that are structur- ally so similar that we have been unable to distin- guish more than one species. This species was included in the Eleiithero- dactylus conspicillatus group by Lynch (1986a). Other than its placement in the E. conspicillatus group, no explicit claims about the relationships of E. achatinus have been made. Eleuthewdactylus actites Lynch Plate 2 Eleuthewdactylus actites Lynch, 1 979b: 230. — Holotype: KU I201II, adult female, from Pilalo, 2486 m, Provincia Cotopaxi, Ecuador. Diagnosis. — A member of the Eleuthew- dactylus (Eleuthewdactylus) conspicillatus group having ( 1 ) skin on dorsum shagreen, that on venter smooth; discoidal fold present; dorsolateral folds thin; (2) tympanic membrane and tympanic annu- lus prominent, its length V^-Vi length of eye; (3) snout acuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, about as wide as lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males with vocal slits and nuptial pads; (7) first finger longer than second; discs on fingers broad; (8) fingers bearing lateral keels; (9) ulnar tubercles absent; ELEUTHERODACTYLIJS IN WESTERN ECUADOR 67 (10) heel tubercles small; outer tarsal tubercles absent; inner tarsal fold thin, present on distal -A of tarsus; (11) inner metatarsal tubercle elongate. 4- 6x subconical outer metatarsal tubercle; supernu- merary tubercles only at bases of toes; (12) toes bearing lateral keels but no webbing; fifth toe slightly longer than third; (13) dorsum gray with darker markings; venter cream with gray spots; posterior surfaces of thighs with black flecks; ( 14) SVL in males 30.0^0.0 mm, in females 48.2-64.2 mm. Eleuthewdactylus actites is most similar to E. w-nignun (and E. achatinus; see previous account). Unlike E. w-nigrum, E. actites has an inner tarsal fold, but the two species are most easily distin- guished by color pattern. In E. actites, the groin and posterior surfaces of the thighs have black flecks rather than having large black spots on a white or yellow background or large cream or yellow spots on a black background, as in E. w-nigrum. Description. — The original description by Lynch (1979b) is complete. Coloration in life. — The dorsum is gray, green- ish brown, pale reddish brown, or dark brown, usually with darker markings. Dark canthal and postorbital stripes and a pale labial stripe are present. The venter is white or yellow with brown flecks ( smaller individuals) to a heavier suffusion of brown in larger individuals. The posterior surfaces of the thighs are yellow to bluish gray with black flecks. The iris is pale yellow with black flecks and a median, horizontal red streak. Coloration of the type series (KU 120111-24): Dorsum gray, pale reddish brown to dark brown; markings darker brown edged with black; canthal and supratympanic stripes black, edged with creamy yellow-bronze; flanks paler than dorsum, usually with faint rose wash; lower flanks pale blue-gray with black spots; limbs colored as dorsum; con- cealed surfaces of thighs pale bluish gray speckled with silver and spotted with black; anal patch gray; venter white to yellow speckled with brown; under- sides of thighs deep flesh; palms, feet, and tarsi black; labial region rose-bronze with black mark- ings; iris pale yellow reticulated with black and with a red horizontal streak (JDL, 25 June 1 968). A topoty pe ( KU 1 3 1 2 1 3 ) was described as having the posterior surface of the thigh yellow, becoming orange distally, heavily marbled with black (JDL, 2 July 1970), and another series of topotypes (KU 131214-58) were described as having a pale cop- per labial stripe and the dorsum dull greenish brown medially, washed with reddish brown later- ally (JDL, 4 July 1970). Natural history. — Eleutherodactylus actites inhabits a region of cool cloud forest, but it is rarely observed within the forest. It seems to have a proclivity for exceedingly moist microhabitats. Most individuals found by day were under rocks along streams or in a moist pasture; two were in a mossy bank at the edge of a stream. All individuals active in grass by day were juveniles. Of 132 specimens with ecological data, only three were collected at night — two on broad-leafed plants (<1 m above ground) at the edge of the forest and one from low (<1 m) vegetation along a stream. Breeding occurs at least in January; many adults in breeding condition but no juveniles are in a series of 96 specimens obtained by James A. Peters in the vicinity ofthe type locality on 18- 19 January 1959. Samples (/z =116) obtained at the type locality in June 1968, July 1970, and July 1971 contain many juveniles. We interpret this seasonal distribution of juveniles as evidence that the breeding season is short and that sexual maturity is achieved in a single year, even by this relatively large species of Eleutherodactylus. Juvenile males lack vocal slits and nuptial pads. In the type series, juvenile males are 23.0-29.0 mm SVL. Females from the type series having only straight, thin oviducts are as large as 41. 1 mm SVL; females with SVLs of 43.3^6.8 mm have only slight convolutions ofthe oviducts. Distribution. — Except for two specimens from La Esperanza (below Pilalo) at an elevation of 1 500 m in the humid subtropical regime, this species is known only from elevations of 1 760-2486 m in the vicinity of Pilalo on the Pacific slope ofthe Cordil- lera Occidental in central Ecuador (Fig. 2 1 ). This is an area of cool, moist cloud forest in the humid temperate regime; Pilalo has a mean annual tem- perature of 12.6°C and an average annual rainfall of 2400 mm. We believe that a record for the species from "Latacunga"" in a much drier inter- 68 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 81° 80 ^^y^ 78° 0 20 40 60 80 100 ^^.^-^ Kilometers /" ji ^01) 111 ( V '1111(1 111 \ \ ■-— ( \®-::-:6-' ''"" -0° ( ■■■'■'■ ^''Ky'K -^^^'' " ■■ 0- -'( v-' -rx s ; / ) ^' • J? ,^°=° 1°- 2°- ^M'^ ^o • E. acthes -3° I ® E. anatipes ® £. anomahis 3°- ^^T>- ,;-,_\',< A E. babax =•/ /--^\ W' 'X-. ' '-^ ■ £. cajamarcensis 78° y 80°,, f?:., 79° Fig. 21. Distribution of five species of Eleitthew- dactylus in western Ecuador. Andean basin to the east represents an error in transcribing field notes. Remarks. — Eleiitherodactylus actites is a mem- ber of the Eleutherodactyhis conspicillatus group (Lynch, 1986a). Lynch (1979b) suggested that E. actites is most closely related to E. w-nigrum, but there is no evidence to support his hypothesis. We consider E. actites to be a geographic replacement of E. w-nigrum, which occurs at both higher and lower elevations to the north and to the south of Pilalo and, thus, has a range encompassing that of E. actites. Our characterization of E. actites as replacing E. w-nigrum in the region of Pilalo is based on their occupancy of the same (or nearly the same) microhabitat, their phenetic similarity, and our failure to find the two species in sympatry. Eleutherodactylus anatipes Lynch and Myers Plate I Eleutherodactylus anatipes Lynch and Myers. 1 983:5 1 6. Holotype: KU 177626. subadult female, from Maldonado. 1410 m. Provincia Carchi, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Craugastor) anomahis group having (1) skin on dorsum finely spiculate in males, bearing low warts in females, that on venter smooth; discoi- dal fold prominent; dorsolateral folds interrupted; (2) tympanic membrane evident, tympanic annulus obscured by spicules, small, its length '/t length of eye; (3) snout rounded in dorsal view, truncate in profile; (4) upper eyelid lacking tubercles, much wider than lOD; cranial crests absent; (5 ) vomerine odontophores triangular in outline; (6) males with vocal slits and nuptial pads; (7) first finger longer than second; discs on fingers relatively small; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) tubercles and folds on heel and tarsus absent; (11) inner metatarsal tubercle elongate, 6x indistinct outer metatarsal tubercle; supernumer- ary tubercles absent; (12) toes webbed; webbing extending to discs except only to distal subarticular tubercle on Toe IV; fifth toe shorter than third; (13) dorsum brown with darker brown blotches; venter cream; posterior surfaces of thighs yellow with brown mottling; (14) S VL in males 42.9-64.3 mm, in one female 98.6 mm. Eleutherodactylus anatipes has more toe web- bing than any other mainland member of the genus except the Colombian E. zygodactylus. Both spe- cies have the toes fully webbed except for the last one or two phalanges of Toe IV. Eleutherodactylus anomalus with slightly less webbing is easily con- fused with E. anatipes, but E. anomalus has a broader head, lacks discs on the fingers (small discs present in E. anatipes), and has an inner tarsal fold (absent in E. anatipes). Description. — Lynch and Myers (1983) de- scribed juveniles and subadults; Lynch and Ardila- Robayo (1993a) provided descriptive notes and illustrated an adult female. Coloration in life. — The colors of the holotype and paratype (KU 177625-26) were described as ELEUTHERODACTYLUS IN WESTERN ECUADOR 69 follows: Dorsum dull greenish brown with orangish warts and ridges; concealed thigh and groin dull yellow reticulated with black; throat white with brown vermiculations; venter and undersides of legs yellow; iris bright copper with black flecks and brown horizontal streak (JDL, 30 May 1977). Natural history. — This is one of the largest species of Eleutherodactylus and is one of the streamside species; others in western Colombia and Ecuador are E. anomalns, hufouifonnis, cheiwplelhus, necems, and zygodactylits. As this common name implies, these frogs never or seldom stray from the immediate vicinity of streams. Adults and juveniles sit on the ground or on rocks within 1 m of streams or in protected places beneath undercut banks of streams. In exceptionally wet areas, mule trails become streams in rainy seasons, and E. anatipes can be found sitting beside, or in, mule trails. All records for the species are for small streams and not any of the rivers that course through its range; the limited data suggest that E. anatipes avoids rivers. Distribution. — Eleutherodactylus anatipes is known from 11 localities between 100 and 1600 m in the foothills and lower cloud forests on the western slopes of the Andes in western Colombia (Departamento Valle de Cauca) and extreme north- em Ecuador (provincias Carchi and Esmeraldas). Both Ecuadorian localities are in the humid sub- tropical regime (Fig. 21). Remarks. — A single adult female is known. The holotype (KU 177626) is a juvenile female having a SVL of 73. 1 mm. In having extensive webbing between the toes, Eleutherodactylus anatipes is like E. zygodactylus, a species distributed in the lowlands of the northern part of Chocoan Colombia (Lynch and Myers, 1 983:map 2). The extensive webbing is considered to be a shared, derived character uniting these two species. Eleutherodactylus anatipes, but not E. zygodactylus, has spiculate dorsal skin — a charac- ter shared with E. anomalus and E. cheiroplethus. These four species also are unusual among mem- bers of the subgenus Craugastor in lacking obvi- ous sexual dimorphism in the size of the tympanic annulus (a plesiomorphic character) and in having exceptionally large eyes resulting in a naiTow lOD. These characters (either unpolarized or plesio- moq^hic) make the species seem similar, but they do not represent evidence that the species are closely related, although we think (perhaps irratio- nally) that the four species are close relatives, and we assign them to the E. anomalus group of the subgenus Craugastor. Eleutherodactylus anomalus (Boulenger) Plate 1 Hylodes anomalusBoulenger, 1898: 11 9. — Syntypes: BM 1 947.2. 16.8-10 from Cachabe, Provincia Esmeraldas, Ecuador. Eleutherodactylus anomalus — Peters, 1 955:339. Diagnosis. — A member of the Eleuthero- dactylus (Craugastor ) anomalus group having ( 1 ) skin on dorsum tuberculate, bearing many short ridges, that on venter smooth; discoidal fold present; dorsolateral folds absent; (2) tympanic annulus evident, tympanic annulus bearing warts, small, its length '/3 length of eye; (3) snout rounded in dorsal view and in profile; head relatively broad, HW 42- 48% SVL; (4) upper eyelid lacking tubercles, much wider than lOD; cranial crests absent; (5) vomerine odontophores broad, nearly arched; (6) males lack- ing vocal slits; nuptial pads present; (7) first finger longer than second; discs absent; (8) fingers lack- ing lateral fringes; (9) ulnar tubercles absent; (10) tubercles on heel and tarsus absent; inner tarsal fold present; (11) inner metatarsal tubercle elongate, 3x oval outer metatarsal tubercle; supernumerary tu- bercles absent; (12) toes webbed; webbing extend- ing to disc on Toe V and lateral edge of Toe III; penultimate subarticular tubercle on Toe IV absent; discs small; fifth toe shorter than third; ( 1 3) dorsum brown with indistinct darker mottling; throat brown with diffuse cream spots; otherwise venter cream; posterior surfaces of thighs cream with brown reticulation; (14) SVL in males 31.5-62.1 mm. in females 76.5-92.4 mm. Eleutherodactylus anomalus can be confused with the larger E. anatipes; both have tuberculate skin on the dorsum and extensive webbing. Unlike E. anatipes, E. anomalus lacks discs on the fingers (small discs on toes), has broader vomerine odontophores, coarser skin on the dorsum (folds and tubercles), and an inner tarsal fold. Eleuthero- 70 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 dactyhis aiionuilus also has a wider head than does E. anatipes, but these differences can be detected only by measurements. Description. — Lynch and Myers (1983) pro- vided a complete account of the species. Cochran and Coin's ( 1 970) description of a syntype is more nearly complete than Boulenger's (1898) descrip- tion, but the illustrations are better in Boulenger ( 1 898) than in Cochran and Coin (1970). Coloration in life. — Color notes on two Ecua- dorian specimens are, as follow. KU 165129 from Estacion Biologica Rio Palenqiie, Provincia Los Rios: Dorsum dull brown; flanks cream with brown reticulations; posterior thighs dark brown with yellow flecks; venter creamy white with brown reticulations on throat; iris or- ange-brown with radiating brown triangles (WED, 28 March 1975). KU 177627 from Santo Domingo de los Colorados, Provincia Pichincha: Iris pale copper with brown triangles fore and aft; pale areas on posterior thigh and in groin dull yellow (JDL, 12 June 1977). Natural history. — As might be guessed from the digital morphology, E. anomalus is strictly terrestrial. Adults and juveniles commonly are found along small, sluggish streams at night. When dis- turbed, the frogs may jump into pools and hide in the leaf litter or debris on the bottom, or they may run along the bank of the stream. Two individuals were in shaded areas on the banks of streams by day. Lynch and Myers (1983) reported breeding in February. A pair in axillary amplexus was observed in a small depression; 2 hr later the female was sitting on a clutch of 69 eggs. This is a large frog; juvenile females (ICNMHN 13204, 13206) having thin, unconvoluted oviducts are as large as 66 mm SVL. Distribution. — Eleiitherodactylus anomalus is distributed in forested lowlands below 1100 m in western Colombia and northwestern Ecuador (Lynch and Ardila-Robayo, 1 993b); of 1 4 localities of known elevation in Ecuador, all are between 1 00 and 700 m. In Ecuador, the distribution of this species is broadest near the Colombian border, and it tapers inland to the south, and thus, is absent in the drier coastal region (Fig. 21 ); six Ecuadorian localities for this species are in the humid tropical regime, and nine are in the humid subtropical regime. Remarks. — This species is most closely related to E. cheiroplethus, which is distributed at higher elevations ( 1 140-1 540 m) than E. anomalus on the western flank of the Cordillera Occidental (Lynch, 1 990). In Ecuador, but not in Colombia, E. anomalus is not found in the same streams as E. anatipes. Eleutherodactylus apicidatus Lynch and Burro wes Plate 7 Eleutherodactylus apiculatus Lynch and Burrowes, 1 990:8. Holotype: IND-AN 1 506, adult female, from the Reserva Natural La Planada, 1780 m, 7 km S of Chucunes. Municipio de Ricaurte, Departamento Nariiio, Colombia. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum smooth; that on venter areolate; well anteriad to groin; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, small, its length '/t-'/s length of eye; (3) snout subacuminate in dorsal view, subtruncate in profile; (4) upper eyelid bearing one small, nonconical tubercle, narrower than lOD; cranial crests absent; (5) vomerine odontophores oblique; (6) males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs broad; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles absent; (10) small tubercles on heel and outer edge of tarsus; inner tarsal fold absent; (11) inner metatarsal tubercle oval, 3x elongate outer metatarsal tubercle; supernumerary tubercles numerous, largest at bases of Toes II-IV; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; (13) dorsum brown with pale interorbital bar and white line above dark cloacal patch; venter pale brown; posterior sur- faces of thighs brown; (14) SVL in males 17.8- 21.8 mm, in females 21.6-26.3 mm. Eleutherodactylus apiculatus is most similar to E. calcarulatus, which differs in having a conical tubercle ( calcar ) on the heel rather than one or more small, nonconical tubercles, small ulnar tubercles, ELEUTHERODACTYLUS IN WESTERN ECUADOR 71 and an inner tarsal tubercle. Elcuthcrodactylus apicuUitus is smaller than E. calcarulatus (males 1 7.8-24.6 mm. females 25.3-28.9 mm SVL). Liv- ing individuals are easily distinguished by the color of the iris — blue-gray in E. calcarulatus and bronze in E. apiculatiis. Description. — The original description ( Lynch and Burrowes. 1990) is complete. Coloration in life. — The dorsum is tan or yel- lowish tan with or without small, irregularly shaped brown blotches; some individuals have a broad, middorsal yellow-tan stripe, and others have many fine yellow stripes on the dorsum. There is a black interorbital bar. The limbs are tan with brown bars, and the venter is white or yellow with brown flecks. A dark brown, triangular cloacal patch is bordered dorsally by a yellowish-tan line. The concealed surfaces of the limbs are brown. The iris is bronze with black reticulations and a median, horizontal red streak (Lynch and Burrowes, 1990). Natural history. — This forest-dwelling species has been studied extensively only in the cloud forest at the type locality, where Patricia A. Burrowes found it to be active every night in April-June, when gravid females were found on nearly every night. Males call from leaves and branches 1 .0-3.5 m above the ground; the call consists of two peeps. Juvenile females are 2L2-21.6 mm in SVL. The two Ecuadorian specimens were in axils of leaves of elephant ear plants by day. Distribution. — This species is known only from the type locality at an elevation of 1 780 m on the western slope of the Cordillera Occidental in south- em Colombia and in cloud forest at 1 750 and 2 1 20 m in Provincia Pichincha, Ecuador (Fig. 22); thus, it occurs in the humid subtropical and humid tem- perate regimes. We anticipate that E. apiculatus will be found to be more widely distributed in northwestern Ecuador. Remarks. — Direct evidence is lacking, but we suspect that this species may have a restricted distribution. Bothf. apiculatus ondE. calcarulatus are peculiar in that each is abundant at some locali- ties and inexplicably scarce elsewhere. Thus, we cannot estimate confidently distributions or be con- fident in inferences about the ecological interac- tions of these two species whose distributions may interdigitate. The indirect evidence is that the spe- — 1 — 81° I 80° 0 20 40 60 80 100 ''3^^ 78° - 1 Kilometers /^ 300 m ( 2000 m S 1 1 >40(IOni 1 1 I Permunenl k \ -::;''--•■•"" /" • fpi^ - 1 — -0° & -\ )aS§ / ; .o ^ -.Jo" 1°- -2° ■/ 2°- -3° "^ '^ 3°- ,^ A E. apiculatus ' % • E. appendiculatus 81° / y 80° ■ E. colomai 78° - 79° 1 Fig. 22. Distribution of three species of Eleuthero- dactylus in western Ecuador. cies thought to be related to E. apiculatus are distributed to the north and to the south of the distribution of E. apiculatus. One assumption un- derlying this indirect evidence is that allied species are distributed allopatrically. However, sampling has not been exhaustive at appropriate elevations in the Departamento Narifio, Colombia, or in provincias Carchi and Imbabura, Ecuador. Eleutherodactylus appendiculatus (Werner) Plate 4 Hylodes appendiculatus Werner, 1 894:4 1 0. — Holotype: NHM 16507, juvenile female, from "Ecuador." Eleutherodactylus appendiculatus — Peters, 1 955:350. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) devillei group having 72 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 ( 1 ) .skin on di)rsuni relatively smooth with large tubercles on eyelids, between eyes, and on limbs, that on venter coarsely areolate; discoidal fold absent; dorsolateral folds prominent; (2) tympanic membrane and tympanic annulus prominent, small, slightly more than V?. length of eye; (3) snout acuminate in dorsal view, pointed or rounded in profile, with fleshy proboscis at tip; (4) upper eyelid bearing long, conical tubercle, wider than lOD; cranial crests absent; (5) vomerine odontophores oblique, slightly elevated well behind choanae; (6) males having vocal slits; nuptial pads absent; (7) first finger much shorter than second; discs large, weakly emarginate; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles present; (10) tubercles on heel absent, on outer edge of tarsus prominent; inner tarsal fold absent; (11) inner meta- tarsal tubercle elongate, 4x elongate outer metatar- sal tubercle; supernumerary tubercles minute; (12) toes bearing lateral fringes; webbing absent; discs large, weakly emarginate; fifth toe longer than third; (13) dorsum brown with brown blotches; narrow bars present on limbs; venter cream with dark brown reticulations; throat, labial bars, and canthal stripe dark brown; posterior surfaces of thighs brown; ( 14) SVL in males 18.8-2 1 .0 mm, in females 30.0-35.0 mm. El e lithe rodactylus appendicnhitus is one of the most distinctive species in western Ecuador. The presence of a fleshy proboscis is unique; this and the presence of elongate tubercles on the upper eyelids serve to distinguish this species from all others in western Ecuador (and elsewhere). Description. — Lynch (1971a) redescribed the species based on the holotype and USNM 2047 1 2- 13. Miyata (1980a) provided additional informa- tion on adults. Coloration in life. — Dorsolateral stripes con- trast with the dorsum and flanks as noted in the following field notes. KU 165 133-38 from 5 km ESE of Chiriboga, Provincia Pichincha: — Dorsum and flanks pale green to reddish tan with reddish-brown markings; dorsolateral stripes orange-tan or pale green; fingers and toes creamy tan; discs black and cream; heels and anal region black; venter bluish gray with black mottling; iris pale bronze with median, horizontal, dull brown streak (WED, 3^ April 1975). KU 165142 from 14 km W of Chiriboga, Provincia Pichincha: — Dorsum tan with dark brown markings; venter bluish gray with black markings; iris greenish gray with black flecks and median, horizontal, orange-tan streak (WED, 8 May 1975). Natural history. — Adults and juveniles have been found on low vegetation (<1 m) in dense forest at night; some were near streams. Miyata (1980a) reported individuals along roadcuts on rainy nights. Juveniles are active on the ground in wet forest by day (Lynch and Burrowes, 1989; WED, pers. obs.). Distribution. — The species is known from the Reserva La Planada in extreme southern Colombia and 12 localities in Ecuador (Fig. 22). All localities are in cloud forest at elevations of 1 460-2800 m on the western slopes of the Cordillera Occidental; six sites are in the humid subtropical regime, and six are in the humid temperate regime. Remarks. — Females having no convolutions of the oviducts are as large as 27.3 mm SVL (MCZ 98030); KU 177637 (27.9 mm SVL) has thin oviducts with only slight convolutions. Two males from Provincia Pichincha (MCZ 92885, 94818) have white testes, whereas another male (KU 165140) has dark flecks on the mesorchium. One mature male having a SVL of 16.2 mm (QCAZ 7968) lacks a fleshy proboscis; there is no apparent trace of injury. In adult males, the vocal slits are short and located posterolateral to the tongue. Al- though we assign E. appendiculatus to the E. devil lei group, we are not confident in doing so. Its nearest relatives are undescribed species from cloud for- ests in western Colombia. Eleuthewdactylus babax Lynch Plate 3 Eleutherodactylus haha.x Lynch, 1989a: 10. Holotype: ICNMHN 1 3592. adult male, from Finca La Planada, 1 970 m, 5 km by road from Chucunes, Municipio de Ricaurte, Departamento Narifio. Colombia. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) dolops group having (1) skin on dorsum bearing low warts, that on venter smooth; discoidal fold present; dorsolateral folds incomplete; (2) tympanic membrane andtym- ELEUTHEKODACIYLUS IN WESTERN ECUADOR 73 panic aniuiliis prominent, small, its length V4-A length of eye; (3) snout subacuminate in dorsal view, rounded to truncate in prohle; (4) upper eyelid lacking tubercles, wider than lOD; cranial crests absent: (5) vomerine odontophores triangu- lar in outline; (6) males having vocal slits; nuptial pads absent; (7) first finger longer than second; discs narrow; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles and folds; (11) inner metatarsal tubercle oval, 3x round outer metatarsal tubercle; supernu- merary tubercles only at bases of toes; (12) toes lacking lateral fringes and webbing; fifth toe slightly longer than third; (13) dorsum brown with darker brown blotches; ventercream; posterior surfaces of thighs brown with cream flecks; (14) SVL in one male 42.4 mm. in females 45.6^8.7 mm. By virtue of having long, slender digits with narrow digital discs, E. babax is distinguished immediately from all other Eleuthewdactylus in western Ecuador. The similar E. dolops from the eastern slopes of the Andes is larger. The smaller Barycholos pidcher superficially resembles E. babax but has an inner tarsal tubercle and a short fifth toe. Description. — Lynch (1989a) described this species on the basis of limited material from three localities in Colombia. Additional material has not amplified the described variation. Coloration in life. — No color notes are avail- able for the two specimens known from Ecuador; the following descriptions are based on specimens from Colombia. La Planada, Departamento Narino: Dorsum yellowish olive to reddish brown with dark brown markings (commonly outlined with cream); flanks and concealed surfaces of hind limbs dull orange to salmon; flanks with brown spots; posterior sur- faces of thighs with brown reticulations; throat and chest brown; belly and undersides of thighs salmon; iris bronze above and black below brown horizon- tal streak (P. A. Burrowes, April-June, 1986). Boqueron, Vereda Las Amardlas, Mimicipio El Cairo, Departamento Valle del Cauca: Dorsum brown with darker brown markings edged with cream; groin and posterior surfaces of thighs brown with cream flecks; throat brown with cream flecks; belly dull pale yellow with some gray flecks; un- dersides of limbs with orange cast; upper iris heavily flecked with pale olive; lower iris brownish gray (JDL 18356-57. 25 June 1991). Natural history. — Because it is terrestrial and nocturnal, this species is easily overlooked. Indi- viduals have been found in dense cloud forests as well as in clearings. It is not restricted to the margins of streams, but individuals at La Planada, Departamento Nariiio, Colombia, and nearTandapi, ProvinciaPichincha, Ecuador, were collected along streams. Distribution. — Eleutherodactylus babax occurs in cloud forests at elevations of 1500-2200 m on the Pacific slopes of the Cordillera Occidental from Departamento Antioquia, Colombia, to Provincia Pichincha, Ecuador. The two Ecuadorian localities are at elevations of 1550 and 1750 m in the humid temperate regime (Fig. 21). Remarks. — Eleutherodactylus babax was placed in the Eleutherodactylus dolops group by Lynch (1989a), who recognized two species in the group — £'.Z7fl/?fljr on the Pacific slopes of the Andes and E. dolops on the Amazonian slopes. Eleutherodactylus cajamarcensis Barbour and Noble Plate 5 Eleutherodactylus cajamarcensis Barbour and Noble, 1920:404. Holotype: MCZ 5407. adult male, from Ruins near Huambos, Departamento Cajamarca. Peru. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) unistrigatus group having (1) skin on dorsum shagreen, bearing ill- defined rows of pustules, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, round, its length V^-Vz length of eye; (3) snout rounded in dorsal view and in profile, canthus rostralis angular; (4) upper eyelid bearing small tubercles, slightly narrower than lOD; cranial crests absent; (5) vomerine odontophores oblique, not prominent; (6) males having vocal slits and nuptial pads; (7) first finger shorter than second; discs small; (8) fingers lacking lateral fringes; (9) ulnar tubercles indistinct or absent; (10) heel and outer edge of tarsus lacking tubercles; inner edge of 74 UNTV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 tarsus usually bearing one tubercle; (II) inner metatarsal tubercle oval. 4-6x oval outer metatar- sal tubercle; supernumerary tubercles numerous; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; di.scs as large as those on fingers; (13) dorsum gray to pale brown with darker brown markings; venter cream with brown or gray spots; posterior surfaces of thighs, groin, and lower flanks black with white spots; (14) SVL in males 19.2-24.1 mm, in females 27. 1-33.8 mm. Eleutherodactylus cajamarcensis clearly is a member of the E. unistrigatus group, but does not seem to be closely allied to any of the other species in western Ecuador. Lynch ( 1 969) compared it with E. unistrigatus and a species that later was named E. modipeplus (Lynch. 1979a). In habitus and mor- phology of the digits, it is most like E. petwbardus, versicolor, and wiensi, but it differs from these by having white spots in fields of red flash colors on the flanks and posterior surfaces of the thighs. The digit discs are slightly nanower than those in E. petrohardus, versicolor, and wiensi. Description. — The redescription by Lynch (1969) is adequate. Coloration in life. — Detailed notes on colora- tion are not available for specimens from western Ecuador; the following descriptions are based on specimens from southern Ecuador. KU 141896-909 from 5 km NE Cariamanga, Provincia Loja: Dorsum gray, tan, dark brown, or reddish brown; venter pale gray; spots in groin and on posterior surfaces of thighs tomato-red; iris bronze with reddish-brown median, horizontal streak (WED, 18 July 1971). KU 165 190-92 from 12.5 km S Loja, Provincia Loja: Dorsum tan to reddish brown with dark brown markings; groin and anterior surfaces of thighs rose; posterior surfaces of thighs black with white flecks or mottling; iris pale bronze with reddish-brown, median, horizontal streak (WED, 9 March 1975). Natural history. — Although E. cajamarcensis sometimes is found on low vegetation at night, most individuals have been found by day; many were under rocks in subparamo, others were in either terrestrial or arboreal bromeliads in forests; as many as 1 1 individuals have been obtained from a single bromeliad. Distribution. — This species is widely distrib- uted on the Pacific slopes in the region of the Huancabamba Depression in northern Peru and the Loja Basin in southern Ecuador. It occurs in tropi- cal dry forest, cloud forest, and subparamo at elevations of 1800-3100 m. The specimens re- ported here are from Luz Maria (02°4 1 ' S ), Provincia Azuay, the northernmost known locality for the species (Fig. 2 1 ). This site is at an elevation of 1 880 m in the humid temperate regime. Remarks. — The presence of E. cajamarcensis in western Ecuador is somewhat surprising. Luz Maria, Provincia Azuay, is the northernmost known locality for the species, which, together with its apparent allies (E. petrobardus, versicolor, and wiensi). are distributed to the south. These four species (and E. lymani) seem to be elements of a Huancabamba Depression fauna (Duellman and Wild. 1993). some of whose members encroach into the area here termed western Ecuador. Eleutherodactylus calcarukitus Lynch Plate 6 Eleutherodactylus calcarukitus Lynch, 1976a:6. — Ho- lotype:KU 1 11218.adultfemale,fromTandapi, 1460 m, Provincia Pichincha. Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum smooth, that on venter areolate; discoidal fold prominent; dt^rsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, round, small, its length '/i-'/? length of eye; (3) snout subacuminate in dorsal view, rounded in prolile, with papilla at tip; (4) upper eyelid bearing one or two tubercles, slightly wider than lOD; cranial crests absent; (5 ) vomerine odontophores oblique; (6) males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs dilated; (8) fingers bearing lateral fringes; (9) ulnar tubercles small, conical; ( 10) heel bearing elongate, conical tubercle; inner tarsal tu- bercle present; (11) inner metatarsal tubercle oval, 4-6x oval outer metatarsal tubercle; supernumer- ary tubercles few; (12) toes bearing narrow lateral fringes; webbing absent; fifth toe much longer than ELEUTHERODACTYLUS IN WHSTHRN ECUADOR 75 third; (13) dorsum pale brown with darker brown markings; venter cream with brown flecks; poste- rior surfaces of thighs uniform brown; ( 14) SVL in males 17.8-26.6 mm, in females 25.3-28.9 mm. Eleutherodactyliis calcarulatus is most easily recognized because it is relatively small and has smooth skin on the dorsum and prominent tu- bercles on the upper eyelid and on the heel. Addi- tionally, if living material is available, the blue- gray iris is distinctive. Several other species have prominent tubercles on the upper eyelids and heels, but among those in western Ecuador, E. appendi- culatus, crucifer, latidiscus, and muricatus have tuberculate skin on the dorsum, or the tubercles are small, as in E. eremitus, laticlovius, and nyctophylax: moreover, E. appendiculatus, labiosus, and tenebrionis have shorter fifth toes. The smaller E. scolodiscus has a bright blue iris, but it has papillae on the tips of some of the digits. Description. — The original description by Lynch ( 1976a) is complete. Coloration in life. — The dorsal coloration is highly variable, ranging from cream with a green tint to dark brown or nearly black; the dorsal markings usually are diffuse and barely distin- guishable in some individuals, as noted in the following descriptions. KU 1 11216-44 from Tandapi, Provincia Pichincha: Dorsum cream with green tint, yellow- ish brown, reddish brown, or dark brown to nearly black; dorsal markings and limb bars same color as ground color but darker, diffuse in some individu- als. Some have a reddish-brown dorsum with brown stripes. Venter dark creamy yellow to brown or grayish black; groin and concealed surfaces of thighs gray to blackish brown. Cream or yellow spots evident on labial border in some individuals. Iris blue-gray, usually with a reddish-bronze, me- dian, horizontal streak (JDL, 15-23 July 1967). KU 165 189 from 3.5 km NE Mindo, Provincia, Pichincha: Dorsum dull olive-green with dark brown markings; venter dull greenish yellow with gray suffusion; iris bluish gray with dull red streak (WED, 8 April 1975). Natural history. — Eleiitherodactyhts calcaru- latus was observed every night during JDL's visits to Tandapi, Provincia Pichincha, in 1967, 1968, 1970, and 1977. The species is arboreal and espe- cially abundant along trails through cloud forest, particularly in the proximity of small streams. In spite of its abundance at Tandapi, no calls were associated with this species. However, at La Delicia, Provincia Imbabura, on 16 January 1978, males were calling at night from high perches (2.4-3.0 m above the ground) in deep forest and at the edge of the forest; the call is an explosive, musical "tink." By day, individuals have been found in arboreal bromeliads, in the axils of elephant ear plants, under bark on logs, and in rotting fence posts. Distribution. — This species occurs in cloud forests at moderate elevations (1140-2700 m) on the western flank of the Cordillera Occidental in Ecuador (Fig. 23); 10 localities are in the humid tropical regime and nine are in the humid temperate regime. The presence of the species on the slopes southeast of Maldonado, Provincia Carchi, almost assures that it will be discovered in adjacent Departamento Nariiio, Colombia. Remarks. — The largest juvenile female (thin, straight oviducts) is 25.0 mm SVL. One specimen (MCZ 98076) from the Rio Corazon, 6.3 km E of Tandapi, Provincia Pichincha, is an adult female having a SVL of 32.4 mm; this specimen is 3.5 mm larger (SVL) than any other female examined and is tentatively referred to E. calcarulatus. 81 80 0 20 40 60 80 100 Fig. 23. Distribution of two species of Eleuthero- dactyliis in western Ecuador. 76 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Eleutlierodcictylus caprifer Lynch Plate 2 EleittlicroJactyliiscdpriferLynch 1977:282. — Holotype: KU 131589, adult male, from Las Palma.s (= La Palma). 920 m, Provincia Pichincha. Ecuador. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) conspicillatus group having ( 1 ) skin on dorsum smooth, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, round, small, its length Va-A length of eye; (3) snout short, subacuminate in dorsal view, truncate in profile; (4) upper eyelid lacking tubercles, slightly wider than lOD; cranial crests absent; (5) vomerine odontophores triangu- lar in outline; (6) males having vocal slits; nuptial pads absent; (7) first finger longer than second; discs broad; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles; (11) inner metatarsal tu- bercle elongate, 4x elongate outer metatarsal tu- bercle; supernumerary tubercles absent; (12) toes bearing narrow lateral keels; webbing absent; fifth toe slightly longer than third; ( 1 3) dorsum tan with numerous thin brown chevrons and broad brown dorsolateral stripe; venter white with brown stripes on throat; posterior surfaces of thighs brown with cream flecks; ( 14) SVL in males 21.0-30.4 mm, in females 40.5^3.8 mm. Like other species in the Eleutherodactylus conspicillatus group, E. caprifer has the first finger longer than the second and the fifth toe only slightly longer than the third. The color dorsal pattern off. caprifer resembles some E. achatinus {'"brederi'" morph) but is readily distinguished by the presence of areolate skin on the venter and short, brown, longitudinal stripes on the throat in females, and by the absence of dorsolateral folds. Description. — Lynch (1977) and Lynch and Myers (1983) provided thorough descriptions. Coloration in life. — The following description is of the type series, which illustrates the sexual dimorphism in color. KV 13 1589-602 from La Palma, Provincia Pichincha: In males, dorsum yellow (with green tint in some individuals) with brown tint on snout; concealed surfaces of thighs yellow; throat flesh- colored. In females, dorsum reddish brown (pale orange in some individuals), with paler, nearly yellowish-brown flanks; groin and concealed sur- faces of thighs dull brown with pale green flecks on latter; throat white with pair of dull gray longitudi- nal stripes. In both sexes, naiTow brown or black chevrons and dorsolateral stripes on dorsum and transverse bars on limbs; face yellow fading to reddish brown; venter (except throat) uniform off- white; iris bright copper, darkest below, with dark brown, median, horizontal streak (JDL, 8 August 1977). Natural history. — At the type locality, Eleutherodactylus caprifer is reasonably abundant in the spray zone of a small waterfall, where indi- viduals have been found on low vegetation (up to 20 cm above the ground). Males have been noted calling in July and August; the call is a series of 8- 10 high, piercing chirps. Distribution. — This species is distributed in lowlands (50-950 m) in southwestern Colombia (Departamento Valle de Cauca southward) and northwestern Ecuador (Fig. 24). Four of the five Ecuadorian localities are in the humid subtropical regime, whereas one is in the humid tropical re- gime. Remarks. — As noted by Lynch and Myers (1983), this species is seldom collected; perhaps this reflects some microhabitat preference not ap- parent to collectors. On various occasions, JDL has found two or three individuals in an hour or so at one site, but there is nothing obvious about why the species is abundant at the type locality. Eleutherodactylus celator Lynch Plate 7 Eleutherodactylus celator Lynch 1 976a:22. — Holotype: KU 13 1573. adult female, from La Delicia, 2700 m, Provincia Imbabura, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having (1) skin on dorsum finely areolate, that on venter coarsely areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, round, its length Vt'-A length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid PLATE 1 ^^'"^h^ E. anatipes. IND-AN (PAB 568). jii\enile. La Planada. Narino. E. anomalus, KU 163130. Icnialc. .5S.4 mm S\ 1.. Km PalcnqL Colombia. PAB. Pichincha, Ecuador. WED. iE:. «^ren(.v, KU 179077, female, 72.3 mm SVL.Maldonado. Car- E. loii;^ir(>stns. iC.N.Mll.N 13215. temalc, 49.6 mm SVL. Rio chi, Ecuador. JDL. Calima. Vaile de Cauca. Colombia. JDL. E. omatissimiis. USNM 2J mm S\ L. Tandapi, Pichincha. Ecuador. RWM. Pichincha. Ecuador. JDL. £. m///7(:flr!«,MCZ 94456. female. 33.8 mm SVL. Rio Palenque. E. mctoplnlcLx. KU 110954, male. 24.8 mm SVL. Tandapi. Pichincha. Ecuador. KM. Pichincha, Ecuador. JDL. E. subsigillatiis, MCZ 97559. male, 29.2 mm SVL. Centinela, E. pteiuloplulus. USNM 286043, male. 24.7 mm SVL. 13.1 km Pichincha, Ecuador. KM. NW Nono, Pichincha, Ecuador. RWM. £. eiigenlae. KU 165899, female, 30.5 mm SVL. Quebrada Za- E. tbymalopsoides. KU 177863, female. 44.9 mm SVL. Pilalo, padores, Pichincha, Ecuador. WED. Cotopaxi, Ecuador. JDL. PLATE 7 E. Itacolaicralis. KU 120154, female. 23.5 mm SVL. Tandapi. E. walkcri. KU 17S()I(). Icmale. 21.9 mm SVL. Santo Domingo Pichincha. Ecuador. JDL. de los Colorados, Pichiiicha. liciiador. .IDL. E. wcilkeh. KU 142017, female, 19.5 mmSVL. Rio Baba, Pichin- E apu iilaiiiy IND-AN 1755, female, 21.6 mm SVL. La Plana- cha, Ecuador. WED. da, Narino, Ecuador. PAB. E. verecimclii.s, ECO 94, female, 2 1 .4 mm SVL. El Cri.stal, E.s- £. panilliis. KU 1 20244, female. 24. 1 mm SVL. Tandapi, Pichin- mcraldas, Ecuador. MLC. cha, Ecuador. JDL. E. wsadoi, KU 2 1 805 1 , female, 25. 1 mm SVL. Rio Canoi, Pichin- E. celaloi: KU 1 65200, male, 20.5 mm SVL. 9.5 km NW Nono, cha, Ecuador. LAC. Pichincha, Ecuador. WED. PLATE 8 / /■/, , /;m, I CO I cu.kIoi mi C, iSVL. ElCristal. Imikt.iM.is E. Icnni. KU 177323. female. 22.3 mm .SVL. 14 km .SE Mal- donado. Carchi. Ecuador. JDL. £./n/77i(W)fn/.s, KU 131607. female. 20.8 mm SVL. Pilalo.Coto- / siiiKnilxilnuii kL 2 1N2^ V male. I'S 7 mm S\ L C aslKa To paxi, Ecuador. JDL. toias, Boli\ai Eeiiadoi I AC E. chalceiis. KU 200152. female. 30.5 mm SVL. La Planada. E. clialceiis. ICNMHN 19186. female, 33.7 mm SVL. Paique Narino, Colombia. WED. Nacional Las Orqui'deas. Antioquia. Colombia. JDL. E. giilaris, ICNMHN 13739. female, 22.6 mm SVL. Isla Gorgo- E. scnidisvus. IND-AN 1778, female. 22.3 mm SVL. La Plana- na, Cauca, Colombia. JDL. da. Narino, Colombia. PAB. ELEUTHERODACTYLUS IN WESTERN ECUADOR 77 lacking tubercles, much narrower than flat lOD; cranial crests absent; (5) vomerine odontophores prominent, triangular in outline; (6) males lacking vocal slits and nuptial pads; (7) first finger shorter than second; fingers short; discs large; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles; (11) inner metatarsal tubercle elongate, 5-6x subconical outer metatarsal tubercle; supernumerary tubercles numerous; ( 1 2) toes bearing narrow lateral fringes; webbing absent; discs as large as those on fingers; fifth toe much longer than third; (13) dorsum brown with darker brown markings; venter cream with minute brown flecks; posterior surfaces of thighs cream with minute brown flecks; (14) SVL in males 19.6-21.4 mm, in females 22.0-24.5 mm. Eleutherodactylus ce/arorresembles E. chalceus in having areolate skin on the dorsum and short digits, but E. chalceus has elongate papillae on Fingers II-IV (absent in E. celator), bifid distal subarticular tubercles on fingers and toes (round in E. celator), and brown mesorchia (white in E. celator). Description. — Lynch's (1976a) original de- scription is adequate. Coloration in life. — Coloration in this small species is highly variable, as noted in the following descriptions. KU 131573-86 from La Delicia, Provincia Imbabura: Dorsum reddish brown, medium brown, or tan with brown, reddish-brown, or black mark- ings (enamel-white flecks in a few individuals; one individual mostly pale grayish cream with soft gray markings; some individuals with pale gray to cream spots bordered by black); dorsolateral stripes black enclosing fawn-yellow to pale rust areas; con- cealed surfaces of limbs colorless or pale orange; venter yellow, dirty cream, or pale gray when collected (later changed to black); throat of males more lemon-yellow than belly; iris deep brown with median, horizontal, dark brown streak ( JDL, 5 August 1970). KU 165200-01 from 9.5kmNWNono, Provincia Pichincha: Dorsum yellowish tan to pale orange; flanks tan; venter pale yellow; iris copper with median, horizontal brown streak (WED. 1 1 April 1975). KU 177683-725 from 14 km SE Maldonado, Provincia Carchi: Dorsum yellow, pale orange. reddish brown, or brown with or without brown marbling; limbs tending to be more orange than body; venter creamy metallic-yellow to nearly black, in which case dark cream flecks obvious; throat dirty dull yellow; limbs gray; iris bright copper with black reticulations and brown, median, hori- zontal streak (JDL, 31 May 1977). Natural History. — All Ecuadorian specimens have been found in terrestrial bromeliads in cloud forest by day; usually only one individual was found in a given bromeliad, but one large brome- liad contained eight individuals. Arboreal bromeli- ads at a site 14 km SE of Maldonado, Provincia Carchi, were searched, but no individuals were found, whereas terrestrial bromeliads yielded 43 individuals. Lynch and Burrowes (1990) reported specimens from terrestrial bromeliads in Colom- bia, but they also noted individuals on low vegeta- tion not associated with bromeliads at night. Distribution. — Eleutherodactylus celator is known from only six localities at elevations of 1750-2800 m on the western slopes of the Cordil- lera Occidental in northwestern Ecuador and adja- cent Colombia (Fig. 24). The five Ecuadorian lo- calities are in the humid temperate regime. Remarks. — The distribution of this small frog is poorly documented because of inadequate searches of terrestrial bromeliads in cloud forest. -0 81 0 20 40 60 80 100 Kilometers 81 • E. c ap lifer A E. celator ■ E. dissimulatiis O E. gentry'i Fig. 24. Distribution of four species oi Eleuthero- dactylus in western Ecuador. 78 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Eleutherodactylus cerastes Lynch Plate 3 Elciitherodactyliis cerastes Lynch, I975a:25. — Holo- type: USNM 195785, juvenile female, from Palma Real. Provincia Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) sulcatus group hav- ing (1) skin on dorsum bearing heterogeneous tubercles and longitudinal ridges, that on venter smooth; discoidal fold absent; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, its length '/^-V? length of eye; (3) snout rounded in dorsal view, truncate in profile; (4) upper eyelid bearing many conical tubercles (one elongate), broader than lOD; cranial crests present; (5) vomerine odontophores arched, broad; (6) males lacking vocal slits and nuptial pads; (7) first finger longer than second; discs barely devel- oped; (8) fingers lacking lateral fringes; (9) ulnar tubercles in a distinct row; (10) heel bearing one small tubercle; outer surface of tarsus having small tubercles; inner tarsal fold and tubercles absent; (11) inner metatarsal tubercle compressed. 3x low, round outer metatarsal tubercle; supernumerary tubercles absent; (12) toes lacking lateral fringes; webbing absent; discs small, but larger than those on fingers; fifth toe shorter than third; (13) dorsum brown with darker markings associated with tu- bercles and ridges; venter cream with brown reticu- lation; throat brown with cream flecks; posterior surfaces of thighs brown with cream spots; iris green in life; ( 14) S VL in two males 28.3-33.4 mm, in nine females 44.4-55.8 mm. West of the Andes, this is the only species of broad-headed Eleutherodactylus in which there is an elongate tubercle on the upper eyelid. Superfi- cially, it resembles E. ruizi, which has granular skin on the venter and an inner tarsal fold. Eleuthero- dactylus helonotus also has granular skin on the venter, but it lacks elongate tubercles on the upper eyelids and has low, flat warts on the dorsum (but only large females are known). Like E. cerastes, E. cadenoi has smooth skin on the venter and an elongate eyelid tubercle, and lacks an inner tarsal fold, but it is easily distinguished by having an ossified cheek region. The absence of webbing on the feet readily distinguishes E. cerastes from the coarsely tuberculate E. anomalus. Eleuthero- dactylus necerus (a large member of the subgenus Craugastor) is similar in skin texture and lacks webbing on the feet, but it lacks conical cubercles on the eyelids. Additionally, E. cerastes is distin- guished from other members of the genus in west- ern Ecuador by having a lime-green iris in life. Description. — Lynch (1975a) based the origi- nal description primarily on specimens from west- ern Ecuador. Coloration in life. — Color notes are not avail- able for Ecuadorian specimens; the following notes are based on specimens from Colombia. Parque Nacional Natural Las Orquideas, Vereda Calles, Municipio Urrao, DepartamentoAntioquia: Dorsum brown to rusty brown with or without black highlights, dark green patch, or orange tint on ridges; posterior surfaces of thigh black with cream flecks; venter cream with brown reticulation; pale yellow line on lips of one individual; iris pale green with brown or black radiations (JDL 16359-62, 23-25 May 1988). Vicinity of Boqueron, Departamento Choco: Dorsum brown to dark brown with black markings beside ridges or ridges rust colored; labial bars black and cream or dark brown and reddish brown; throat brown or dull brownish yellow with brown marbling; posterior surfaces of thighs dark brown with cream flecks; iris lime-green with dark gray radiations or black reticulation (JDL 18539 and 18686, 24-25 June 1991). Natural history. — This species inhabits cloud forests and clearings therein. Occasionally, indi- viduals of this terrestrial, primarily nocturnal frog are found in leaf litter by day, but it is assumed that these were disturbed and fled their diurnal retreats. At night, the frogs have been observed sitting in an alert posture on upper surfaces of leaves, walking across trails, and climbing banks of streams. Distribution. — Ecuadorian records for the spe- cies are at elevations of 500-1200 m in the humid subtropical regime on the western face of the Andes (Fig. 23), but in Colombia, where the species is better known, it extends to about 2000 m. Present records suggest a continuous distribution from northwestern DepartamentoAntioquia, Colombia, to the Ri'o Guayllabamba in Provincia Pichincha, Ecuador. ELEUTHERODACTYLUS IN WESTERN ECUADOR 79 Remarks. — This species was included in the Eleuthewdactylus sulcatus group as redefined by Lynch (1986a). Eleuthewdactylus sernai Rivero from the extreme northern part of the Cordillera Occidental in Colombia probably is not distinct from E. cerastes, but that issue will be addressed in a monograph of the frogs of the E. sulcatus group by JDL. At present, the relationships of several species in the E. sulcatus group, including E. cerastes and E. helonotus (the only members of the group in western Ecuador), are not resolved. Eleutherodactylus chalceus (Peters) Plate 8 PhyUohateschalceusV^iQK. 1 874:609. — Syntypes: Lost, from "Pastasasathal" (= Pastassa-Tal, Colombia, tide Nieden, 1923:402). Syrrhophus chalceus — Boulenger, 1888:206. Syrrhophiis oreolatus Boulenger, 1 898: 1 22. — Syntypes: BM 1947.2.15.38-39 from Cachabe, Provincia Esmeraldas, and "Chimbo" (= Puente del Chimbo), Provincia Guayas, Ecuador. Synonymy fide Lynch, 1980b: 179. Eleutherodactylus areolatus — Lynch, 1968:291. Eleutherodactylus chalceus— L^nch 1968:291; 1980b: 179. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) diastema group hav- ing ( 1 ) skin on dorsum and venter areolate; discoi- dal fold prominent; dorsolateral folds absent; (2) tympanic membrane absent but tympanic annulus evident beneath skin, its length about Vi length of eye; (3) snout short, subacuminate in dorsal view, truncate in profile; (4) upper eyelid lacking tu- bercles, as wide as lOD; cranial crests absent; (5) vomerine odontophores oblique; (6) males having vocal slits and large, external vocal sac; nuptial pads absent; (7) first finger slightly shorter than second; discs round, with papillae at tips of discs on Fingers II-IV; (8) fingers broad, bearing narrow lateral fringes; distal subarticular tubercles bifid; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles and folds; (11) inner metatarsal tubercle oval, 3x flat, round outer metatarsal tu- bercle; supernumerary tubercles not obvious; ( 12) toes lacking lateral fringes; webbing absent; discs round, bearing distal papillae of Toes II-IV; distal subarticular tubercles weakly bifid; fifth toe much longer than third; ( 1 3) dorsum cream to brown with or without reddish brown spots or reticulations; venter white; posterior surfaces of thighs unpigmented; (14) S VL in males 1 7.5-26.9 mm, in females 27.7-31.2 mm. Eleutherodactylus chalceus is distinctive in the genus by having areolate skin on the dorsum. Also, the presence of elongate papillae on the tips of the digits and bifid distal subarticular tubercles readily distinguish this species from all others. Eleuthero- dactylus scolodiscus is similar, but it has short (not elongate) papillae only on Finger III and Toes II- IV, round instead of bifid distal subarticular tu- bercles, less prominent granulations on the dor- sum, orange in the groin and on the concealed surfaces of the thighs, and a blue iris in life. Description. — Lynch's ( 1 97 1 ) redescription (as E. areolatus) is complete and accurate, except that he did not distinguish adequately between adults and subadults and juveniles. The size range give here is based only on calling males and ovigerous females. Coloration in life. — Many Eleutherodactylus chalceus have no color pattern; others have red- dish-brown flecks on the dorsum (most common in Departamento Valle de Cauca, Colombia), bold reddish-brown spots, or reddish-brown reticula- tions. The variation and ability to change color are documented by the following field notes: KU ] 17487-91 from Santo Domingo de los Colorados, Provincia Pichincha: At night, dorsum uniform creamy yellow to pale reddish-bronze or pale reddish-brown; by day, some individuals creamy yellow with reddish-brown spots; flanks pale yellow; hands and feet of yellow phase, or- ange, or of reddish phase, yellow; colors well set off from color of shank and forearm; venter uni- form white, but throat may have yellowish tint; posterior surfaces of thighs unpigmented (JDL. 5 March 1968). KU ]] 9474-85 from Santo Domingo de los Colorados, Provincia Pichincha: Dorsum reddish brown at night; by day creamy yellow with or without reddish-brown spots; venter white; hands and feet yellow or orange (JDL, 2 August 1968). KU J 65 J 43^8 from 3.5 km NEMindo. Provincia Pichincha: Dorsum pale greenish tan with reddish- 80 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 brown spots; thighs creamy white; venter white; palpebrum bronze; iris black with faint, dull bronze reticulations (WED, 7 April 1975). One individual (KU 1 4 1 773 ) from the Rfo Baba, Provincia Pichincha, and two (KU 141774-75) from 10.5 km N Quininde. Provincia Esmeraldas, were noted to have a metallic green dorsum (WED, 2and4July 1971). Another individual (KU 120252) from Santo Domingo de los Colorados, Provincia Pichincha. was noted to have the groin and con- cealed surfaces of the thighs red (JDL, 3 August 1968). Natural history. — During the day, these small frogs take shelter in the axils of leaves of bromeli- ads and elephant ear plants. At night, they are active on native and cultivated vegetation, usually no more than 1.5 m above the ground. During 1977- 1978, JDL marked large numbers ofE. chalceus in a banana grove at the edge of Santo Domingo de los Colorados, Provincia Pichincha. As many as 30 individuals were encountered in a single evening on herbaceous vegetation in the banana grove, as well as on the leaves of the banana plants. The ease of collecting in the banana grove may contribute to JDL's impression that E. chalceus was more abun- dant in the banana grove than in primary forest 1 or 2 km away. Eleutherodactylus chalceus often is found on vegetation beside streams, but the observation of the species there probably reflects collectors' pref- erences for streams rather than that of the frogs, which also are abundant in the forest away from streams. Males were found calling in May at Maldonado, Provincia Carchi, and in June at Santo Domingo de los Colorados, Provincia Pichincha. Distribution. — This species occurs at eleva- tions of 50-1970 m from southwestern Departa- mento Antioquia, Colombia, to Provincia Guayas, Ecuador (Fig. 25). Twenty of the 25 Ecuadorian localities for this species are below 1000 m, where it seems to be far more abundant than it is at higher elevations. The species is absent from the drier regions of southwestern Ecuador, but one of the syntypes supposedly originated from Chimbo, Provincia Guayas, in the dry subtropical regime. Six Ecuadorian localities are in the humid tropical regime, whereas 18 are in the humid subtropical Remarks. — The two trivial names, areolatus and chalceus, were carried in Syrrhophus for much of this century, but little literature accumulated on either name until Lynch (1971a) provided a rede- scription of what he called E. areolatus, although Lynch (1968) briefly confused E. chalceus with what is now called E. lachmosus, a species in the Amazon Basin. Lynch (1980b) subsequently de- cided that the purported type locality ofPhyllobates chalceus (Pastassa-Tal, Colombia) must be some unknown locality on the western side of the Andes rather than the Pastaza Valley of eastern Ecuador; thus, he combined the two species because no features were available to distinguish Syrrhophus areolatus from Phyllobates chalceus. 81 80 0 20 40 60 80 100 • E. chalceus O E. duellmani Fig. 25. Distribution of two species of Eleuthero- dactylus in western Ecuador. ELEUTHERODACTYLUS IN WESTERN ECUADOR 81 Lynch and Burrowes (1990:20) included E. chalceiis and E. scolodiscus in a "small group of species having papillae at the digital tips (E. chalceus, E. gularis [Boulenger], and E. vocator [Taylor])." They added that these four species usually were associated with two species in lower Central America — E. diastema (Cope) and E. hylaeformis (Cope) — in spite of the absence of explicit shared characters. Eleutherodactylus chalceus has papillae on the tips of the digits on Fingers II-IV and Toes II-IV. The digital pads and disc covers oiE. vocator axe pointed but do not bear papillae. Lynch and Burrowes (1990) were con- fused about the condition of the digits in E. gularis as well; this resulted from Lynch's ( 1976b) incor- rect report concerning E. gularis. In the latter, the fingers bear no papillae on the pads or disc covers, but Toes II-IV have papillae. Eleutherodactylus scolodiscus is like E. gularis, except that it has papillae on the third fingers. Eleutherodactylus diastema and E. hylaeformis lack papillae on either fingers or toes. Lynch thinks that there are no fewer than two or three undescribed species in Colombia and Panama that have been confused (by various authors) withf. diastema, E. gularis, andf. vocator; two of these undescribed species have papillae on the fingers. Although most students of Costa Rican frogs have mentioned an Eleutherodactylus diastema group, none has been explicit about its characters. Cochran and Coin (1970) attempted to be explicit and cited the shape of the vocal sac (folds) and the short fingers and toes as supporting evidence. How- ever, Savage (1968) argued against the inclusion of E. moro in the group; although his comments were directed against the original description of a syn- onym off. moro, his comments represent an effec- tive criticism of the subsequently published ac- count by Cochran and Coin (1970). Savage (1968) noted that frogs of the E. diastema group have short, stubby fingers but pointed out that E. moro does not have short, stubby fingers and therefore is not a member of the E. diastema group. The fingers of E. chalceus are short and stubby and resemble, in some ways, those of E. diastema and E. gularis, but it is premature to suggest that we have any idea about the homologies among "short, stubby fingers" and "longer, less stubby fingers," let alone anything about polarities among the states of the character(s). Eleutherodactylus colomai sp. nov. Holotype.— QCAZ 1289, adult male, from Alto Tambo, 830 m (00°5r42" N, 78°30'54" W), Provincia Esmeraldas, Ecuador, one of a series collected on 6 December 1990 by Stella de la Torre, Diego Lombeida, Jean-Marc Touzet. and Felipe Campos. Paratype.— QCAZ 1296, collected with the holotype. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum smooth, that on venter finely areolate; discoidal fold prominent; dorsolat- eral folds absent; (2) tympanic membrane and tympanic annulus prominent, its length about '/3 length of eye; (3) snout subacuminate in dorsal view, protruding in profile; (4) upper eyelid lacking small tubercles, about as wide as lOD; cranial crests absent; (5 ) vomerine odontophores elongately oval, prominent; (6) males having vocal slits and white nuptial pads; mesorchium white; (7) first finger shorter than second; discs broad, round; (8) fingers lacking lateral fringes; (9) ulnar tubercles low; ( 10) heel bearing short calcar; outer edge of tarsus bearing indistinct tubercles; inner edge of tarsus lacking one tubercle and fold; (11) inner metatarsal tubercle oval, IVi-Ax round outer meta- tarsal tubercle; plantar surfaces areolate; (12) toes bearing indefinite lateral fringes near bases of dig- its; webbing absent; fifth toe much longer than third; (13) dorsum tan with dark sacral and suprainguinal spots; side of head and flanks darker brown with white flecks; venter cream with brown flecks; (14) SVL in two males 16.5 and 1 7.8 mm, females unknown. Eleutherodactylus colomai is most readily con- fused with E. rosadoi. These small frogs have similar color patterns, at least in preservative. How- ever, E. colomai has a protruding snout (rounded in E. ra^fl Jo/ ) and a calcar on the heel (small tubercles in E. rosadoi ), and lacks tubercles on the upper eyelid (present in E. rosadoi ). Moreover, the mesorchium is white in E. colomai and black in E. rosadoi. 82 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Description. — n = 2 males. Head narrower than body, slightly wider than long; HW 34.5 and 35 A%: snout subacuminate in dorsal view, protruding in profile, acuminate at tip (Fig. 26); E-N 84.8 and 95.2% length of eyes; nostrils slightly protuberant (primarily laterally), directed laterally; canthus rostralis angular, straight or weakly sinuous; loreal region flat, sloping abruptly (nearly vertically) to lips; lips not flared; upper eyelid and rest of head lacking tubercles; upper eyelid width 87.5 and 100.0% lOD; cranial crests absent; temporal re- gion vertical; supratympanic fold poorly defined, obscuring upper edge of tympanic annulus, extend- ing posteriorly to about level of upper arm but not curving ventrally; tympanic annulus adpressed against skin, clearly visible except uppermost por- tion; tympanic annulus round, its length 32.0 and 38.1% length of eye, separated from eye by dis- tance about half length of tympanic annulus; postrictal tubercles small, separate nonconical. Choanae round, not concealed by palatal shelf of maxillary arch; vomerine odontophores elongately oval, transverse, posteromedian to choanae, rela- tively prominent, separated medially by distance equal to about half width of odontophore, each bearing four or five teeth in transverse row; tongue wider than long, its posterior border not notched, posterior V-x not adherent to floor of mouth; vocal slits posterolateral to tongue; vocal sac single, subgular. Dorsum smooth, lacking tubercles and folds except for pair of supra-inguinal tubercles just lateral to midline; cloacal sheath and enlarged tubercles in cloacal region absent; skin on throat Fig. 26. Eleiitherodactyhis colomai, C, QC AZ 1 289, showing top and side of head and dorsal view of left foot, tarsus and heel. Scale bar = 2 mm. ELEUTHERODACTYLUS IN WESTERN ECUADOR 83 and belly finely areolate; discoidal fold well anteriad to groin. Upper and lower surfaces of limbs smooth, except skin below and ventrolateral to vent ar- eolate; three ulnar tubercles in row, antebrachial tubercle largest; thenar tubercle oval, smaller than bifid palmar tubercle; supernumerary palmar tu- bercles numerous; subaiticular tubercles low, round; fingers lacking lateral fringes (QCAZ 1296 might be scored as having thickened lateral folds on Fingers II and III); first finger shorter than second; discs on fingers broad, rounded, those on Fingers III and IV about twice width of digit proximal to disc, on Finger II about 1.5x width of digit (same size as tympanic annulus), on Finger I slightly wider than width of digit; all discs bearing broad ventral pads having complete circumferential grooves; nuptial pad white, on median surface of thumb above joint with metacarpal. Heel bearing short calcar (width at base equals length); outer edge of tarsus bearing row of indistinct tubercles; inner surface of tarsus lacking tubercle and fold; inner metatarsal tubercle twice as long as wide, about of round outer metatarsal tubercle; plantar surfaces areolate; subarticular tubercles round, low; toes bearing indefinite lateral fringes, most obvious at bases of toes; webbing absent; discs on toes slightly smaller than those on outer fingers; tip of Toe V extending to middle 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 overlapping; shank 49.4 and 49.7% SVL. Dorsum tan with diffuse darker markings across interocular region, in scapular region, and forming vague X-shaped mark postsacrally; dark brown spot above sacrum and darker suprainguinal spot; side of head dark brown with black spots above, and posterior to, tympanic annulus; brown facial mark extending onto flank to about level of sacrum, bordered above by slightly darker line, with white fleck within brown region; forearms dark brown with some darker brown spots (possibly remnants of cross bars); hind limbs pale brown with darker brown bars — oblique on shank, narrower than interspaces; knees dark brown; posterior surfaces of thighs brown heavily punctated with cream or cream with dense brown flecking. Throat and chest brown with cream flecks; posterior part of belly unpigmented except for brown flecks; undersides of hind limbs cream with brown flecks; region below vent brown but no defined triangular patch. Measurements of holotype: SVL 17.8, shank 8.8, HW 6.3, head length 6.0, upper eyelid width 1.8, lOD 1.8, tympanic annulus length 0.8, eye length 2.5, E-N 2.1. Coloration in life. — Color notes on three indi- viduals from Km 18 on Altaquer-Tumaco Road, Departamento Nariiio, Colombia, were provided by Pedro M. Ruiz-Carranza: Dorsum brown to coppery brown with dark brown transverse bands and yellow dorsolateral line; flanks olive-green with diagonal brown bars; loreal region brown; tympanum ochre; posterior surfaces of thighs dull yellow; throat dark brown; chest, belly, and under- sides of limbs red; iris bronze or brown above, red below. Natural history. — Both specimens were found on leaves of bushes 50-60 cm above the ground at night; one was in primary forest and one in second- ary forest. Distribution. — This species is known from the humid subtropical regime at elevations of 830- 1 200 m in the foothills of the Andes in northwestern Ecuador and southwestern Colombia (Fig. 22). Etymology. — The specific name is a noun in the genitive case and is a patronym for our Ecuadorian colleague, Luis A. Coloma, who made many col- lections of specimens available for our use. Remarks. — In Ecuador, this species is known from only the two males (holotype and paratype having a SVL of 16.5 mm) collected in 1990 and a juvenile female (ECO 217) having a SVL of 14.5 mm collected in 1994. Four specimens collected in 1995 in Departamento Nariiio, Colombia, include two males having SVLs of 13. 1 and 14.5 mm, one subadult female, 17.1 mm, and one juvenile fe- male, 1 1 .3 mm. Based on the somber dorsal color and pattern, we anticipate that this small species inhabits the leaf litter on the forest floor by day. Eleuthewdactylus cremmgiiis Lynch Plate 3 Eleuthewdactylus crenunguis Lynch, 1976a:2. — Holo- type: KU 1 20 126, adult male, from Tandapi, 1460 m, Provincia, Pichincha, Ecuador. 84 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Diagnosis. — A member of the Eleuthero- dactyhis (Eleutherodactylus) cerasinus group hav- ing (1 ) .skin on dorsum finely shagreen with .scat- tered warts and ridges, that on venter smooth; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus prominent, round, small, its length Vy--A length of eye; (3) snout subacuminate in dorsal view, rounded in profile, lips flared; (4) upper eyelid lacking tubercles, much broader than lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males having vocal slits; nuptial pads absent; (7) first finger longer than second; discs large, emarginate; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus bearing small, conical tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle elongate, 4- 6x round outer metatarsal tubercle; supernumerary tubercles few; (12) toes lacking lateral fringes; webbing absent; fifth toe longer than third; (13) dorsum gray with brown blotches; venter creamy white with brown marbling; posterior surfaces of thighs black with cream flecks; (14) SVL in males 35.2^9.2 mm, in females 59.1-64.5 mm. Eleutherodact}'liis crenungiiis is most similar to E. ocellatus. The two species can be distinguished by E. cretnmgius having tubercles on the heels (absent in E. ocellatus) and having the first finger longer than the second (shorter than the second in E. ocellatus). Juveniles oi E. labiosus can be con- fused with E. crenunguis, but E. labiosus has a conical tubercle on the upper eyelid and the first finger shorter than the second. Moreover, E. crenunguis usually has dark marbling on the ven- ter, whereas the venter in E. labiosus usually is white or cream without dark reticulations. Description. — The original description (Lynch, 1976a) is adequate. Coloration in life. — Detailed color notes in life are available for specimens from two localities; the differences may indicate geographic variation in color or different perceptions of color by two observers. KU 111213 from Tandapi, Provincia Pichincha: Dorsum pale brown with black mottling and minute yellow flecks; flanks gray, invaded by brown above and spotted with brown and black; lip barred with cream; groin gray; posterior surfaces of thigh black with white flecks; limbs pale burnt-orange with brown and black bars; throat creamy white with gray mottling; venter same but with pale orange blotches; under surfaces of hind limbs black with white spots; iris heavily reticulated with black, upper edge bright yellow bronze; most of iris brown-bronze, lowermost edge gray-bronze (JDL, 24 July 1967). KU 120125-29 from Tandapi, Provincia Pichincha: Dorsum medium brown to rust-brown with rust-orange to dark brown markings (one individual with white spots; another individual with yellow flecks and black mottling); flanks with black spots; hind limbs pale brown with darker brown bars; posterior surfaces of thighs dull brown to black with cream to white spots; venter dull yellow to orange (chin and throat black with white flecks in one individual); iris gold above and gray below median, horizontal brown streak (JDL, 27- 29 July 1968). KU 1 65223-29 from 3. 5 km NEMindo, Provincia Pichincha: Dorsum brown, reddish brown, or ol- ive-tan with orange-brown tubercles and ridges; upper lips with black and cream bars; posterior surfaces of thighs black with bluish white flecks; throat dull bluish white with gray mottling; belly dull pink to pale orange; iris bronze with black reticulations (WED, 7 April 1975). Natural history. — All individuals for which ecological data are available were found on low vegetation along, or on banks of, streams at night. During field work at Tandapi, Provincia Pichincha, in 1967 and 1968, adult males and juveniles of both sexes were found on the ground; subsequently, adult females were found on leaves and branches. A male was calling at 3 .5 km NE of Mindo, Provincia Pichincha, on 8 April 1975. Distribution. — This species occurs at moder- ate elevations (800-1 640 m) in the humid subtropi- cal regime on the western slopes of the Andes in Ecuador (Fig. 27). Remarks. — When first described, Eleuthero- dactylus crenunguis was associated with the E. fitzingeri group (Lynch. 1976a) because the first finger is much longer than the second, and the skin on the venter is smooth. During the next decade, JDL, encouraged by the criticism of J. M. Savage, began to abandon and then to dismantle that con- ELEUTHERODACTYLUS IN WESTERN ECUADOR 85 -1 -0 81 80 0 20 40 60 80 100 Kilometers 300 m 2000 m > 4000 m t^f'-yi Permanent V^ E. crenunguis E. degener E. hamiotae Fig. 27. Distribution of three species of Eleitthero- dactylus in western Ecuador. struction (the E.fitzingeri group sensu Lynch), and E. crenunguis was associated with E. rubicundus from the Amazonian slopes of the Andes. Finally, Lynch et al. (1994) associated E. crenunguis with other Chocoan taxa — E. cerasinus, labiosus, ocellatus, orpacobates, and tenebrionis. We think that E. ocellatus is the closest relative to E. crenunguis. The range of E. ocellatus is immedi- ately to the north of that off. crenunguis in Provincia Carchi. Ecuador, northward to Departamento Cauca. Colombia; hence, we do not expect E. crenunguis to range northward into Colombia. Eleutherodactylus crucifer (Boulenger) Plate 5 HylodescruciferBouXcngQx, 1899:456. — Holotype: BM 1947.2.16.91 from Porvenir. 1760 m. Provincia Bolivar, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum finely tuberculate with scattered conical tubercles on eyelids, back, and shanks, that on venter areolate; discoidal fold promi- nent; dorsolateral folds absent; (2) tympanic mem- brane and tympanic annulus prominent, round. small, its length Va-A length of eye; (3) snout short, rounded in dorsal view, truncate in profile; (4) upper eyelid bearing conical tubercles, broader than lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males hav- ing vocal slits and external vocal sac; nuptial pads present; (7) first finger shorter than second; discs on outer fingers broad; (8) fingers bearing crenulated lateral fringes; (9) ulnar tubercles conical; (10) heel bearing short calcar; inner and outer edges of tarsus bearing conical tubercles; (11) inner metatarsal tubercle oval, 4-6x round outer metatarsal tu- bercle; supernumerary tubercles absent; plantar surface areolate; (12) toes bearing crenulated lat- eral fringes; webbing absent; fifth toe much longer than third; (13) dorsum cream with brown mark- ings; venter cream with dark gray flecks; posterior surfaces of thighs cream with brown reticulation (uniform deep blue in life); (14) SVL in males 18.3-20.6 mm, in females 22.8-34.5 mm. Eleutherodactylus crucifer \s distinctive in hav- ing conical tubercles on the upper eyelids, forearm (ulnar tubercles ), heel, and outer edge of the tarsus. This pattern of conical tubercles also is present in the Colombian E. calcaratus, but that species lacks crenulated lateral fringes on the digits and has a much longer calcar. Living individuals are easily distinguished because the dorsum is green and the groin and concealed surfaces of the thighs are blue in E. crucifer as contrasted with a brown dorsum and concealed surfaces of the thighs in E. calcaratus. Description. — Lynch (1976a) redescribed the species based on a series of specimens fromTandapi, Provincia Pichincha, Ecuador. Coloration in life. — This species has a distinc- tive coloration — green dorsum with dark blue in the groin and on the concealed surfaces of the thighs; the blue is paler in juveniles and absent in the smallest individuals. Color notes on specimens from two localities are, as follow. KU 120133-40 from Tandapi, Provincia Pichincha: Dorsal surfaces of body and limbs bright green with dark green to black markings on dorsum, flanks, and limbs; groin, distal part of posterior surface of thigh, and underside of shank dark blue; proximal part of posterior surface of thigh cream with brown reticulations; venter dull cream (whitish cream on throat) with brown spots; 86 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 gray tint along lip; iris red above and white-bronze below median, horizontal brown streak. One indi- vidual with almost black venter; one individual pale green with white venter, and another with a cream patch on the middle of the back (JDL, 29 July 1968). KU 165235 from 3.5 km NE Mimlo, Provincia Pichincha: Dorsum bright green with dark green markings on body and dark brown bars on limbs; flanks pale yellowish-green with dark brown mark- ings; groin and underside of knee dark blue; venter silvery-white; other ventral surfaces greenish yel- low, all with brown flecks; iris dark red with pale bronze ventrally (WED. 7 April 1975). Natural history. — At night, individuals have been found on ferns and leaves up to 2 m above the ground, in dense forest, and along streams. The occurrence of a green frog with many conical warts on moss-covered leaves suggests that the frog is practicing some sort of crypsis, not unlike that of highland populations of Hyla loncasteri reported by Trueb ( 1968). By day. the frogs were found in arboreal bromeliads. Distribution. — Eleuthewdactyhis crucifer is known only from cloud forest in the humid sub- tropical regime at elevations of 1200-1800 m on the Pacific slopes of the Cordillera Occidental in Ecuador (Fig. 28). Remarks. — Two gravid females (ECO 087 and 123) from El Cristal, Provincia Esmeraldas. are remarkable in their small size (SVLs of 23.0 and 22.8 mm. respectively) in comparison with gravid females from Tandapi. Provincia Pichincha (SVLs of 26.2-28.5 mm). Eleutherodactylus degener sp. nov. Plate 5 Holotype.— QCAZ 1 297. adult male, from Alto Tambo, 830 m (00°51'42" N, 78°30'54" W), Provincia Esmeraldas. Ecuador, obtained on 6 December 1990 by Stella de la Torre, Jean-Marc Touzet, Diego Lombeida, and Felipe Campos. Paratypes. — ECO 77, 1 22. ( See Appendix 1 for localities.) Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum smooth, that on venter 80 0 20 40 60 80 100 Kilometers y 300 m ( 2(100 m CZ] >4(J00 m L_J Permanenl I iJ !• E. crucifer o \ — '-79 78 Fig. 28. Distribution of Eleutherodactylys crucifer. areolate; discoidal fold absent; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, round, its length '/w/? length of eye; (3) snout rounded in dorsal view, truncate in profile; (4) upper eyelid lacking tubercles, as wide as or slightly wider than lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs round; (8) fingers bearing lateral fringes; (9) ulnar tubercles small; (10) heel bearing minute tubercles; tarsus lacking tubercles; (11) inner metatarsal tubercle oval, 4x subconical outer metatarsal tubercle; su- pernumerary plantar tubercles indistinct; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; (13) dorsum brown or cream with brown flecks; venter white, with brown reticulation in females; posterior surfaces of thighs brown in male, cream with brown reticulations in females; (14) SVL in one male 22.2 mm, in two females 3 1 .0-32.7 mm. Eleutherodactylus degener is most similar to E. subsigillatus and, hence, presumably most closely related to E. eugeniae, nyctophylax, pho.xocephalus, and subsigillatus. Unlike the other species, E. degener lacks nuptial pads and has a smooth dor- sum (shagreen or finely tuberculate in the others). Moreover, E. subsigillatus differs by having small ELEUTHERODACTYLUS IN WESTERN ECUADOR 87 tubercles on the tip of the snout, upper eyeUds, and tarsi. In western Ecuador, the only other species having an orange iris is E. sobetes, which differs from E. degener hy having cranial crests, dorsolat- eral folds, and a tuberculate dorsum. Description. — (n = 3; 1 male, 2 females) Head wider than long, narrower than body: HW 39.6- 40.8% {x = 40.3) SVL; snout rounded in dorsal view (margin interrupted by protruding nostrils), truncate in profile (Fig. 29): E-N 81.2-100% ( f = 92.0) length of eye: snout longer than length of eye: nostrils small, protuberant, directed laterally: can- thus rostralis angular, weakly concave: loreal re- gion concave, sloping abruptly to lips: upper eyelid width equal to lOD in females, 112% in male: interorbital space flat: cranial crests absent: supratympanic fold low, angling posteroventrally behind tympanic annulus: tympanic membrane prominent, directed dorsolaterally: tympanic an- nulus visible except uppermost edge, round, its length 35.9^1 .7% (i = 38.5) length of eye, sepa- rated from eye by distance about 1 Vix tympanic membrane: postrictal tubercles low. Choanae round, not concealed by palatal shelf of maxillary arch; vomerine odontophores posteromedian to choanae, separated medially by a distance equal to width of one odontophore, each bearing five or six teeth in diagonal row: tongue round in male, longer than wide and feebly emarginate behind in females, posterior -A not adherent to floor of mouth: vocal slits short, posterolateral to tongue: vocal sac single, subgular. Skin on dorsum smooth, that on venter areolate: dorsolateral and discoidal folds absent: cloacal sheath and tubercles absent; ulnar tubercles small; palmar tubercle bifid, slightly larger than oval thenar tubercle: supernumerary palmar tubercles numerous: subarticular tubercles broader than long, simple, not elevated; first finger shorter than sec- ond: fingers bearing lateral fringes; similar fringe Fig. 29. Eleuthewdactylus degener, 9, ECO 77. showing top and side of head and pahnar view of hand. Scale bars = 2 mm. UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 evident along outer edge of palm; discs round. those of male 1 .3x width of digit proximal to disc, of females about 2x width; ventral surfaces of discs bearing pads dehned by circumferential grooves; male lacking nuptial pads. When hind limbs flexed perpendicular to body axis, heels broadly overlap- ping; shank 47.0-50.3% (i = 49.1) SVL; heel bearing minute tubercle on upper edge; tarsus lack- ing tubercle and folds; inner metatarsal tubercle about 2.5x as long as wide, about 4x size of rounded, subconical outer metatarsal tubercle; su- pernumerary plantar tubercles low; subarticular tubercles low, round, simple; toes bearing lateral fringes; webbing absent; discs on toes as large as those on outer fingers; tip of Toe V extending to distal edge of distal subarticular tubercle on Toe IV; tip of Toe III not extending to middle of penultimate subarticular tubercle on Toe IV. In preservative, dorsum of holotype brown with diffuse, slightly paler markings (interorbital bar and W-shaped mark in occipital region) and small dark brown flecks; limb bars and canthal stripe absent; white flecks along upper lip and on most tubercles; posterior surfaces of thighs uniform brown; preaxial surfaces of arms, groin, anterior surfaces of thighs, concealed surfaces of shanks, preaxial surfaces of tarsi, and feet unpigmented; brown encroachment on chin, but otherwise vocal sac and venter unpigmented. Dorsum of adult fe- male paratypes cream with brown flecks forming pattern — interorbital bar and chevrons in scapular and sacral regions; bars on limbs narrow; upper lips lacking flecks; labial bars and canthal and supratympanic stripes absent; throat uniform white; flanks and anterior surfaces of thighs with brown or black marks forming a reticulum; belly and under- sides of hind limbs white with brown spots and reticulations on belly and less distinct reticulations on hind limbs; posterior surfaces of thighs with large pale spots separated by brown lines having clusters of dark flecks within them; undersides of shanks patterned like posterior surfaces of thighs but less clearly so. Measurements of holotype: SVL 22.2, shank 11.1. HW 8.8, head length 8.3, upper eyelid width 2.8, lOD 2.5, tympanic annulus length 1.2. eye length 3.2. E-N 2.6. Coloration in life. — Based on 35-mm trans- parencies of an adult female provided by Martha L. Crump, the dorsum is brown with vague darker markings; the flanks are white with black reticula- tions, and the iris is orange. Color notes on a specimen collected in 1995 at Altaquer, Departa- mento Narino, Colombia, were provided by Pedro M. Ruiz-Carranza, as follow. Dorsum carmel col- ored, with a broad occipital triangle outlined with yellow; venter and hidden surfaces of hind limbs creamy yellow with red vermiculations; periphery and anterior part of chin reddish ochre; iris bright reddish-orange with brown flecks. Natural history. — The holotype was on a leaf of a bush 50 cm above the ground in primary forest at night. Distribution. — Eleiithemdactylus degener is known from only three localities at 830 and 1200 m in the humid subtropical regime on the lower slopes of the Andes in northwestern Ecuador and adjacent Colombia (Fig. 27). Etymology. — The specific epithet is a Latin noun meaning "different." The name is used in allusion to the orange eyes, which are strikingly different from the colors of the irises in most Eleutherodactylus. Remarks. — There is striking sexual dimorphism in coloration in the small sample of one male and two females. The dorsal pattern is subdued in the male in contrast to that in females. The groin and posterior surfaces of the thighs in females have reticulations, whereas these surfaces are uniform brown in the male; moreover, the belly is reticu- lated in females and unpigmented in the male. In the better known E. subsigillatus, a ventral pattern of reticulation develops with size in females, and the absence of dark markings on the belly in males is taken to be dependent on their smaller size than that of adult females. Eleutherodactylus dissimulatus sp. nov. Plate 5 Holotype.— KU 179088. adult female, one of a series collected at Quebrada Zapadores, 5 km ESE Chiriboga, 2010 m (00° 15' S. 78°43' W), Provincia Pichincha, Ecuador, on 8 July 1977 by David C. Cannatella, Marsha Lynch, and John D. Lynch. Paratopotypes.— KU 165923-25, 179087, 179089-94. ELEUTHERODACTYLUS IN WESTERN ECUADOR 89 Diagnosis. — A member of the Eleuthero- dactylus (Eleuthewdactxlus) unistrii>cifiis group having ( 1 ) skin on dorsum smooth except tor minute ridges, that on venter areolate; discoidal fold well anteriad of groin; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus present, round, its length V3--/5 length of eye; (3) snout long, subacuminate in dorsal view, angularly rounded in profile; (4) upper eyelid bearing minute tubercles, narrower than lOD; cranial crests absent; (5 ) vomer- ine odontophores oval; (6) adult males unknown; (7) first finger shorter than second; discs on outer fingers large, apically rounded; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles absent except for low antebrachial tubercle; (10) heel bearing subconical tubercle; outer edge of tarsus bearing poorly defined tubercles; inner edge of tarsus lacking tubercle or fold; (11) inner metatar- sal tubercle elongate, 6-8x subconical outer meta- tarsal tubercle; supernumerary plantar tubercles at bases of toes; (12) toes lacking lateral fringes; webbing absent; fifth toe much longer than third; (13) dorsum cream to brown with brown interorbital bar and scattered spots on flanks and thighs; venter cream with small brown spots; posterior surfaces of thighs, groin, and concealed shanks cream (or- ange in life) with black spots and irregular lines; ( 14) S VL in adult males unknown, in females 27.4— 32.5 mm. Eleutherodactylus dissimulatus is most similar to the slightly smaller E. calcarulatus. In addition to the slight difference in size, E. dissimulatus has a longer snout, straight rather than concave canthus rostralis, oval rather than oblique vomerine odontophores, and a pattern of spots or lines on an orange background in the groin and on the con- cealed surfaces of the hind limbs (uniform brown in E. calcarulatus). Description. — {n = 8; 1 male, 7 females) Head wider than long, narrower than body; HW 38.2- 42.0% (J = 39.5) SVL; snout longer than length of eye (Fig. 30); E-N 83.3-100% {x = 91 .2) length of eye; nostrils small, protuberant, directed laterally; canthus rostralis sharply angular, nearly straight; loreal region weakly concave, sloping abruptly to nonflared lips; upper eyelid width 72.7-87.5% ( x = 80.3) lOD; interorbital space flat; most individuals with minute conical tubercle on upper eyelid; Fig. 30. Eleutherodactylus dissimulatus. 9. KU 179088. Scale bar = 2 mm. supratympanic fold indistinct; tympanic annulus, round, its length 31.4^0.5% (v = 35.8) length of eye, separated from eye by distance equal to 1 Vi-lx tympanic annulus; postrictal tubercles small, coni- cal. Choanae moderate-sized, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores posteromedian to choanae, separated medially by a distance equal to width of one odontophore, each as large as a choana, each bear- ing three or four teeth in transverse row; tongue longer than wide, its posterior border feebly notched, posterior -A not adherent to floor of mouth. Skin on dorsum smooth except for minute eye- lid tubercle and a system of fine anatomizing ridges; discoidal fold well anteriad to groin; skin on flanks relatively smooth, that on venter areolate; cloacal sheath and tubercles absent. Palmar tubercle bifid, larger than oval thenar tubercle; supernumerary palmar tubercles indistinct; subarticular tubercles round; first finger shorter than second; fingers bear- 90 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 ing narrow lateral fringes; disc on thumb scarcely wider than digit proximal to disc, that on Finger II as wide as tympanic annulus; discs on Fingers III and IV larger than tympanic annulus; fingers long, slender. When hind limbs flexed perpendicular to body axis, heels broadly overlapping; shank 54.8- 60.0% (.f = 57.7) SVL; inner margin of tarsus lacking tubercle or fold; inner metatarsal tubercle three times as long as wide, not compressed; subarticular tubercles low, round; supernumerary plantar tubercles at bases of toes; fifth toe much longer than third; toes lacking lateral fringes; web- bing absent; discs on toes as large as those on fingers. In preservative, dorsum cream to brown with a dark brown interorbital bar and scattered small brown spots, especially on flanks and thighs; ob- lique bars on shanks brown, naiTower than pale interspaces; canthal and supratympanic stripes and labial bars indistinct or absent. Venter cream with small brown spots; groin, posterior surfaces of thighs, and concealed surfaces of shank nearly colorless with black spots or irregular line; in KU 179089, posterior surfaces of thighs black with cream spots. Measurements of holotype: SVL 30.7, shank 16.9, HW 12.1, head length 11.7, upper eyelid width 2.4, lOD 3.2, tympanic annulus length 1.7, eye length 4.2, E-N 3.8. Coloration in life. — The dorsum and venter are pale tan, orange, olive, or brown with an orange tint above and minute black flecks on the venter. The throat is orange with black flecks. The concealed surfaces of the hind limbs are orange or yellow with black spots or marbling. The iris is bright copper with a median, horizontal brown streak. Natural history. — All individuals were on low vegetation along streams in cloud forest at night. Distribution. — This species is known from only two localities at 1 920 and 2020 m in cloud forest in the humid temperate regime on the western flanks of the Andes in Provincia Pichincha, Ecuador (Fig. 24). Etymology. — The specific name is a Latin ad- jective meaning "disguised" or "hidden." The name is used in reference to the bright color pattern hidden on the concealed surfaces of the hind limbs. Remarks. — Only 1 3 specimens are known; the single male (KU 179094) is a juvenile having a SVL of 13. 1 mm. Juvenile females having minute, white, ovarian eggs and narrow, straight oviducts are as large as 25.4 mm in SVL (KU 165923). Eleutherodactylus dissimulatus seems to be most closely related to E. apiculatus and E. calcanilatus. Our association of these three species is not based on any synapomorphies. Eleutherodactylus duelhnani Lynch Plate 4 Eleutherodact}'lus diiellmani Lynch, 1980c:332. — Ho- lotype: KU 179325, adult male, from Quebrada Zapadores, 5 km ESE Chiriboga, 2010 m, Provincia Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) surdus group having ( 1 ) skin on dorsum bearing numerous small, flat warts, that on venter areolate; discoidal fold ill- defined; dorsolateral folds short, extending to level of sacrum; (2) tympanic membrane and tympanic annulus absent; (3) snout short, rounded in dorsal view, rounded to truncate in profile; (4) upper eyelid lacking tubercles, as wide as lOD; interorbital region furrowed; (5) vomerine odontophores large, subtriangular in outline; (6) males lacking vocal slits; nuptial pads small; (7 ) first finger shorter than second; discs broad; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus bearing small tubercles; inner edge of tarsus bearing large tubercle; (11) inner metatarsal tubercle oval, at least 8x indistinct outer metatarsal tubercle; supernumerary tubercles few at base of Toe IV; (12) toes having lateral fringes and basal webbing; fifth toe longer than third; (13) dorsum brown with diffuse darker markings; ven- ter gray to cream with diffuse brown spots or reticulations; posterior surfaces of thighs brown with a few cream flecks; ( 14) SVL in males 24.9- 36.0 mm, in females 36.6-45.8 mm. Eleutherodactylus duelhnani is most similar to E. hamiotae and E. surdus in that all three species have relatively short fifth toes that are only slighter longer than the third toes, and all three lack tym- panic annuli. Eleutherodactylus duelhnani differs from the others in having basal webbing on the toes and distinct, short dorsolateral folds, as contrasted ELEUTHERODACTYLUS IN WESTERN ECUADOR 91 to dorsolateral folds weak or absent and webbing between toes absent or extremely basal in the other species. Also, the fingers in E. duellmani are longer than those in E. hamiotae. Description. — The original description by Lynch ( 1980c) is adequate. Coloration in life. — Although this species lacks distinctive coloring, subtle differences in colora- tion exist, as noted in the following descriptions. KU 165908 from 9 km SE Tandayapa, Provincia Pichincha: Dorsum dull gray with tan dorsolateral folds; flanks grayish tan anteriorly, yellowish tan posteriorly, with dark olive-brown mottling; dorsal surfaces of limbs olive-tan with dark olive-brown bars; venter grayish tan with dark brown mottling; iris dull reddish-brown (WED, 9 April 1975). KU 165913-15 from Qiiebrada Zapadores, Provincia Pichincha: Dorsum olive-brown with dark brown interorbital crest and W-shaped scapu- lar crest, and dark brown and orange tubercles; posterior surfaces of thighs grayish tan with yellow flecks; venter dull yellow with cream flecks and greenish suffusion on belly; iris copper with black reticulations (WED, 3 April 1975). KU 179299-315 from La Delicia, Provincia Imhahura: Dorsum green, rust, or pale to dark brown with darker brown to black markings; venter gray to black with dirty white spots; throat same with white flecks; posterior surfaces of thighs brown with pale yellow spots; facial markings tinged with yellow; iris chocolate-brown to copper with black reticulations and hint of median, horizontal streak (JDL, 16 January 1978). KU 179338-46 from 14.8 km E Chiriboga, Provincia Pichincha: Dorsum green in males, dull brown in females; flanks bearing nearly yellow spots; posterior surfaces of thighs fleshy brown with cream flecks; dark labial bars and postocular stripe, latter bordered anteriorly by cream or yel- lowish mark; venter dull gray with cream flecks; iris chocolate-brown (JDL, 2 January 1978). Natural history. — At night, E. duellmcmi perches on branches and herbs immediately above small streams and sits on rock faces in the spray zones of waterfalls; some have been found in crevices or between rocks in small streams. This species apparently is sensitive to drying and re- mains where humidity approaches 100%. By day. these frogs were found under rocks, humus, or leaves in seepages, where water was running under the covers over the frogs. Some frogs escaped by hopping into streams. Distribution. — Eleutherodactyhis dueUmani occurs in cloud forests at elevations of 1550-2700 m on the west flank of the Andes from Departamento Cauca. Colombia, southward to Provincia Pichincha, Ecuador (Fig. 25). One locality is in the humid subtropical regime, whereas eight are in the humid temperate regime. Remarks. — Lynch (1980c) included E. dueUmani in his E. surdus group, then defined on the basis of having lost the middle ear and having a short, blunt snout. That species group (and asso- ciation) now seems to be an artificial grouping of most of the Ecuadorian Eleutherodactyhis lacking a middle ear. Species such as E. baryecuus, pugna.x, and ruidus have long fifth toes in contrast to the shorter fifth toes in the four species here restricted to the E. surdus group. Presently we suspect that Lynch's (1980c) original grouping confounded a series of species united only by inhabiting seep- ages. The character of relative lengths of the third and fifth toes rather sharply cleaves the E. surdus group of Lynch (1980c); many additional earless species, such as E. diaphonus and E. gracilis that are not members of the E. surdus group, have been found in Colombia. Eleutherodactylus duellmani, hamiotae, and surdus closely resemble one another and presumably are closely related; however, all three species occur in seepages and small streams, and their similarities may reflect only morphologi- cal adaptations to these microhabitats. Until addi- tional skeletal material becomes available, it is premature to include more than E. duellmani, hamiotae, sobetes, and E. surdus in the E. surdus group. Eleutherodactylus eremitus Lynch Plate 5 Eleutherodactylus eremitusLynch, 1980d:2. — Holotype, MCZ 92 1 03, adult female, from La Palma-Chiriboga road. 25.7 km above (NE) La Palma, 1 820 m, Provincia Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) unistrigatus group 92 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 having ( 1 ) skin on dorsum finely areolale, that on venter areolate; discoidal fold well anteriad to groin; dorsolateral folds absent; (2) lower part of tympanic annulus visible, round, its length, '/s-'/: length of eye; (3) snout subacuminate in dorsal view, rounded or weakly protruding in profile; (4) upper eyelid bearing conical tubercle, narrower than lOD; cranial crests absent; (5) vomerine odontophores oval; (6) males having vocal slits and nuptial pads; (7) first finger shorter than second; discs broad; (8) fingers bearing lateral fringes; (9) ulnar tubercles small, subconical; (10) heel and outer edge of tarsus bearing small tubercles; inner tarsal tubercle or fold absent; (11) inner metatarsal tubercle oval. 4x subconical outer metatarsal tu- bercle; supernumerary tubercles numerous; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; (13) dorsum cream with faint brown flecks and dark canthal stripe continu- ing onto flank; venter white; posterior surfaces of thighs and areas on flanks colorless (pale yellow in life); (14)SVLin males 17.2-21.8 mm, in females 27. 1-27.6 mm. Eleutherodactylus eremitus is a small green frog having either a loose reddish-brown reticulum on the dorsum or broad reddish brown dorsolateral stripes. This coloration sets it apart immediately from all other species in western Ecuador. Because E. eremitus has small tubercles on the heels and tarsi and a relatively large tympanic annulus, it might be confused with the lowland E. subsigillatus, which lacks a tubercle on the upper eyelid and usually has a pattern in the groin and on the con- cealed surfaces of the thighs. Description. — The original description by Lynch ( 1980d) is adequate. Coloration in life. — The dorsum is green, usu- ally with darker markings, and the venter is pale yellow; descriptions of individuals are, as follow. KU 165884 from 3.5 km NE Mindo, Provincia Pichincha: Dorsum pale green with dull red mark- ings; limbs greenish yellow; venter pale yellow; iris copper (WED, 7 April 1975). KU 179085-86 from Qiiehrada Zapadores, Provincia Pichincha: Head brown or reddish brown ; body green, latter with black spots in one and reddish-brown dorsolateral stripes in one; venter pale yellow or white with minute brown flecks on throat; concealed surfaces of hind limbs pale yel- low; iris bright copper with brown flecks (JDL, 8, 9 July 1977). Natural history. — Eleutherodactylus eremitus is relatively uncommon in collections. An observa- tion by Lynch and Burrowes (1990) that one was seen 7 m above the ground on an observation tower suggests that the species may seldom occur near the ground. By day, several individuals have been found in arboreal bromeliads, and one was in a terrestrial bromeliad; others were sitting on leaves of herbs and bushes to heights of 2 m above the ground. Distribution. — This small species occurs in cloud forests at elevations of 1540-2100 m on the western flanks of the Andes in extreme southern Colombia (Departamento Narino) and northwest- ern Ecuador (provincias Cotopaxi and Pichincha) (Fig. 31). Three localities are in the humid sub- tropical regime, and five are in the humid temperate regime. Remarks. — The relatively broad and flat head of this species led Lynch (1980d) to consider Eleutherodactylus eremitus to be a member of the E. lacrimosus group (or assembly). The features cited by Lynch (1980d) and Lynch and Duellman ( 1 980) possibly reflect the bromeliad habitat rather -0 81 80 0 20 40 60 80 100 -1 Kilometers Z' 300 m (^ --- 2000m N . [ 1 >40(Klm i \ ] Permanent ^ snow j^ Fig. 31. Distribution of two species oi Eleuthero- dactylus in western Ecuador. ELEUTHERODACTYLLJS IN WESTHRN KCUADOR 93 than relationships. As noted by Lynch ( 1 98()d), this species may have been under collected because few collectors sample bromeliads in the cloud forest. Ek'Hthewdactyliis eugeniae sp. nov. Plate 6 Holotype.— KU 165899, adult male, one of a series collected at Quebrada Zapadores. 5 km ESE Chiriboga, 2010 m (00° 15" S, 78°43'W), Provincia Pichincha, Ecuador, on 3 April 1975 by William E. Duellman. Paratypes.— KU 165900-904, 179382-83. and QCAZ 2629. (See Appendix I for localities.) Diagnosis. — A member of the Eleuthero- davtylus (Eleutheroclactylus) imistrigatus group having ( 1 ) skin on dorsum finely shagreen, becom- ing coarse on flanks, that on venter areolate; discoi- dal fold well anterior to groin; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, round, its length V^-A length of eye; (3) snout moderately long, subacuminate in dorsal view, rounded in profile, with papilla at tip; (4) upper eyelid lacking tubercles, narrower than lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males hav- ing vocal slits, subgular vocal sac, and white nup- tial pads; (7) first finger shorter than second; Fin- gers Il-IV bearing large, apically rounded discs; (8) fingers bearing lateral fringes; (9) ulnar tu- bercles absent, except for antebrachial tubercle; ( 1 0) heel and outer edge of tarsus lacking tubercles; inner tarsal tubercle indistinct; (11) inner metatar- sal tubercle oval, 4-6x oval outer metatarsal tu- bercle; supernumerary plantar tubercles low, at bases of toes; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; ( 1 3) dorsum cream to pale brown with brown spots; bars absent on limbs; venter cream with brown flecks on throat; posterior surfaces of thighs and flanks cream with brown reticulations; (14) SVL in males 25.2-26.0 mm, in six females 30.5-37.6 mm. Eleutheroclactylus eugeniae is most similar to E. nyctophylax, a species of equal size on the Pacific versant of the Ecuadorian Andes. The two differ in that E. nyctophylax has small tubercles on the upper eyelid and heel, a more angular canthus rostralis, areolate plantar surfaces, and the poste- rior surfaces of thighs brown with cream flecks. Description. — (/; = 7 for proportions of females [including one immature] or 8, if sex not specified) Head as wide as body, as wide as long; HW 36.5- 40.8% {x = 39.2) SVL; papilla at tip of snout (Fig. 32); E-N 90.6% length of eye in one male, 89.5- 1 13.9% (x = 104.6) in females; snout longer than length of eye; nostrils weakly protuberant, directed dorsolaterally; canthus rostralis weakly angular, nearly straight; loreal region flat, sloping abruptly to lips; lips weakly flared; upper eyelid width 65.9- 80.6% (f = 7 1 .8) lOD; interorbital space flat; most individuals with minute conical tubercle on upper eyelid; supratympanic fold indistinct; tympanic annulus round, superficial, indistinct externally in Fig. 32. Eleutlicrodacrsliis eugeniae. 9, KU 1 65899. Scale bar = 2 mm. 94 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 only male, prominent in females, its length 31.2^^ length of eye in one male. 36.7^2.7% ( .r = 4 1 .5) in females, separated from eye by distance equal to 1 '/;x length of tympanic annulus; postrictal tu- bercles small, conical; tongue as long as wide, its posterior border feebly notched, posterior V?. not adherent to floor of mouth; choanae moderate sized, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores posteromedian to choanae, separated medially by a distance equal to width of one odontophore, each larger than a choana, each bearing five or six teeth in transverse row; male with vocal slits lateral to tongue. Skin on dorsum of body finely shagreen, coarser on flanks, and finely granular on upper surfaces of limbs; skin on belly areolate, that on throat smooth; discoidal fold well anteriad to groin; cloacal sheath and tubercles ab.sent. Palmar tubercle bifid, slightly larger than oval thenar tubercle; supernumerary palmar tubercles small, indistinct; subarticular tu- bercles round, pungent; fringe along outer edge of palm and Finger IV; disc on thumb narrow, that on Finger II as wide as tympanic annulus; discs on Fingers III and IV larger than tympanic annulus. Heel of adpressed hind limb reaching point just anterior to eye; when hind limbs flexed perpen- dicular to body axis, heels overlapping; shank 5 1 .4-56.6% ( .V = 53.2 ) SVL; inner margin of tarsus having indistinct tubercle (difficult to detect); inner metatarsal tubercle twice as long as wide, not compressed; subarticular tubercles round, not coni- cal; supernumerary plantar tubercles at bases of toes: discs on toes as large as those on fingers. In preservative, dorsum cream, gray, or pale brown with scattered brown spots on the limbs and (or not) on digits; flanks with brown reticulations; dorsum with or without creamy-white spots out- lined in brown and/or white interorbital bar edged with brown. Faint supratympanic stripe evident; labial bars, canthal stripe, and limb bars absent. Venter cream, with or without faint brown reticula- tion on belly; brown flecks on throat; ventral sur- faces of thighs and shanks unmarked; unpigmented area in groin; anterior surfaces of thighs brown, posterior surfaces cream with brown reticulations, except for brown patch behind knee. Measurements of holotype: SVL 30.5, shank 16.4, HW 12.1, head length 10.1, upper eyelid width 2.9. lOD 3.9, tympanic annulus length 1.4, eye length 3.8. E-N 3.4. Coloration in life. — The dorsum is pale creamy tan to dusky yellow; if pale spots are present, they are pale yellow to pale orange. There is a faint brown reticulum on the flanks; the throat and venter are white, and the concealed surfaces of the hind limbs are a flesh color to brown. The iris is pale greenish-gray to coppery orange with brown flecks or fine black reticulation. Natural history. — Eleiithewdactylus eugeniae inhabits cloud forests away from streams. In July 1977, one individual was 2 m and another 4 m above the ground; JDL no longer recalls what possessed him to climb the tree to find the latter specimen, but this observation suggests that the apparent rarity of this species may reflect the frog's use of a stratum above the collecting zone of most biologists. Distribution. — This species is known only from three localities in a limited area of cloud forest at elevations of 1700-2010 m in the upper valley of the Rio Pilaton in Provincia Pichincha, Ecuador (Fig. 33). One locality is in the humid subtropical regime, and two are in the humid temperate regime. Etymology. — The specific name is a noun in the genitive case and is a patronym for Eugenia del Pino of the Pontificia Universidad Catolica del Ecuador. For more than a decade she extended many courtesies to us while we were working in Ecuador, and she accompanied WED to the type locality in 1977. Remarks. — Only nine specimens are known. The only juvenile (KU 165901) is a female 25.5 mm in SVL. We think that E. eugeniae is most closely related to E. nyctophyla.x, but we are un- aware of any synapomorphy to link the pair of species. Eleutherodactylus floridus sp. nov. Holotype.— USNM 239672, adult male, ob- tained in "region of Sigchos, [Provincia] Cotopaxi," Ecuador, in June 1957 by Manuel Olalla. Paratypes.— USNM 239674-92. (See Appen- dix I for localities.) Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) myersi group having ELEUTHERODACTYLUS IN WESTERN ECUADOR 95 81' \ 80" 0 20 40 60 80 100 ^3^^^ 78° -1° Kilometers /" j^ V^\ " ^ 300 ni ( 2000 m \ r^ r~l >4(K)()m 1 ' j 1 Permanent k _ sndw ^ ^^o:'""' -'-■ '^^ ^' -0° d-^'- /\^"X o' , 1^ '( r^W CO ', ' ~ ^ : T- -2 ] ^l 1 / ; ^'^'5; 2- -3 ■^ , .■■■^ 3°- ■';'' ▲ £. eugeniae /wL- , - ry • £. gidaris ■ £■. lielonotHS (T^^''^^^ '' ' --'^)L'' 81° / :V: y 80\ 'j~ O E. labiosus 78° 79° \^ ■•' ' rr^y l-V l_ Fig. 33. Distribution of four species of Eleuthew- dactylus in western Ecuador. (1) skin on dorsum shagreen with many rounded warts, that on venter areolate; discoidal fold absent; dorsolateral folds absent; (2) tympanic membrane prominent, and tympanic annulus thin and not prominent, higher than long, its length Vy--/> length of eye; (3) snout short, subacuminate in dorsal view, rounded in profile; (4) upper eyelid bearing low tubercles, about as wide as lOD; cranial crests absent; (5) vomerine odontophores prominent, oval to triangular in outline; (6) males lacking vocal slits and nuptial pads; (7) hrst finger shorter than sec- ond; discs on outer fingers round, about twice width of digit proximal to pad; (8) fingers bearing narrow lateral keels; (9) ulnar tubercles absent; ( 1 0) heel and inner edge of tarsus lacking tubercles; outer edge of tarsus bearing row of small tubercles; (II) inner metatarsal tubercle oval, about 4x round outer metatarsal tubercle; supernumerary plantar tubercles absent; ( 12) toes bearing narrow lateral keels; webbing absent; fifth toe slightly longer than third; discs smaller than those on outer fingers; (13) dorsum brown with diffuse darker markings; ven- ter cream with brown marbling; throat brown; groin and posterior surfaces of thighs cream with brown marbling; ( 14) SVL in two males 17.4 and 18.3 mm. in 17 females 23.3-26.7 mm (.v = 25.5). Eleuthewdactylusfloridus differs from all other species in the E. myersi group by having larger discs. It also differs in color pattern, but probably has red areas on the concealed surfaces of the hind limbs. Eleutherodactyhis floridus lacks dorsolat- eral folds (present in E. hectiis) and has white mesorchial lining (black in E. leoni). Because of its large digital discs, E. floridus superficially resembles species in the E. leptolophus group in Colombia, but in all of those frogs the fifth toe is much longer than the third (only slightly longer in E. floridus). Description. — (/; = 19; 2 males, 17 females) Head wider than body in males and immature females, slightly narrower than body in gravid females, wider than long; HW 40.2 and 40.4% in males, 38.9-42.5% (v = 40.3) in females; snout short, subacuminate in dorsal view, rounded in profile (Fig. 34); E-N 70.8 and 7 1 .4% length of eye in males, 65.7-80.6% (v = 72.7) in females; nos- trils protuberant, directed laterally; canthus rostralis angular, concave; loreal region weakly concave, sloping abruptly to lips; lips not flared; upper eyelid bearing small, pustular tubercles (not larger than those on rest of head); upper eyelid width 1 00.0 and 135.3% lOD in males, 88.0-107.7% (f = 98.2) in females; cranial crests absent; supratympanic fold indistinct but obscuring upper edge of tympanic annulus; side of head vertical; tympanic annulus thin, only anterior and ventral parts visible when pressed against skin; tympanic membrane not pustular or thickened; tympanic annulus round in males, higher than long in females; length of tym- panic annulus 33.3 and 39.3% length of eye in males, 3 1 .4-39.2% ( r = 34.0) in females; postrictal tubercles subconical, posterior to tympanic annu- lus; skin on head bearing low warts, even on face, but those above tympanic annulus most prominent. Choanae oval, not concealed by palatal shelf of 96 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 34. Eleutherodactylus floridus, USNM 239673, 9, showing top and side of head, and USNM 239674. 9, showing palmar view of hand. Scale bars = 2 mm. maxillary arch, especially small in adult females, proportionately larger in small individuals; vomer- ine odontophores posteromedian to choanae, oval to nearly triangular in outline, each 3-5x size of a choana, separated medially by distance equal to width of odontophore, each bearing three to five teeth in transverse row; tongue longer than wide, its posterior border not notched, posterior -A-A not adherent to floor of mouth; males lacking vocal slits. Dorsum shagreen with many, scattered large warts; largest warts in areas of darker pigment; warts forming postocular ridges and in some indi- viduals shorter ridges postsacrally; dorsolateral folds absent; flanks with higher density of large warts than on dorsum; cloacal sheath and enlarged tubercles in cloacal region absent; skin on throat smooth or wrinkled; skin on venter areolate; dis- coidal fold absent. Upper surfaces of arms tubercu- late; ulnar tubercles absent or in low row along postaxial surface of forearm; thenar tubercle oval, much smaller than bifid palmar tubercle; supernu- merary palmar tubercles prominent, round, smaller than round, nonconical subarticular tubercles; fingers bearing narrow lateral fringes; first finger shorter than second; disc on thumb not expanded; discs on Fingers III and IV about twice width of digits, that on Finger II smaller; all fingers having ventral pads defined by circumferential grooves; nuptial pads absent in males. Upper surfaces of ELEUTHERODACTYLUS IN WESTERN ECUADOR 97 hind limbs warty, largest warts on limb bars; heel tubercles absent; outer edge of tarsus bearing row of small tubercles; inner surface of tarsus lacking tubercles; inner metatarsal tubercle two and one- half times as long as wide, about four times size of round outer metatarsal tubercle; supernumerary plantar tubercles absent; subarticular tubercles round, nonconical; toes bearing narrow lateral keels; webbing absent; discs on toes expanded but smaller than those on outer fingers; tip of Toe V extending just beyond penultimate subarticular tubercle on Toe IV; tip of Toe III extending to base of that tubercle; when hind limbs flexed perpendicular to axis of body, heels overlapping; shank 53.4 and 56.5% SVL in males, 50.0-57.1% (v = 53.6) in females. Dorsum brown with subdued, darker brown markings (chevrons, blotches, labial bars, canthal- supratympanic stripe); cloacal triangular dark brown; bars on shanks (when present) transverse, about equal to width of interspaces; throat brown with cream flecks; belly and ventral surfaces of limbs cream with brown marbling; areas at bases of arms, on lower flanks, in groin colorless (cream); anterior and posterior surfaces of thighs cream with dark brown marbling. Measurements of holotype: SVL 26.0, shank 13.6, HW 1 0.2, head length 9.6, upper eyelid width 2.2, lOD 2.5. tympanic annulus 1 .0. eye length 3.3, E-N 2.5. Coloration in life. — Unknown. Natural history. — Unknown Distribution. — This species is known only from provincias Cotopaxi, Imbabura, and Pichincha. Many of the specimens have elevational data and were collected by members of the Olalla family. Paynter and Traylor (1977) discussed many of these localities; thus, it seems that E. floridus oc- curs at elevations of 700-2000 m on the western flank of the Andes (Fig. 35). Nine of the 12 locali- ties seem to be in the humid subtropical regime; the other three are in the humid temperate regime. Etymology. — The specific name is a Latin ad- jective meaning "abounding with flowers." The name is used in allusion to Glen Flores in recogni- tion of his contributions to the study of Eleuthewdactylus and whose surname is Spanish for "flowers." -0 81 80 0 20 40 60 80 100 1 Kilometers Z' 100 m ( 200(1 ni \ [ IZ] >40(10 m 1 r PerniaiiL-nl >v smiw ^ Fig. 35. Distribution of two species of Eleiithero- dactylus in western Ecuador. Remarks. — One juvenile female (USNM 239791) has a SVL of 16.1 mm, and one nearly aduh female (MCZ 90320) has a SVL of 22.2 mm. We think that E. floridus is the nearest relative off. pyrrhomerus, even though E. floridus has much larger digital discs. However, most of the features held in common by the two species seem to be plesiomorphic. Eleutherodactyhis gentryi sp. nov. Plate 3 Holotype. — KU 1 3 1 540. adult female, one of a series collected "5 km E (by road) Pilalo" (esti- mated distance), 3200m (-00°56' 15" S,78°45' 30" W), Provincia Cotopaxi, Ecuador, on 7 July 1970 by John D. Lynch. Paratypes.— KU 131541-50, USNM 239763, 239768-7 1 . (See Appendix I for localities.) Diagnosis. — A member of the Eleuthero- dactyhis (Eleutherodactylus) curtipes group hav- ing (1) skin on dorsum bearing low warts (most obvious on flanks), that on venter areolate; discoi- dal fold well anteriad to groin; dorsolateral folds present; (2) tympanic membrane and usually tym- panic annulus absent; (3 ) snout shoit, subacuminate 98 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 to rounded in dorsal view, rounded in protile; (4) upper eyelid lacking tubercles, narrower than lOD; cranial crests and bony tubercles between eyes; (5) vomerine odontophores oblique; (6) males lacking vocal slits and nuptial pads; (7) first finger shorter than second; discs on outer fingers narrow, about 1.5x width of digit proximal to pad; (8) fingers bearing nanow lateral keels; (9) ulnar tubercles absent except for low antebrachial tubercle; (10) heel bearing nonconical tubercle; inner and outer edges of tarsus bearing minute tubercles; (11) inner metatarsal tubercle oval, about 3x subconical outer metatarsal tubercle; supernumerary plantar tu- bercles numerous; ( 1 2) toes bearing lateral fringes; webbing absent; fifth toe longer than third; (13) dorsum brown with little pattern; pale labial stripe present; venter cream with small brown flecks; posterior surfaces of thighs brown; (14) SVL in males 23.0-28.5 mm, in females 29.6-35.5 mm (.f = 33.4). Eleutherodactylus gentry i is similar to E. curtipes but differs from that species in having obvious (albeit small) digital discs, bony tubercles on the frontoparietals between the cranial crests, longer hind limbs, and in lacking a tympanic annulus; in E. curtipes, the tympanic annulus is fully formed but lies concealed beneath the skin. Description. — (n = 12; 4 males, 8 females) Head no broader than body (less so in gravid females), wider than long; HW 39.4-40.0% (.v = 39.7) SVL in males, 36.3^1.7% (,r = 39.2) in females; snout short, subacuminate to rounded in dorsal view, rounded in profile; E-N 78.6-82.8% (J = 80.9) length of eye in males, 70.7-86.5% (,r = 81.6) in females; nostrils weakly protuberant, di- rected dorsolaterally; canthus rostralis angular, straight or weakly concave; loreal region concave, sloping abruptly to lips; lips not flared; upper eyelid bearing low tubercles, its width 65.7-86.2% (,f = 76.5) lOD in males, 63.0-78.8% (r = 68.3) in females; interorbital region broad, its surface re- flecting structure of underlying bones (low cranial crests just medial to eyes, bony tubercles between crests, extending onto nasals in some specimens); supratympanic fold indistinct, made less evident by tubercles above and below fold; tympanic annu- lus never visible externally; when detectable, an- nulus weak, higher than long; postrictal tubercles round, not prominent. Choanae small, round, not concealed by palatal shelf of maxillary arch; vomer- ine odontophores posteromedian to choanae, ob- lique, each about three times size of a choana, separated medially by distance equal to half width of odontophore, each bearing four or five teeth in oblique row; tongue broader than long, its posterior border not notched, posterior -/s not adherent to floor of mouth; males lacking vocal slits and sac. Dorsum smooth to feebly warty; dorsolateral folds usually distinct, fragmented posterior to sacrum; area between dorsolateral folds bearing small warts; skin on flanks bearing larger warts; skin of venter areolate; discoidal fold well anteriad to groin; cloacal sheath and tubercles absent; skin on upper surfaces of limbs smooth or shagreen; ulnar tubercles indistinct except for small antebrachial tubercle. Thenar tubercle oval, about Vi size of bifid palmar tubercle; numerous, low supernumerary palmar tubercles; subarticular tu- bercles round to wider than long, low; first finger shorter than second; fingers bearing fleshy lateral keels; tip of thumb not expanded into disc; Fingers II-IV bearing round discs, those on Fingers III and IV about V/2X width of digit proximal to disc; all digits having broad ventral pads; nuptial pads ab- sent in males. Upper edge of heel bearing low tubercle; less prominent tubercles on inner edge of tarsus and forming row on outer edge of tarsus; inner metatarsal tubercle oval, about 3x subconical outer metatarsal tubercle; numerous low supernu- merary plantar tubercles, indistinct except at bases of toes; subarticular tubercles round, not conical; toes bearing lateral fringes; webbing absent; toe discs round, bearing ventral pads, smaller than those on outer fingers; tip of Toe V extending to distal base of distal subarticular tubercle on Toe IV; tip of Toe III extending to proximal border of penultimate subarticular tubercle on Toe IV; when hind limbs flexed axis of body, heels not overlap- ping (touching in a few specimens); shank 46.5- 50.0% (,f = 47.8) SVL in males, 41.6-50.9 {x = 46.0) in females. In preservative, dorsum brown with darker brown markings consisting of rectangular blotch extend- ing from eyes to sacrum, scattered small spots on body, limb bars fragmented into spots, indistinct canthal-supratympanic stripe, and hints of labial ELEUTHERODACTYLIIS IN WESTERN ECUADOR 99 bars; throat paler brown; belly cream with small brown flecks or uniform brown; anterior and pos- terior surfaces of thighs pale brown in most indi- viduals; pale patch behind knee and in groin, and venter pale in USNM 239763. Measurements of holotype: SVL 34.4, shank 17.5, head width 13.5, head length 12.0, chord of head length 1 3.0. upper eyelid width 2.9, lOD 4.6 eye length 4.2. E-N 3.6. Coloration in life. — The dorsum is variable, and the venter and concealed surfaces of the hind limbs usually are at least tinted with a salmon color, as noted in the following description. KV 1 3 1 540-65 from 5 km E Pilalo, Provincia Pichinclui: Dorsum red to brown with darker brown to black markings (usually a dark reddish-brown blotch); flanks paler than dorsum; venter dirty- cream to salmon; posterior surfaces of thighs salmon-brown to brown; labial stripe creamy bronze; iris pale copper with reddish-brown, me- dian, horizontal streak (JDL, 7 July 1970). Natural history. — All individuals were under rocks by day. Males collected in July have swollen testes with dark mesorchia (either brown reticula- tion or uniform stippling). The smallest gravid female (USNM 239763, 29.5 mm in SVL) is only slightly larger than a female estimated to be enter- ing reproductive condition (USNM 239738, 28.2 mm). Distribution. — This species is known only from elevations of 2850-3380 m in the upper reaches of cloud forest, principally in the humid subtemperate regime, west of the Paramo de Apagua, Provincia Cotopaxi, Ecuador (Fig. 24). Etymology. — The specific name is a noun in the genitive case and is a patronym for our late friend and colleague, the intrepid botanist Alwyn H. Gen- try, who was killed while conducting field work in western Ecuador on 3 August 1 993 . We think of this species as residing high in the cloud forests of western Ecuador and figuratively looking west- ward over the remnants of forest on the lowlands of western Ecuador, and dedicate this species to the memory of "Al" Gentry, who loved the tropical forests and who enriched the lives of everyone who knew him. Remarks. — Numerous specimens referred to this species are either juveniles or, in the case of USNM 239738-62. partially crushed. Although specific identity is not a problem, we are reluctant to designate juveniles and poorly preserved speci- mens as paratypes. Eleutherodactylus gentryi is a member of the E. curtipes group, which also in- cludes E. buckleyi, cryophilus, curtipes, and two recently described species {E. satagius andf. xestus) from Colombia (Lynch, 1995) The sharing of broadly expanded frontoparietals that extend over the posteromedial portions of the orbits unites E. curtipes and E. gentryi as probable sister species. Eleutherodactylus gularis (Boulenger) Plate 8 //y/oJe'.v g/z/^m- Boulenger. 1898:463.— BM 1947.2.15.82, adult male, from Cachabe. Provincia Esmeraldas, Ecuador. Hyloxalus huigrae Fowler. 1913:165 . — Holotype: ANSP 18113, adult male, from Huigra, 4000 ft, Provincia Chimborazo. Ecuador. Synonymy fide Lynch, 1976b: 14. Eleutherodactylus diastema — Dunn, 1942:2. Eleutherodactylus gularis — Cochran and Coin, 1970:441; Lynch. 1976b: 14. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) diastema group hav- ing ( 1 ) skin on dorsum smooth, that on venter areolate; discoidal fold absent; dorsolateral folds absent; (2) tympanic membrane absent; tympanic annulus visible through skin, round, '/^--/s length of eye; (3) snout subacuminate in dorsal view, trun- cate in profile; (4) upper eyelid lacking tubercles, narrower than lOD; cranial crests absent; (5 ) vomer- ine odontophores oval in outline, narrowly sepa- rated; (6) males having vocal slits and large subgular vocal sac; nuptial pads absent; (7) first finger shorter than second; discs slightly wider than digits; disc covers feebly pointed but lacking papillae; (8) fingers broad, bearing weak lateral fringes; (9) ulnar tubercles absent; ( 1 0) heel and tarsus lacking tubercles and folds; (11) inner metatarsal tubercle oval, 3x round outer metatarsal tubercle; supernu- merary plantar tubercles absent; (12) toes broad, lacking lateral fringes; webbing basal; fifth toe much longer than third; discs lanceolate with minute papillae at tips of Toes II-IV; (13) dorsum pale brown with vague markings; snout pale; venter 100 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 cream with brown stippling on thrt)at; posterior surfaces of thighs brown; (14) SVL in males 20.2- 21.6 mm, in females 23.3-24.8 mm. Eleuthewdactylus gidaris is most similar to E. diastema but differs in having fleshy, basal web- bing on the foot and in having pointed disc covers, which on Toes III-IV bear small papillae. Description. — {n = 16: 9 males, 7 females) Head as broad as body (less so in gravid females), longer than broad; HW 34.0-39.1% (x = 36.7 ± 0.4) SVL; snout long, subacuminate in dorsal view, truncate in profile (Fig. 36); nostrils not protuber- ant, directed laterally (with slight dorsal vector), near tip of snout; E-N 64.7-92.6% ( x = 78.4 ±1.9) length of eye; canthus rostralis rounded, straight; loreal region flat, sloping abruptly to lips; lips not flared; upper eyelid lacking tubercles, its width 55.2-80.8% (x = 66.0 ± 2.8) lOD; interorbital region flat; cranial crests absent; temporal region nearly vertical; supratympanic fold low, poorly defined; lower 3/4 of tympanic annulus evident, round, separated from eye by distance equal to % of its diameter; length of tympanic annulus 28.6- 42.4% ( X = 34.5 ±1.3) length of eye; two postrictal tubercles posteroventral to tympanic annulus. Choanae long, oval, not concealed by palatal shelf of maxillary arch; vomerine odontophores posteromedian to choanae, oblique, each larger than a choana, separated medially by distance equal to half width of odontophore, bearing five or six teeth in oblique row; tongue much longer than broad, its posterior border not notched, posterior half not adherent to floor of mouth; males having short vocal slits lateral to tongue; external vocal sac Fig. 36. Eleutherodactylus gidaris. cS. QCAZ 43 1 6. showing top and side of head and palmar view of hand. Scale bars - 2 mm. ELEUTHERODACTYLUS IN WESTERN ECUADOR 101 commonly forming longitudinal folds median to mandibles in preserved specimens. Dorsum smooth except for some low tubercles on back postsacrally in some specimens; dorsolat- eral folds absent; skin on lower flanks and venter with low areolations (most evident laterally); dis- coidal fold absent; cloacal sheath and tubercles absent; forearm slender; ulnar tubercles absent. Thenar tubercle oval, smaller than oval palmar tubercle; supernumerary palmar tubercles absent; subarticular tubercles round, flattened; flrst finger shorter than second; fingers broad, bearing weak lateral keels; digital discs only slightly wider than digits (Fig. 36); disc covers feebly pointed; nuptial pads absent in males. Heel and tarsus lacking tubercles and folds; inner metatarsal tubercle about 2'/! times as long as wide, not compressed, 3x round, unelevated outer metatarsal tubercle; super- numerary plantar tubercles absent; subarticular tu- bercles low, round; digits flattened, lacking lateral keels; webbing basal, formula: I 2 — 2^ II 2 — 3* III 2'/: — 4^ IV 4^ — 2- V; toe discs lanceolate with minute papillae at tips of Toes Ill-V; fifth toe much longer than third; when hind limbs flexed against axis of body, heels touching or barely overlapping; hind limbs short, shank 39.5^4.6% (r = 42.2 ± 0.4) SVL. Dorsum pale brown with small brown spots and nearly black interorbital bar; snout paler than rest of dorsum; limbs pale brown with dark brown bars and white flecks on forearms; supratympanic stripe and spot at anterior corner of eye dark brown; subocular spot white with brown border; all ventral surfaces white except for brown stippling along jaws; posterior surfaces of thighs pale brown; disc covers and outer digits dark brown; innermost digits cream; white line separating disc cover from brown on top of digit. Coloration in life. — A color photograph of an individual from Isla Gorgona, Colombia, reveals a yellowish-tan dorsum and yellow flanks; reddish- brown dorsal markings consist of an interorbital bar, narrow canthal stripe, broader postorbital stripe, short, irregular marks in the scapular region, and transverse dots on the shanks. The iris is golden- bronze with a reddish-brown horizontal streak. Natural history. — Eleuthewdactylus gitlaris inhabits lowland tropical rainforest. Kenneth Miyata obtained two specimens on vegetation along a small stream at San Miguel, ProvinciaEsmeraldas, on the night of 20 June 1977. Distribution. — This species occurs at low el- evations in the humid tropical regime in northwest- ern Ecuador and Colombia northward to at least Pizano (Departamento Choco) (Fig. 33). The only record above 400 m is the purported type local ity of Hyloxalus huigrae at 1 200 m in the dry subtropical regime. Remarks. — Dunn ( 1 942 ) combined Eleuthew- dactylus gularis with E. diastema (Cope). Cochran and Coin (1970) did not reference Dunn's paper and reported E. gularis and E. diastema from Colombia. They stated (1970:374) that the two species could be distinguished by the "disc of third finger and of fourth toe two or three times the size of tympanum" in E. diastema and "not quite as large as tympanum" in E. gularis. However, the discs in E. diastema never are that large, even in the specimens that they reported and described. Lynch (1976b) suggested that E. gularis differs from E. diastema in that the former has papillae on the tips of [some] discs; he used this observation to remove Hylodes huigrae Fowler from the synonymy of E. diastema and to allocate it to the synonymy of £. gularis. However, not all specimens from the lowlands of Colombia and none of those from Ecuador have discs with papillae as long as those illustrated by Lynch( 1976b). Specimens from Ecuador and south- ern Colombia have papillae only on Toes II-IV, usually in the form of rounded knobs at the apex of the pad on the underside of the disc cover, but sometimes they are more prominent. Frogs of this group from the Atrato and San Juan drainages in Chocoan Colombia have larger papillae on the toes (some on fingers as well) and many of these frogs are small in contrast to those from Ecuador and Isla Gorgona. There are probably three species in- volved in western Colombia but only one in Ecua- dor. In E. gularis, the digital tips are somewhat pointed, and casual observation (not under a dis- secting microscope ) would lead one to consider the discs to be papillate; the holotypes oiE. gularis and Hyloxalus huigrae do not have papillae at the tips of the digits. On the strength of the presence of short papillae 102 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 on some toes and fleshy basal webbing between the toes, we recognize E. i>iil(irls as distinct from E. diastema. The latter is known as far east as extreme eastern Panama and is expected to be found in Colombia. Along the Pacific coast, but apparently not inland, E. gidaris ranges as far north as Pizarro in Departamento Choco, Colombia. Two unnamed species occur in Colombia and Panama. One of those was illustrated by Lynch ( 1 976b) as E. i>ularis. but it is a smaller frog with prominent papillae on the finger tips but not the toe tips. Eleuthewdactylus gidaris is the same size as E. diastema but may be more closely related to E. vocalis Taylor, a small species with pointed disc covers. Eleuthewdactylus Immiotae Flores Plate 4 Eleutherodactylus hamiotae Flores, 1993:427. — Holo- type: MCZ 98027. adult female, from 13.1 km NW None, 2140 m. Provincia Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) surdus group having ( 1 ) skin on dorsum shagreen with flat warts poste- riorly and slightly more pungent pustules on flanks, that on venter areolate; discoidal fold poorly de- fined; dorsolateral folds absent; postocular folds pustular; fold lower on flank extending to level of sacrum, becoming lost posteriorly amidst pustules; (2) tympanic membrane and tympanic annulus absent; (3) snout short, rounded in dorsal view, rounded to slightly sloping in profile; (4) upper eyelid bearing flattened warts, as wide as or slightly broader than lOD; cranial crests absent; (5 ) vomer- ine odontophores oval in males, triangular in fe- males; (6) males lacking vocal slits; thumb of males swollen, lacking nuptial pads; testes white; (7) first finger shorter than second; thumb lacking disc; discs on outer fingers nearly twice width of digit, all bearing ventral pads; (8) fingers bearing fleshy lateral keels; (9) ulnar tubercles absent (Flores, 1993:430, reported "indistinct ulnar tu- bercles"); ( 10) heel and outer edge of tarsus bearing minute tubercles; inner edge of tarsus bearing thick- ened area along distal half; (11) inner metatarsal tubercle twice as long as wide; outer metatarsal tubercle not apparent (Flores ( 1993:430, reported outer metatarsal tubercle as "variably present, minute"); supernumerary plantar tubercles absent; (12) toes having lateral fringes and basal webbing; webbing formula for outer toes IV 4^ — 2/2 V; fifth toe longer than third, not reaching distal subarticular tubercle on Toe IV; ( 1 3) dorsum brown to tan with scattered dark areas; facial markings poorly de- fined; venter cream with dense brown stippling; posterior surfaces of thighs brown with cream spots (or uniform brown, USNM 239843); (14) SVL in males 28.2-30.9 mm, in females 35.7-36.2 mm. Eleutherodactylus hamiotae is most similar to E. duellmani, but it is smaller with less well devel- oped basal webbing between the toes, a more sloping snout (truncate in E. duellmani), less obvi- ous dorsolateral folds, more robust fingers, and no tubercles on the heel and outer edge of the tarsus. Breeding males of E. duellmani have nuptial pads (absent in E. hamiotae). Description. — The type description (Flores, 1993) is complete. This species is represented by few specimens; we provide measurements for the second known male (USNM 239843): SVL 30.9, shank 13.4, HW 1 1.3, head length 10.0, chord of head length 10.5, upper eyelid width 2.9, lOD 2.3, eye length 4.0, E-N 2.6. Coloration in life. — Based on field notes of Kenneth Miyata, Flores (1993:430-31) reported colors in life of the two paratypes, as follow. "MCZ 97486: 'Dorsum dark greenish olive mottled with dark brown. Limbs dark brown barred with black. Venter gray with dark gray reticulations. Iris dark brown.' MCZ 98052: 'Dorsum clay colored mottled with cinnamon brown. Venter translucent smoke gray. Iris deep coppery brown'." Natural history. — The type locality is a well- watered roadcut; the type series was collected on the dripping rock face (Flores, 1993). James A. Peters collected a single specimen ( USNM 239843 ) at a locality " 1 2 km W Nono (Nono-Nanegal road)" on 26 October 1958. Peters ' field notes suggest that he collected at the same site visited 19 yr later by Kenneth Miyata, who collected the type series. Distribution. — Eleutherodactylus hamiotae is known only from the vicinity of the type locality in cloud forest at an elevation of 2140 m in the humid temperate regime on the western slopes of the Andes in Ecuador (Fig. 27). ELEIJTHERODACTYLIIS IN WESTERN ECUADOR 103 Remarks. — The distribution of Eleuthero- dactylus duellmcmi, which occurs at elevations of 1550-2500 m, encompasses that of E. hamiotoe. The closest record of E. duellmani to that of E. hamiotae is 9 km SE Tandayapa (2150 m ). which is only about 6 km (airline) from the type locality of E. hamiotae. Eleuthewdactylus hamiotae seems to be more closely related to E. dueUmani than to either of the other species in the E. surdus group (£. sobetes and E. surdus). Eleuthewdactylus hectus Lynch and Burrowes Plate 8 Eleutherodactylus hectus Lynch and Burrowes, 1990:11.— Holotype: IND-AN 1947. adult female, from Reserva Natural La Planada. 1 780 m. Municipio de Ricaurte. Departamento Nariiio. Colombia. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) myersi group having (1) skin on dorsum tuberculate to granular, that on venter areolate; discoidal fold absent; dorsolateral folds prominent; (2) tympanic membrane and tym- panic annulus prominent, y?-'/: length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid bearing two or three subconical tubercles, narrower than lOD; cranial crests absent; (5) vomerine odontophores oval in outline; (6) males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs small; (8) fingers bearing lateral fringes; (9) ulnar tubercles present; (10) heel and outer edge of tarsus bearing small tubercles; inner edge of tarsus bear- ing small, foldlike tubercle; (II) inner metatarsal tubercle oval, 4x conical outer metatarsal tubercle; supernumerary plantar tubercles numerous, minute; (12) toes bearing narrow lateral fringes and lan- ceolate discs; webbing absent; fifth toe slightly longer than third; (13) dorsum gray to brown with darker brown markings; venter pale brown with vague blotches; posterior surfaces of thighs brown with or without cream spots; (14) SVL in males 13.6-16.8 mm, in females 19.4-22.5 mm. Eleutherodactylus hectus is most similar to frogs included in the E. pyrrhomerus group by Lynch (1976c), but it differs from each of them by having dorsolateral folds and larger digital discs, although some specimens of £. leoni may have dorsolateral folds (fide L. A. Coloma, in litt.). In addition, E. hectus lacks red or orange on the concealed sur- faces of the hind limbs and/or in the groin; instead, E. hectus has white spots. Description. — The type description by Lynch and Bunowes (1990) is complete, but see Re- marks. Coloration in life. — According to Lynch and BuiTowes (1990), the dorsum is brown, tan, or reddish brown with creamy white, black, or brown markings. An orange-tan blotch on the head or a black interorbital bar usually are present. The limbs are brown or tan with darker brown or black trans- verse bars. The groin and hidden surfaces of the hind limbs are brown with white flecks, which may be clumped into spots, and the flanks are brown with varying degrees of creamy white mottling. The venter is yellow-tan with brown flecks or brown with creamy-white flecks. The iris is dull brown to greenish or bluish gray with a median, horizontal red streak. Natural history. — In contrast to other Eleutherodactylus on the western slopes of the Andes in Ecuador, this species is diurnal in cloud forest. During the day, individuals were observed in recesses formed by buttresses and roots, and hopping in the leaf litter. Because of the cryptic coloration, stationary individuals were difficult to detect. At night, the frogs were observed sleeping on low herbaceous vegetation, a habit reminiscent of many dendrobatids. Distribution. — Eleutherodactylus hectus is known from two localities at intermediate eleva- tions (1200-1780 m) in cloud forests in the humid tropical regime on the western slopes of the Andes in extreme southern Colombia (type locality) and northwestern Ecuador ("above" Lita) (Fig. 31 ). Remarks. — Specimens from Ecuador and re- cently collected topotypes (UVC) do not have the inner metatarsal tubercles projecting medially; this condition, cited as diagnostic by Lynch and Burrowes ( 1990), probably is an artifact of preser- vation. Eleutherodactylus hectus probably is most closely related to E. leoni, from which it differs in coloration and the presence of lanceolate discs on the toes. The limited distributional data for E. 104 UNIV. KANSAS NAT. HIST. MUS. SPEC. PIJBL. NO. 23 hectus suggest that this species may be restricted to elevations below the altitudinal range ( 1 96()-34()() m) of E. leoni. Eleuthcwdactylus Iwlonotiis Lynch Amblyphrynus hekmotiis Lynch, 1 ^)l>iv. 1 9. — Holotype: BM 1970: 178. adult female, from the Rio Pitzara ( = Rfo Pitsara). Provincia Pichincha. Ecuador. Elentlu'wdactyhis Iwlonotiis — Lynch. 1 98 1 h: 3 1 8. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) sidcatus group hav- ing (1) skin on dorsum with heterogeneous tu- bercles, that on venter coarsely areolate; discoidal fold not apparent; dorsolateral folds absent; (2) tympanic membrane and tympanic annuius promi- nent, higher than long. -A-Vi length of eye; (3) snout subacuminate in dorsal view, truncate in protile; head broad (HW ± 50% SVL); (4) upper eyelid bearing flattened warts, much broader than lOD; cranial crests low, parasagittal; (5) vomerine odontophores arched; (6) males unknown; (7) first finger longer than second; discs absent; (8) fingers bearing weak lateral keels; (9) ulnar tubercles present; (10) heel lacking tubercles; outer edge of tarsus bearing row of small tubercles; inner edge of tarsus lacking fold or tubercles; (11) inner metatar- sal tubercle weakly compressed, 2-3x round outer metatarsal tubercle; supernumerary plantar tu- bercles absent; (12) toes bearing lateral fringes; tips of toes not expanded; webbing absent; fifth toe slightly shorter than third; (13) dorsum brown with indefinite black markings associated with tubercles and folds; venter cream with large brown blotches; posterior surfaces of thighs brown with black re- ticulation; (14) SVL in males unknown, in one female 69.6 mm. Eleutherodactylus helonotus is the only broad- headed species of Eleutherodactylus in western Ecuador having areolate skin on the venter. We expect that males and juvenile females, when dis- covered, will have tuberculate skin on the dorsum comparable to that in E. anomalus, cerastes, and necerus. but the two large females now known have low warts on the dorsum and appear to be smoother skinned. Unlike E. anomalus, the digital tips in E. helonotus are not expanded, and the toes lack any trace of webbing. The upper eyelid does not bear elongate tubercles as in E. cerastes. Eleuthero- dactylus necerus differs from E. helonotus by hav- ing the posterior surfaces of the thighs black with pale spots, an unpatterned venter, and numerous short ridges on the dorsum. Coloration in life. — Unknown. Natural history. — This species is known from two females. The holotype is gravid, and the paratype is a slightly smaller (60.6 mm SVL) female having moderate oviductal convolutions and small ovarian eggs. The paratype was on a mule trail in cloud forest. The short limbs, narrow digital tips, and bulk indicate that it is a terrestrial species. Distribution. — One individual is from the Ri'o Pitsara. and the other is from the immediate envi- rons of Mindo (1410 m), both in Provincia Pichincha on the western slopes of the Andes in Ecuador ( Fig. 33 ). Presumably, this species has a limited distribu- tion at intermediate elevations in cloud forest in the humid subtropical regime. Remarks. — When first described (Lynch, 1975a), E. helonotus was placed in the genus Amblyphrynus because no ventral pads could be seen on the tips of the digits. The two species of Amblyphrynus were considered by Lynch (1975a) as somehow being derived from the broad-headed group of Eleutherodactylus; such a classification is incompatible with a cladistic interpretation. It was not until freshly collected specimens of Ambly- phrynus ingeri became available that Lynch (1981b) realized that ventral pads were present in that species and discontinued recognizing the genus Amblyphrynus. No additional material of E. helonotus has become available, but it too probably will be found to have traces of ventral pads. What little anatomical data are available place E. helonotus securely within the E. sulcatus group. More precise placement is not possible, although E. helonotus does have areolate skin on the venter, as do E. ingeri, ruizi, and sulcatus, in contrast to smooth skin on the venter in E. cadenai, cerastes, and E. cornutus. Eleutherodactylus illotus sp. nov. Plate 2 Holotype. — KU 1 6588 1 . adult female, one of a series collected 3.5 km NE (by road) Mindo, 1540 ELEUTHERODACTYLUS IN WESTERN ECUADOR 105 m (00°{)2'53" S, 78°46"2{)" W), Provincia Pichincha, Ecuador, on 7 April 1975 by W. E. Duellman. Paratypes.— KU 165882-83, USNM 239722- 33. (See Appendix I for localities.) Diagnosis. — A member of the Eleuthew- dactyhis (Eleutherodactylus) conspicillatus group having ( 1 ) skin on dorsum shagreen with scattered tubercles, that on venter smooth; discoidal fold present; dorsolateral folds pustular, complete; (2) tympanic membrane and tympanic annulus promi- nent, its length '/^--/s length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid bearing tubercles, about as wide as lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males lack- ing vocal slits and nuptial pads; (7) first finger longer than second; outer fingers bearing small discs about twice width of digit; (8) fingers bearing lateral keels; (9) ulnar tubercles absent; (10) heel bearing small subconical tubercle; outer edge of tarsus bearing minute tubercles; one inner tarsal tubercle; (11) inner metatarsal tubercle elongate, 4-6x conical outer metatarsal tubercle; supernu- merary plantar tubercles few; (12) toes bearing narrow lateral fringes; webbing absent; fifth toe slightly longer than third; (13) dorsum brown with darker brown markings; venter cream with brown wash or mottling; groin and posterior surfaces of thighs brown without pale flecks or spots; ( 14) SVL in males 25.9-29.3 mm, in seven females 38.3- 44.6 mm (f = 41.0). Eleutherodactylus illotus is similar to E. achatinus, but differs by having a small tubercle on the heel and brown coloration on the venter; also the former lacks vocal slits and nuptial pads (both present in E. achatinus). Eleutherodactylus illotus is similar to. but differs from. E. thectoptermis from western Andean Colombia (Lynch, 1975b) by hav- ing a small tubercle on the heel (rather than a calcar), larger tympanic annulus. and dorsolateral folds, and by lacking white spots on the posterior surfaces of the thighs. Description. — {n = 10; 2 males. 8 females) Head broader than body (except in gravid females), broader than long; HW 42.3 and 44.8% SVL in males, 40.2^3.1% (r = 41.9) SVL in females; snout relatively long, subacuminate in dorsal view. rounded in profile; nostrils slightly protuberant, directed dorsolaterally: E-N 92.5 and 95.6% SVL in males. 90.3-103.8% (.v = 100.0) length of eye; canthus rostralis angular, prominent, sinuous; lo- real region slightly concave, sloping abruptly to lips; lips not flared; posterior portion of upper eyelid bearing low to subconical tubercles (anterior terminus of postocular fold), width of upper eyelid 88.9 and 1 13.3% lOD in males. 83.9-121.9% (.v = 104.5) lOD in females; interorbital region flat; cranial crests absent; temporal region nearly verti- cal; supratympanic fold curving posterior to tym- panic annulus; tympanic membrane and tympanic annulus prominent, round in males, slightly higher than long in females, separated from eye by dis- tance equal to length of tympanic annulus; length of tympanic annulus 35.0 and 40.0% length of eye in males, 32.3-^0.5% (v = 36.8) length of eye in females; two subconical postrictal tubercles. Choanae approximately triangular (apex posteriad), not concealed by palatal shelf of maxillary arch; vomerine odontophores posteromedian to choanae, elevated and triangular in outline, each slightly smaller than a choana, separated medially by dis- tance equal to one-half to one width of odontophore, bearing 8- 1 0 teeth in transverse row; tongue longer than wide, its posterior border not notched, poste- rior -A- A not adherent to floor of mouth; males lacking vocal slits and nuptial pads. Skin on dorsum of head shagreen except for round tubercles on upper eyelids, on occipital W (postocular folds), immediately adjacent to supratympanic fold, and immediately anterior to tympanic annulus; skin on dorsum of finely shagreen, becoming more coarse posteriorly; dor- solateral folds low, granular, extending to level of groin; skin on venter smooth; discoidal fold well anteriad to groin; cloacal sheath and tubercles absent; ulnar tubercles absent. Thenar tubercle oval, much smaller than oval palmar tubercle; supernumerary palmar tubercles not evident; subarticulartubercles round, notconical; first finger longer than second; fingers bearing faint lateral keels; outer fingers bearing expanded, round to subtruncate discs about twice as wide as digit proximal to disc; inner digits bearing much nar- rower discs, only slightly wider than digit; all fingers bearing pads on digital tips; pads defined by 106 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 circumferential grooves; nuptial pads absent in males. Skin on dorsal surfaces of hind limbs finely shagreen; heel bearing subconical tubercle; up to six minute tubercles along outer border of tarsus (at level of color change between black subtarsal color and paler color on dorsal and lateral surfaces); inner edge of tarsus bearing round tubercle on distal third of tarsus (some specimens with smaller tubercle between round one and inner metatarsal tubercle, but no specimen having tarsal fold); inner metatarsal tubercle thrice as long as wide, about 6¥ subconical outer metatarsal tubercle (longer than wide); supernumerary plantar tubercles few, low, at bases of Toes II-IV; subarticular tubercles round to longer than wide, not conical; digits bearing nar- row lateral fringes; webbing absent; toe discs ex- panded (largest discs about twice width of digit proximal to disc); fifth toe slightly longer than third; tip of Toe V extending to distal edge of penultimate subarticular tubercle on Toe IV. tip of Toe III extending to proximal border of penultimate subarticular tubercle on Toe IV; when hind limbs flexed against axis of body, heels broadly overlap- ping; hind limbs long, shank 57.3 and 60.6% SVL in males, 54.3-61.3% (.r = 57.3) SVL in females. Dorsum brown with darker brown interorbital bar, dorsal chevrons, and inguinal spots; dark brown border below dorsolateral folds grading into brown spots on flanks; canthal stripe and labial bars brown; supratympanic stripe black; cloacal patch black; limbs pale brown with darker brown bars, those on shanks oblique. naiTower than interspaces; under- sides of forearms and tarsi black; groin, anterior and posterior surfaces of thighs, and concealed surfaces of shanks brown without pale flecks; throat pale brown with vague cream areas; belly cream with brown reticulation. Measurements of holotype: SVL 38.3. shank 22.5. HW 16.5, head length 16.0. eyelid width 3.8. lOD 4.1, tympanic annulus length 2.0. eye length 5.3. E-N 5.5. Coloration in life. — The only color notes for living individuals are for KU 165881-82 from 3.5 km NE Mindo, Provincia, Pichincha. as follow. Dorsum brown with dark brown markings; throat and belly bluish white with brown mottling; other ventral surfaces dull gray; iris dark bronze with black reticulations (WED, 7 April 1975). Natural history. — This species is an inhabitant of cloud forest. Limited data on habitat reveal that the species is found on low vegetation at night. Distribution. — For this species, explicit elevational data are available for only five collec- tions (1380, 1540, 1770, 1780, and 2560 m). The latter is the only record from the humid temperate regime; other localities lie at elevations in the humid tropical regime on the Pacific slopes of the Andes from Provincia Carchi southward to Provincia Pichincha, Ecuador (Fig. 37). Etymology. — The specific name is a Latin ad- jective meaning "unwashed" or "dirty"; the name is applied in allusion to the dark marbling on the belly. Remarks. — We are unaware of any synapo- morphy to link E. illotiis and E. thectoptemus, but suspect that the two are sister species. Eleuthero- dactylus thectoptemus occurs on the Pacific slopes of the Andes in Colombia as far south as Departamento Cauca. Eleutherodactylus labiosus Lynch, Ruiz, and Ardila Plate 3 Eleutherodactylus labiosus Lynch, Rufz, and Ardila, 1994:29.— Holotype: KU 131612, juvenile female, from La Palma, 920 m. Provincia Pichincha, Ecua- dor, on 8 August 1970 by John D. Lynch. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) cerasinus group hav- ing ( 1 ) skin on dorsum shagreen with low occipital folds, that on venter areolate; discoidal fold well anteriad to groin; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus promi- nent, higher than long, its length Va-A length of eye; (3) snout subovoid in dorsal view, rounded in profile, lips flared in adult females; (4) upper eyelid bearing one conical tubercle, broader than lOD; cranial crests low in females, absent in males; (5) vomerine odontophores triangular in outline; (6) males having vocal slits; nuptial pads absent; (7) first finger slightly shorter than second; discs broad on outer fingers, usually emarginate; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel bearing small, conical tubercle; outer ELEUTHERODACTYLUS IN WESTERN ECUADOR 107 n 80 0 20 40 60 80 100 Kilometers 30(1 111 2(K)() 111 I I >4()(K)ni I ,,| Perniancnl A E. Hiatus • E. latidiscus -1 ■ E. leoni Fig. 37. Distribution of three species oi Eleuthero- dactylus in western Ecuador. edge of tarsus bearing row of small tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle oval, 4x round outer metatarsal tubercle; super- numerary tubercles only at bases of toes; (12) toes lacking lateral fringes; webbing absent; fifth toe longer than third; (13) dorsum brown with darker markings; venter cream with brown flecks; poste- rior surfaces of thighs brown with cream flecks; (14) SVL in males 35.4-50.8 mm, in females 48.5- 52.3 mm. Eleuthewdactylus labiosus is most similar to E. crenunguis and E. ocellatus, but differs from both by having a conical tubercle on the upper eyelid. Furthermore, the first finger is longer than the second in E. crenunguis, and E. ocellatus lacks a tubercle on the heel. Superficially, E. labiosus is like the smaller E. tenebrionis, which differs by having smooth skin on the dorsum and lacks flared lips in adult females. Description. — The original description (Lynch et al., 1994) is complete. Coloration in life. — The dorsum is various shades of brown from olive-brown to reddish brown with indistinct slightly darker markings. The de- tailed description of holotype is, as follows. KU 131612 from La Palma, Provincia Pichincha: Dor- sal surfaces of body and limbs brown with a rusty cast, and greenish tint on shanks; body with ill- defined darker brown markings; labial stripe dull yellow; posterior surfaces of thighs dull brown; groin, anterior surfaces of thighs, and lower flanks soft yellow with brown marbling; iris bright red- dish-rust in upper quarter, nearly gray elsewhere (JDL, 8 August 1970). Natural history. — Eleutheroclactylus labiosus is primarily an inhabitant of lowland rainforest, where individuals perch on broad leaves or branches along streams at night. This seems to be an uncom- mon species. Our field work has yielded few speci- mens, usually no more than one or two individuals at a given locality, but the late Kenneth Miyata collected 14 specimens, mostly juveniles, at Tinalandia, Provincia Pichincha. Distribution. — This species occurs at eleva- tions of 150-1500 m on the Pacific versant of Colombia (south of Buenaventura) and Ecuador. Only one individual is known from an elevation of more than 1000 m (Fig. 33). Three of the Ecuador- ian localities are in the humid tropical regime, and 14 are in the the humid subtropical regime. Remarks. — For several years JDL confused this species with Eleuthewdactylus latidiscus, a smaller species with lateral fringes on the digits and a much longer fifth toe. The large digital discs are consistent with an arboreal existence. The arboreal habit of this species is important in establishing the fact that variation in the length of the fifth toe does not reflect degree of arboreal ity. Lynch et al. ( 1 994) suggested that E. labiosus is replaced at higher elevations by E. crenunguis; that seems to be a fair description of the distributions of the two species, but there is no evidence of interspecific competi- tion. Some large females have thin, unconvoluted oviducts; Lynch et al. (1994) suggested that fe- males may undergo regression of the oviducts following reproduction. Eleuthewdactylus laticlavius Lynch and Burrowes Plate 5 Eleutherodacrylus laticlavius Lynch and Burrowes, 1990:14.— Holotype: IND-AN 1564. adult female, from Reserva Natural La Planada, 1 780 m, Municipio de Ricaurte, Departamento Narino, Colombia. 108 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Diagnosis. — A menibei" of the Eleiiihero- dactylus (Eleuthewdactylus) unisfrii>atus group having ( 1 ) skin on dorsum smooth with nonconical tubercles on dorsum posterior to sacrum and along dorsolateral stripes, that on venter areolate; discoi- dal fold just anteriad to groin: dorsolateral folds low; (2) tympanic membrane and tympanic annu- lus prominent, its length V4-A length of eye: (3) snout rounded in dorsal view and in prohle: (4) upper eyelid bearing small tubercle, nanower than lOD: cranial crests absent: (5) vomerine odontophores triangular in outline: (6) males lack- ing vocal slits: nuptial pads absent: (7) hrst finger shorter than second; discs broad: (8) fingers bear- ing lateral keels: (9) ulnar tubercles absent: (10) heel bearing small, conical tubercle; outer edge of tarsus bearing row of small tubercles; inner edge of tarsus bearing short tuberclelike fold; (11) inner metatarsal tubercle oval. 6x elongate outer meta- tarsal tubercle: supernumerary tubercles numer- ous, low; ( 12) toes bearing lateral fringes: webbing absent: fifth toe much longer than third: (13) dor- sum either brown with broad pale dorsolateral stripes or brown with darker dorsolateral stripes and brown flanks; venter cream with brown mot- tling: posterior surfaces of thighs brown with cream spots; (14) SVL in males 22.5-26.3 mm, in females 35.0-42.9 mm. Eleutherodactylus laticlavius is most similar to the lowland E. latidiscus. The two species can be distinguished by E. laticlavius having smoother skin on the dorsum and narrower digital discs, and by lacking nuptial pads in males. Description. — The original description by Lynch and Bunowes ( 1990) is complete. Coloration in life. — According to Lynch and Burrowes (1990). living individuals of the striped morph have a chocolate-brown dorsum with wide, yellow canthal-dorsolateral stripes. The other moiph has a tan dorsum with thin, reddish-brown canthal- dorsolateral lines and a narrower, creamy-yellow middorsal line. In both morphs, the dorsal surfaces of the limbs are tan with brown bars edged with creamy white; the flanks and posterior surfaces of the thighs are yellow with brown marbling, or brown to reddish brown with whitish blotches. The venter is white to tan with pink, orange, or greenish yellow tint and brown flecks. The iris is golden- copper with fine black reticulations and a median, horizontal red streak. Natural history. — This species has been found only on low vegetation near streams at night in cloud forest. Distribution. — Eleutherodactylus laticlavius is known from five localities at elevations of 1200- 2565 m on the western flanks of the Andes in extreme southwestern Colombia and in provincias Carchi and Pichincha in northern Ecuador, where it occurs in the humid subtropical and humid temper- ate regimes (Fig. 35). Remarks. — Lynch and Burrowes (1990) re- ported 1 0 specimens from the type locality. Subse- quently, E. laticlavius was found at three localities in Ecuador, but essentially no information is avail- able on the biology of the species. At the time of the original description. E. laticlavius was not com- pared with E. latidiscus, which may be its closest relative. This suggestion is based solely on phe- netic similarity, which includes sharing pattern polymorphs. Eleutherodactylus latidiscus (Boulenger) Plate 6 Hylodes latidiscus Boulenger. 1 898: 1 2 1 . — Syntypes: BM 1947.2.15.66-67. adult females, from Cachabe, Provincia Esmeraldas, Ecuador. Eleutherodactylus latidiscus — Dunn, 1 933:68. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) urustrigatus group having (1) skin on dorsum tuberculate in males, shagreen with scattered tubercles in females, that on venter areolate; discoidal fold prominent, well anteriad to groin; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus promi- nent, round, its length V4--/5 length of eye; (3) snout subacuminate to rounded in dorsal view, rounded in profile; lips flared in adult females; (4) upper eyelid bearing subconical tubercle, broader than lOD; cranial crests absent: (5) vomerine odontophores triangular in outline: (6) males lack- ing vocal slits: nuptial pads present: (7) first finger slightly shorter than second: discs broad on Fingers Il-lV; (8) fingers bearing lateral fringes: (9) ulnar tubercles absent: ( 10) heel bearing small tubercle; ELEUTHERODACTYLVS IN WESTERN ECUADOR 109 outer edge of tarsus bearing row ofsmall tubercles; inner edge of tarsus bearing thick tuberclelike fold; (11) inner metatarsal tubercle oval, 4x round outer metatarsal tubercle; supernumerary plantar tu- bercles indistinct; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; (13) dorsum pattern polymorphic, usually with pale lines along outer edges of W-shaped occipital mark; venter white with some brown stippling; posterior surfaces of thighs brown with small cream spots; ( 14) SVL in males 2 1 .9-25.9 mm, in females 35.2-53.4 mm. Among species in western Ecuador, E. latidiscus is most similar to E. laticlavius, which has smaller digital discs and less coarse skin on the dorsum and lacks nuptial pads in males. Description. — The species was redescribed thoroughly by Lynch et al. (1994). Coloration in life. — The coloration in life is highly variable as attested to by the following descriptions. KU ] 09059 from Santo Domingo de los Colorados, Provincia, Pichincha: Dorsum olive- green with brown markings; throat bright orange; rest of venter brown, darkest on limbs; flanks and anterior and posterior surfaces of thighs with bluish gray flecks; iris pale bronze with black reticula- tions and median, horizontal, reddish-brown streak (WED, 29 March 1967). KU 120222-26 from Santo Domingo de los Colorados, Provincia, Pichincha: By night, dor- sum largely gray. By day, dorsum dark brown with black spots; paler brown on flanks with green wash and brown markings; limbs dark brown with black markings; one individual with reddish-brown head cap; venter and posterior surfaces of thighs dusky black; iris pale golden-bronze with red horizontal stripe and black reticulation (JDL, 3 August 1968). KU 120232-33 from Santo Domingo de los Colorados, Provincia, Pichincha: Dorsum pale rust-brown with dark brown markings; flanks with slight green wash and pale brown markings; limbs pale brown with dark brown bands; venter dirty yellow mottled with greenish brown in one indi- vidual, dark gray in the other; iris bright bronze above, reddish brown below with reddish horizon- tal streak (JDL, 4 August 1968). KU 141957-61 from Rio Baha, Provincia Pichincha: At night, dorsum pale greenish-tan; by day, dorsum olive-brown to reddish brown with darker brown markings; venter dull yellow to creamy white with brown to black markings; iris bronze with median, horizontal red streak (WED, 5 July 1971). KU 165468-70 from Estacion Biologica Rio Palenque, Provincia Los Ri'os: Dorsum tan with olive-brown markings; groin reddish brown; throat and inner two fingers yellow; rest of venter gray; iris dull bronze with black reticulations and hori- zontal red-brown streak (WED, 30 March, 1975). KU 165470 from Estacion Biologica Rio Palenque, Provincia Los Rfos: At night, tan with creamy-yellow dorsolateral stripes; venter pale gray; by day, dorsum brown and venter dark gray (WED. 31 March 1975). Natural history. — Most individuals have been observed at night perched on low vegetation or on tree trunks in forest. By day. this species has been found in leaf litter on the forest floor and in brome- liads; one individual was in the axil of a palm frond 5 m above the ground. An amplectant pair collected 5 km W La Florida, Provincia Pichincha, deposited eggs in a plastic bag on the night of 28 December 1989. The female (KU 218017) has a SVL of 42.4 mm; the loosely adherent clutch contained 40 eggs with capsules about 4.5 mm in diameter and ova about 2.4 mm in diameter. Distribution. — This species occurs in lowland tropical rainforest and humid lower montane forest at elevations of 20-1 230 m in northwestern Ecua- dor and western Colombia as far north as the Rio San Juan (Fig. 37). The highest reported elevation for the species in Ecuador is 830 m in the humid subtropical regime, where seven other localities are recorded; four localities are in the humid tropi- cal regime, and one is in the dry tropical regime. Remarks. — Eleutherodactylus latidiscus most likely is related to E. laticlavius and E. cruentus. The three species are approximately the same size and have much the same shape, but they differ in disc size, skin texture, relative sizes of the tubercles on the heel and outer edge of the tarsus, distinctness of the tympanic annulus. and color pattern on the concealed surfaces of the hind limbs. In Ecuador and southern Colombia, E. laticlavius occurs at higher elevations than E. latidiscus; at present. 110 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 collections arc inadequate to a.sses.s the nature of the contact between E. latidiscus and E. crueutiis in northwestern Colombia (Lynch et al.. 1994). EleHtlwrodactylus leoni Lynch Plate 8 Eli'iitlwrodactyliis leoni Lynch. 1 976c:3 1 3. — Holotype: KU 130870, adult female, from the north slope of Nudo de Mojanda, 3400 m, Provineia Imbabura, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus {Eleuthewdactylus) myersi group having (1) skin on dorsum shagreen with numerous short ridges, that on venter areolate; discoidal fold ab- sent; dorsolateral folds absent; (2) tympanic mem- brane absent but tympanic annulus visible through skin, higher than long, its length V^--A length of eye; (3 ) snout subacuminate in dorsal view, semitruncate in profile; (4) upper eyelid bearing many subconical tubercles, narrower than lOD; cranial crests ab- sent; (5) vomerine odontophores low, oblique; (6) males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs narrow; (8) fingers lacking lateral fringes; (9) ulnar tubercles indistinct; (10) heel lacking small tubercle; outer edge of tarsus bearing small tubercles; inner edge of tarsus bearing ridgelike tubercle; (11) inner metatarsal tubercle elongate, 2x round outer meta- tarsal tubercle; supernumerary plantar tubercles small; (12) toes bearing narrow lateral fringes; webbing absent; fifth toe slightly longer than third; (13) dorsum brown with darker brown markings; venter gray to brown with cream spots; white spots and/or colorless areas (salinon or bright red in life) in axilla, groin, and on thighs; ( 14) SVL in males 14.8-18.3 mm, in females 19.7-25.0 mm. Eleudiewdactylus leoni is most similar to E. floridus, hectus, and pyrrhomenis, but differs by having black testes (white in others) and red spots in the axilla and groin and on the concealed sur- faces of the hind limbs; these areas are white in E. hectus. Eleuthewdactylus hectus, leoni, and pyrrhomenis have narrower digital discs that E. floridus. Description. — The original description ( Lynch. 1976b) is adequate. Coloration in life. — The known variation is covered adequately in the following descriptions. KU 130871-72 from the north slope Nudo de Mojanda, Provineia Imbabura: Dorsum of body and limbs brown or reddish brown with gray mark- ings; lips barred cream and brown; vocal sac yel- low; rest of venter gray with a few yellow spots or washed with salmon; groin and lower flank with yellow spots; posterior surface of thigh gray with yellow spots in one individual, salmon-red in the other; iris powder-blue with black punctations and black streak (JDL, 2 August 1970). KU 165897-98 from 14 km W Chiriboga, Provineia Pichincha: Dorsum grayish brown with black and orange markings or brown with dark brown markings; venter dull gray with bluish- or silver-white flecks; groin and hidden surfaces of hand limbs orange-red with black markings; iris pale green with median, horizontal, red stripe. KU 117320-^0 from 14 km SE Maldonado, Provineia Carchi: Dorsum brown or rust-brown or washed with green; groin and concealed limbs red in females, brown or cream in males; venter of females dull cream heavily flecked with brown, of males bluish with brown flecks or reticulations; throat dusted with brown; iris blue-gray with red horizontal streak (JDL, 31 May 1977). Natural history. — Most specimens of Eleu- therodactylus leoni have been collected by day. Adults are relatively common under rocks and logs in paramo and upper cloud forest, but individuals have been found on low vegetation (<30 cm) in cloud forest and in paramo at night. Males were calling from the ground by day on the north slope of Nudo de Mojanda, Provineia Imbabura, on 2 August 1970 and 5 km W La Gruel, Provineia Carchi, on 26 February 1984. The call is a distinct "tink." Distribution. — Although this species is re- stricted geographically to a relatively small area of the Andes in northern Ecuador and southern Co- lombia, it occurs at elevations of 1960-3400 m. It seems to have two disjunct populations — one on the eastern slopes ofthe Andes in Provineia Carchi, Ecuador, and Departamento Putumayo. Colombia, and the second on the western slopes of the Andes in provincias Carchi and Imbabura, Ecuador (Fig. 37). Ofthe nine known localities in western Ecua- ELEUTHERODACTYLUS IN WESTERN ECUADOR II dor, six are in the humid temperate regime and three in the humid subtemperate regime. Remarks. — Specimens from cloud forests are indistinguishable from those from paramo. The only efforts to determine relationships among the species of the E. myersi group are rather unsatisfac- tory. Phenetically, E. leoni is most similar to E. hectiis and E. repens. Eleuthewdactylus longirosths (Boulenger) Plate 1 Hylodes longirostris Boulenger, 1898:120. — Syntypes: BM 1947.2.15.56-60 from Cachabe. Provincia Esmeraldas, Ecuador. Eleittherodacrxhts longirostris — Dunn. 193 1 :41 1 . Diagnosis. — A member of the Eleuther- odactylus (Cr augastor) fitzingeri group having ( 1 ) skin on dorsum smooth or finely shagreen with low ridges outlining scapular markings, that on venter smooth; discoidal fold well anterior to groin; dor- solateral folds absent; (2) tympanic membrane and tympanic annulus prominent, round, its length V2- Va length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, as wide as lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males with vocal slits and nuptial pads; (7) first finger longer than second; discs on outer fingers broad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel tubercles absent; outer tarsal tubercles absent; fold on distal -A of inner edge of tarsus; (11) inner metatarsal tubercle elon- gate, 4-5x round outer metatarsal tubercle; super- numerary tubercles absent; (12) toes bearing lateral fringes; webbing encompassing all basal subartic- ular tubercles; fifth toe shorter than third; (13) dorsum reddish tan to dark brown, usually with darker brown hourglass-shaped mark; throat cream with dark spots/stripes extending onto chest; rest of venter cream or white; posterior surfaces of thighs uniformly brown; (14) SVL in males 28.8-34.4 mm, in females 43. 1-59.6 mm. Eleutherodactylus longirostris is the only mem- ber of the genus in the lowlands of western Ecuador with moderate webbing (enclosing all basal subarticular tubercles but not reaching the distal subarticular tubercles except on Toe V ). A different species in the cloud forests (E. loiistes) has nearly as much webbing but has a small tympanic annulus under the skin and a rounded snout in dorsal view. A smaller frog, E. achatiniis, superficially resembles E. longirostris but has a longer fifth toe ( longer than the third), lacks webbing between the toes, and usually has pale flecks on the posterior surfaces of the thighs. Description. — The species was redescribed by Lynch and Myers (1983). Coloration in life. — The dorsum is various shades of gray to brown with darker markings; the ventral surfaces of the hind limbs are pinkish-red to orange. Some ontogenetic change is evident in the following description. KU 165475-79 from Estacion Biologica Rio Polenque, Provincia Los Rios: Dorsum tan to brown with dark brown markings; upper lip creamy white, barred with brown in larger individuals; throat white with dark brown to black markings; belly white in juveniles, yellow in adults; groin and ventral surfaces of hind limbs pale rose; iris pale gold above and red below median, horizontal, brown streak (WED. 28 March 1975). Natural history. — In Colombia and northern Ecuador, this species occurs throughout the low- land rainforest, but in the southern part of its range in Ecuador, it usually is found near streams. At night, individuals sit on the ground or perch on low (<0.5 m) vegetation; by day. individuals take shel- ter under debris on the forest floor. Apparent recent hatchlings (SVL 8.5-8.8 mm) were found on 6 July 1976 at Rfo Palenque. Provincia Pichincha. Distribution. — Eleutherodactylus longirostris has an extensive distribution in the lowlands (< 1 200 m) from eastern Panama to southern Provincia Guayas, Ecuador; a disjunct population occurs in the middle Magdalena Valley in Colombia. Locali- ties where this species has been found in Ecuador are at elevations of 20-600 m. Most records for the species in Ecuador (20) are from the humid tropical regime, and 14 are from the humid subtropical regime. However, two are from the dry tropical regime, and one of these (Naranjito, Provincia Guayas) is about 150 km south of the next most southern locality (Fig. 38). 112 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 — 1 \ 81° 80 79^ 78° 0 20 40 60 80 100 ~ ' Kilometers /^ MM) 111 C ■-I1II1II ill V -^^^ ''■' ''' " |.. -0° y/ -(^Jfy jiw ){{ ^''-' . T\ \M L 3°- .^^~12S-^/ • £. longiwstris ■ £. loustes 81°/ } 80° A E. liiteolateralis 79° 78° 1 Fig. 38. Distribution of three species of Eleiitliero- dactylus in western Ecuador. Remarks. — Lynch and Myers ( 1 983 ) suggested that E. longiwstris is the sister species of E. talamancae. Among the many specimens reported by Lynch and Myers (1983), three (UIMNH 53393 and 53424 purportedly from the Rio Caoni, Provincia Esmeraldas, and UIMNH 55721 pur- portedly from San Javier, Provincia Esmeraldas) are misidentified; these specimens are E. malkini, a species known only from the upper Amazon Basin. The misidentification was noted by JDL upon reex- amining these specimens in 1 984. Eleutherodactylus malkini has appreciable toe webbing, but the pos- terior surfaces of the thighs are mottled cream and black. Eleutherodactylus loustes Lynch Plate 3 Eleutherodactylus loustes Lynch, iy79c:498. — Holo- type: KU 179234. adult female, from Maldonado, 1410 m, Provincia Carchi, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) loustes group having ( 1 ) skin on dorsum smooth, that on venter areolate; discoidal fold anterior to groin; dorsolateral folds absent; (2) tympanic inembrane absent; tympanic annulus sinall; (3) snout short, rounded in dorsal view, angularly rounded in profile; (4) upper eyelid bearing small tubercles, broader than lOD; low cranial crests present in adult females; (5) vomer- ine odontophores oval; (6) males with vocal slits; nuptial pads absent; (7) first finger slightly longer than second; discs broad; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing flaplike fold; (11) inner metatarsal tubercle oval, 8x conical outer metatarsal tubercle; supernumerary plantar tubercles indistinct; (12) toes bearing prominent flaplike lateral fringes, ba- sally webbed; webbing encoinpassing all basal subarticular tubercles; fifth toe longer than third; (13) dorsum brown with darker brown markings; throat gray; rest of venter cream with diffuse brown reticulations; posterior surfaces of thighs brown; (14) SVL in males 31.2-27.1 mm, in one female 46.7 mm. Eleutherodactylus loustes most closely re- seinbles E. jaimei from southwestern Colombia; the latter differs by having a visible tympanic membrane and less webbing between the toes. Description. — The original description by Lynch ( 1979c) is complete. Coloration in life. — Description of the types series ( KU 1 7923 1-49) from Maldonado. Provincia Carchi. indicated that in an adult female, the dor- sum was reddish brown with black spots; green wash on spaces between bars on limbs and on lower flank; concealed surfaces of thighs black with green- ish flecks; throat brown with cream flecks; rest of venter cream with brown spots; webbing pale yel- lowish-green; iris deep brown. In males, the dor- sum was greenish brown to green with gold or ELEUTHERODACTYLUS IN WESTERN ECUADOR 113 copper wash, especially anteriorly, and brown spots; venter dirty cream with brown flecks; tips of digits cream; iris dark brown with black flecks. In juve- niles, the heels and rump were reddish orange (JDL. 30-31 May, 1977). Natural history. — At Maldonado, Provincia Carchi, E. loustes was encountered only in the vicinity of small waterfalls. Adults and juveniles were clinging to wet rocks and perched on vegeta- tion in the spray zone of the falls. Likewise, at La Planada, Departamento Narifio, Colombia, frogs of this species were found on rocks in streams and on mud banks beneath waterfalls (Lynch and Burrowes, 1990). Distribution. — Eleuthewdactyhis loustes is known from two geographically close localities in cloud forest on the western slopes of the Cordillera Occidental — La Planada, 1200 m. Departamento Nariilo, Colombia, and Maldonado, 1410 m, Provincia Carchi, Ecuador; both localities are in the humid subtropical regime (Fig. 38). Because of its specialized habitats (spray zone of waterfalls), the species may enjoy a wider distribution that is not obvious from present collections. Remarks. — Lynch (1992a) placed this species and two Colombian species (E. hybotragus and E. juiinei) is the E. loustes species group. These three species share a synapomorphy of an abnormally large zygomatic ramus of the squamosal, which is evident externally as a swelling just in front of the tympanic region. Eleutherodactylus jaimei occurs at elevations comparable to those of E. loustes in Departamento Cauca, Colombia, but E. hybotragus inhabits the lowlands of Departamento Valle del Cauca. Colombia. Eleutherodactylus luteolateralis Lynch Plate 7 Eleutheroclactyliis luteolateralis Lynch, 1 976a: 1 3. — Ho- lotype: KU 131674, adult female, from Tandapi. 1460 m, Provincia Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum shagreen, that on venter areolate;discoidal fold prominent; dorsolateral folds absent; (2 ) tympanic membrane and tympanic annu- lus prominent, round, its length Va-A length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, slightly broader than lOD; cranial crests absent; (5) vomer- ine odontophores oblique; (6) males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs broad; (8) fingers bearing nar- row lateral fringes; (9) ulnar tubercles absent; ( 1 0) heel bearing one large, nonconical tubercle; tarsus lacking tubercles; (11) inner metatarsal tubercle oval, 2-3x oval outer metatarsal tubercle; supernu- merary tubercles numerous; (12) toes bearing lat- eral fringes; webbing absent; discs smaller than those on fingers; fifth toe much longer than third; (13) dorsum gray with pale and dark brown stripes; pale dorsolateral bands from eye to groin; venter cream with brown flecks; posterior surfaces of thighs brown; lower flanks, groin, and anterior surfaces of thighs with cream spots (lemon-yellow to orange-yellow in life) edged in black; ( 14) SVL in males 16.6-23.6 mm, in females 25.6-29.5 mm. Eleutherodactylus luteolateralis closely re- sembles the smaller E. walked (males 15-18 mm, females 2 1-25 mm SVL) occuiring at lower eleva- tions. Structurally, the two species seem to be identical. However, dorsal stripes are present in all E. luteolateralis, whereas this pattern occurs in few E. walked. All individuals of £. luteolateralishave. a pale band extending from the eye to the groin; no such band occurs in E. walked, even those indi- viduals having a striped pattern. The smaller f.parvillus also has yellow spots on the flanks, groin, and anterior surfaces of the thighs, but these spots are not outlined in black. Description. — The original description by Lynch (1976a) is complete. Coloration in life. — The variation in coloration is documented as follows. KU 111380-84 from Tandapi, Provincia Pichincha: Dorsum pale and dark brown; flanks and anterior and posterior surfaces of thighs with bright yellow spots; side of head black; venter cream and brown; iris pale bronze-brown with brown, median, horizontal streak (JDL, 27 July 1967). KU 120254 from Tandapi, Provincia, Pichincha: Dorsum striped with medium brown and pale brown or reddish brown; pale creamy-brown stripes above 114 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 flanks; limb.s reddish brown with dark brown bars; groin, anterior surfaces of thighs, and concealed surfaces of shanks with lemon-yellow spots; poste- rior surfaces of thighs dark gray; some narrow white bars on lips; iris gold above and gray below reddish-brown median, horizontal streak, hnely stippled with black (JDL. 29 July 1968). KU 165505-14 from 4 km NE Dos Rios, Provincia Pichincha: Dorsum reddish tan to yel- lowish tan; groin and anterior and posterior sur- faces of thighs with orange-yellow spots; iris dull bronze (WED. 1 April \915). Natural history. — All individuals have been found on low vegetation (<2 m) at night in cloud forest; many individuals were on herbs and fems near streams. This species is collected infrequently. The observation that the species became more abundant as trails became more overgrown at Tandapi, Provincia Pichincha. between 1967 and 1970 (Lynch. 1976a) hints that E. luteolateralis inhabits deep forest rather than forest edge. Distribution. — This species is restricted to cloud forests at elevations of 1140-1960 m in the Rio Blanco drainage in Provincia Pichincha, Ecuador (Fig. 38). One locality is in the humid tropical regime, and 10 are in the humid temperate regime. Remarks. — This species has worried JDL for nearly 20 years because of the absence of structural differences from E. walkeri. The larger size of E. luteolateralis may reflect only the higher eleva- tions of collections, and the monomorphy (striped pattern only) could be a consequence of fixation by drift; E. walkeri has a low frequency of striped morphs (Lynch, 1 974). The pale dorsolateral bands of E. luteolateralis are not associated with any pattern polymorphs and represent the only un- equivocal (albeit weak) evidence that E. luteolateralis is a species apart from the lowland E. walkeri. Eleutherodactylus lymani Barbour and Noble Plate 2 Eleutherodactylus lymani Barbour and Noble, 1920:403.— Holotype: MCZ 5422. young female, from Perico. Depailamento Cajamarca. Peru. Eleutherodactylus carrioni Parker. 1 932:23. — Holotype: BM 1947.2. 15.99, adult female, from Loja, Provincia Loja, Ecuador. Synonymy fide Lynch, 1969:263. Eleutherodactylus lymani — Lynch, 1969:236. Diagnosis. — A member of the Eleuthero- dactylus (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, its length V2--A length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, usually narrower than lOD; cra- nial crests absent; (5) vomerine odontophores subtriangular in outline; (6) males lacking vocal slits; nuptial pads present; (7) first finger longer than second; discs small; (8) fingers bearing lateral keels; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus lacking tubercles; inner sur- faces of tarsal bearing fold; (11) inner metatarsal tubercle elongate, 4x round outer metatarsal tu- bercle; supernumerary plantar tubercles absent; (12) toes bearing lateral fringes; webbing absent; fifth toe slightly longer than third; (13) dorsum brown with dark brown or black markings; venter cream to white; groin and posterior surfaces of thighs black with white spots or reticulations; (14) SVL in males 25.7-43.6 mm. in females 52.9-69.3 mm. By having the first finger shorter than the second and the fifth toe only slightly longer than the third, E. lymani in western Ecuador requires comparison only with E. achatinus, actites, and w-nigrum. From all of these, E. lymani difl^rs by having an inner tarsal fold. Inf. lymani, the posterior surfaces of the thighs are black with white spots or reticula- tions; this pattern enables quick distinction from E. achatinus (brown with small cream flecks), E. actites (black flecks), and E. w-nigrum (yellow with black spots or reticulation). Eleutherodactylus lymani resembles the smaller E. achatinus in hav- ing dorsolateral folds; these are absent in E. actites and E. w-nigrum, both of which are comparable in size to E. lymani. Eleutherodactylus lymani closely resembles two other species in the E. conspicillatus group — E. citriogaster and E. condor 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. ELEUTHERODACTYLUS IN WESTERN ECUADOR 115 Description. — The redesciiption of the species by Lynch ( 1969) is adequate. Coloration in life. — The doisuni is oHve-green to reddish brown. Descriptions otTiving specimens from western Ecuador are not available; the fol- lowing description is based on specimens from southern Ecuador. KU 165539^0 from 9 km S Loja, Provincia Loja: Dorsum reddish brown with darker reddish- brown chevrons on back and dark gray bars on limbs narrowly outline with creamy gray; dorsolat- eral fold dull red; flanks and posterior surfaces of thighs mottled cream and black; palmar and plantar surfaces black; other ventral surfaces creamy white; iris bronze with black flecks ( WED, 9 March 1 975 ). Natural history. — Most individuals have been found by day beneath rocks and logs in pastures. Most records are from disturbed areas in relatively dry forest. A female (KU 165540) was found with what is presumed to be her egg mass ( KU 1 6623 1 ) beneath a rock on 9 March 1975. Distribution. — This species is known from el- evations of 610-3000 m on the western slopes of the Andes in southern Ecuador and up into subparamo in Provincia Loja, Ecuador, and north- ern Peru (mapped by Duellman, 1992). In western Ecuador, the species has been taken at elevations of 610-1600 m in the dry subtropical regime (Fig. 39). The specimen reported here from Cataviiia, Provincia Azuay (3° 15' S Lat.) is the northernmost record for the species. Remarks. — No cladistic analysis has been per- formed of the species in the mostly lowland Eleiithewdactylus conspicillatus group, but we follow Duellman ( 1 992) in suggesting that E. lymani is most closely related to E. citriogaster and E. condor, species found in the Amazonian, rather than the Pacific, drainage. Eleutherodactylus muricatus Lynch and Miyata Plate 6 Eleutherodactylus muricatus Lynch and Miyata, 1980:2.— Holotype: MCZ 94469. adult male, from theRfoFaisanes, 14.4km(byroad)ELaPalma, 1380 m, Provincia, Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) unistrigatus group — 1 r 81° 80' 79^.^ 78° 0 20 40 60 80 100 y^ ^ Kilometers /^ T '(Id 111 ( 2(10(1 111 N \ _)v-i A>^^ "^ .40(1(1111 1 I 1 IVriiUiiicnl I V, ) '■•""■'"' '' ' * ''r^ """" 1^ :/ x~^ ~'''' t -0° y 4 ■■■ J • \ / 1 '^ ': - 1/^ / / iff / ; - i°- -2° ] \ f >4 - - .'i-f^ •4» • E. lymani 3°- --•- ■ E. nniricatus 81°/ } 80° •V A E. ocellutus 79° 78° 1 / ■■/-TTr> 1 . ' ^: 1 1 1 Fig. 39. Distribution of three species of Eleuthero- dactylus in western Ecuador. having ( 1 ) skin on dorsum smooth with many conical tubercles, that on venter feebly areolate; discoidal fold well anteriad to groin; dorsolateral folds absent; (2) tympanic membrane and tym- panic annulus moderately distinct, its length '/4 length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid bearing many conical tubercles, much broader than lOD; cranial crests absent; (5) vomerine odontophores triangular in outline; (6) males usually having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs broad; (8) fingers bear- ing lateral keels; (9) one or two small ulnar tu- bercles present; ( 10) heel bearing one conical tu- bercle; outer edge of tarsus bearing low tubercles; inner tarsal fold present, short; (11) inner metatar- 116 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 sal tubercle elongate. lOx subconical ouler meta- tarsal tubercle; supernumerary tubercles at bases of toes; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; ( 13) dor- sum brown with black spots; venter brown with cream flecks on belly and darker brown chevrons on throat; posterior surfaces of thighs brown; lower flanks, groin, and anterior surfaces of thighs with creain spots (lemon-yellow to orange-yellow in life) edged in black; ( 14) SVL in males 3 1 .8^0.7 mm, in females 36.8^6.3 mm. Eleiitherodactylus muricatus perhaps is most closely related to E. laticUiviiis and E. latidiscus, but differs from both by having vocal slits. Lynch and Miyata (1980) compared E. muricatus with frogs of the E. cerasinus group (especially E. crenungais and frogs now known as E. lahiosus), but the similarities among these three species are superficial. Skin texture, tubercle development, andcolor patterns of £. laticlavius, latidiscus, and muricatus are alike in most cases, so much so as to cause skepticism as to how many species are in- volved. This problem is exacerbated by size differ- ences between lowland and cloud forest samples and the apparent rarity of E. muricatus. Description. — The original description by Lynch and Miyata (1980) is adequate. Coloration in life. — According to Lynch and Miyata (1980), the dorsum in topotypes is pale to dark brown with black rings around large tubercles, and the flanks are paler brown with a yellowish wash; the venter is dark puiplish brown with yel- lowish-brown mottling, and the iris is gold with black flecks. Specimens from the lowlands lack the black rings around the tubercles on the dorsum, and the venter is muddy yellow with dark brown mot- tling; the groin and ventral surfaces of the hind limbs are purplish brown, and the iris is copper. Natural history. — One individual was about 60 cm above the ground at the edge of a banana grove at night, and some were on vegetation in primary lowland rainforest at night; others were on streamside vegetation within 1 .5 m of the surface of the water at night. Distribution. — This species is known from only three localities at elevations of 200-1380 m in Provincia Pichincha. Ecuador (Fig. 39). One local- ity is in the humid tropical regime and two are in the humid subtropical regime. Remarks. — An obvious difference in size ex- ists between specimens from the lowlands (2 adult females 33.8-36.0 mm SVL) and the type locality at 1380 m (I female 46.3 mm SVL). However, structurally these individuals are alike. One adult male (MCZ 94710) lacks vocal slits. Eleiitherodactylus necerus Lynch Plate 1 EleutlierodactylusnecerusLynch, 1975a:32. — Holotype: USNM 1 95798, adult female, from Mindo. Provincia Pichincha. Ecuador. Diagnosis. — A member of the subgenus Eleiitherodactylus (Craugastor) hufoniformis group having (1) skin on dorsum tuberculate, bearing many short ridges, that on venter smooth; discoidal fold prominent; dorsolateral folds absent; (2) tym- panic membrane and tympanic annulus prominent, higher than long, its length /6--/s length of eye; (3) snout short, rounded in dorsal view, truncate in profile; (4) upper eyelid tuberculate (none elon- gate), much broader than lOD; low parasagittal crests on frontoparietals; ( 5 ) vomerine odontophores arched; (6) males having vocal slits and white nuptial pads; (7) first finger longer than second; discs naiTow; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles and folds; (11) inner metatarsal tubercle compressed. 6-7x obscure outer metatarsal tu- bercle; supernumerary tubercles absent; (12) toes bearing thin lateral keels; webbing absent; discs small, but larger than those on fingers; fifth toe shorter than third; (13) dorsum brown with indefi- nite darker brown mottling; venter cream or white with brown spots along margin of jaw and bases of arms; posterior surfaces of thighs black with large cream spots; ( 14) SVL in males 44.9-68.4 mm. in females 82.2-93.3 mm. Eleiitherodactylus necerus is difficult to con- fuse with other species, especially if an adult fe- male is at hand, because of the large size. It is a species with a broad head and thus likely to be confounded only with other broad-headed spe- cies— E. cerastes and E. helonotus. Eleiithero- dactylus necerus lacks an elongate tubercle on the upper eyelid (present in E. cerastes) and has smooth ELEUTHERODACTYLUS IN WESTERN ECUADOR 117 skin on the venter (areolate in E. hehmotus). In E. uecenis. the inner metatarsal tubercle is compressed, whereas in E. cerastes and E. hehmotus (and in most South American Eleiitheroclactyliis) the inner metatarsal tubercle is not compressed. Description. — The original description (Lynch, 1975a) is largely complete, even though it was based on a single adult female and several juve- niles. Proportions for four males and eight females are: tibia length/SVL 58.5-63.2 {x = 61.0), 55.3- 65.8 (.V = 59.6 ± 1.2); HW/SVL 45.9-50.8 (r = 47.3), 43.0-46.9 (.f = 45.0±0.4); upper eyelid/IOD 137.1-216.2 (.v = 168.6), 141.7-197.5 (v = 162.0 ± 7.7); tympanic annulus/eye 35.4-50.0 ( J = 44.7). 30.8-43.4 (.V = 37.2 ± 1.5); E-N/eye 71.0-85.5 (.? = 77.7), 74.7-92.7 ( x = 83.7 ± 1 .9). Aside from size and nuptial pads in males, there is little external sexual dimorphism. Males have vocal slits posterolaterally in the floor of the mouth. The tympanic annulus is slightly larger in males than in females, but E. uecerus is like E. aiuitipes, anomalus, cheiropletluis. and zygodactylus in not exhibiting pronounced sexual dimorphism in tympanic annu- lus size. Coloration in life. — The variation is noted in the following descriptions. KU 166067 from 4 km E Dos Rios, Provincia, Pichincha: Dorsum dull brown; throat cream with brown reticulations; belly creamy yellow; anterior surfaces of thighs and ventral surfaces of hind limbs pale orange with dark brown flecks; posterior surfaces of thighs orange with dark brown reticula- tions; iris dark brown with radiating cream bars (WED, 2 April 1975). KU 179076-77 from Maldonado, Provincia Carchi.: Dorsum brown with some black markings in one individual and faint green wash on flanks in another; throat brown with white flecks or spots; belly dull yellow or pale yellow-orange with cream spots; ventral surfaces of hind limbs orange; poste- rior surfaces of thighs orange with brown reticula- tions; iris bronze with black flecks and anterior and posterior brown triangles (JDL, 27, 29 May 1 977). KU 1 79080 from Tamlapi, Provincia Pichincha: Dorsum greenish and brown; belly off-white; throat brownish with cream flecks; concealed surfaces of limbs pale orange; upper edge of iris copper with brown reticulation, gray below with brown tri- angles anterior and posterior to pupil (JDL. 21 June 1977). Natural history. — Most individuals have been observed in the immediate vicinity of streams, where they sit on the bank or on rocks in the stream; they have been observed to leap into the water to evade capture. One was under a log at the base of a waterfall by day. Distribution. — Eleutherodactylus necerus in- habits humid lower montane forest at elevations of 600-1540 m on the lower Pacific slopes of the Andes in Ecuador from Provincia Cotopaxi north- ward to Provincia Carchi (Fig. 40); all localities are in the humid subtropical zone. We anticipate that it will be found in southern Colombia, because it has been found at Maldonado, 1 km from the Colom- bian border. Remarks. — Eleutherodactyhis necerus is the nearest relative of £. bufoniformis, an inhabitant of lowlands in Colombia and Panama. Its apparent rarity probably is a consequence of being over- looked by collectors. It was not found at Tandapi in 1967-1970 when we thought the locality had been sampled thoroughly, but it was found there in 1 977. Lynch is guilty of spending too much time gazing at vegetation in quest of dainty centrolenids and 81 80 -1 0 20 40 60 80 100 Kilometers y .<()l)m ( 2000 111 \ _ 1 >4000 111 Permanenl \ • E. iu'ccrus\ 9' 78' Fig. 40. Distribution of Eleutherodactyhis necerus in Ecuador. 118 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 small Ek'utlwrodiutylus and will always be thank- ful to Thomas J. Berger for showing him large toadlike frogs on the ground along the stream where he had Just passed; .search images sometimes are so powerful that a frog weighing a half kilogram somehow is not seen. Eleuthewdactyhis nyctophyUix Lynch Plate 6 Eleutherodacryhts palmcri — Lynch 1971b: 146^7. fig. 29. 226. Eleuthewdactyhis nyctophylax Lynch. 1 976a: 1 6. — Ho- lotype: KU 1 10909. adult male, from Tandapi, 1460 m, Provincia Pichincha. Ecuador. Diagnosis. — A member of the Eleuthero- dactylus {Eleuthewdactylus) imisthgatus group having ( 1 ) skin on dorsum finely shagreen, that on venter areolate; discoidal fold prominent; dorsolat- eral folds absent; (2) tympanic membrane and tympanic annulus prominent, higher than long, its length, '/s-'/2 length of eye; (3) snout long, subacuminate in dorsal view, pointed in profile; (4) upper eyelid bearing one small tubercle, slightly narrower than lOD; cranial crests absent; (5) vomer- ine odontophores triangular in outline; (6) males having vocal slits, subgular vocal sac, and nuptial pads; (7) first finger shorter than second; discs broad; (8) fingers bearing lateral fringes; (9) ulnar tubercles small; (10) heel bearing small conical tubercle; tarsus lacking tubercles and folds; (11) inner metatarsal tubercle elongate, 5-6x round outer metatarsal tubercle; plantar surface areolate; (12) toes bearing prominent lateral fringes; web- bing basal (not encompassing basal subarticular tubercles); discs large; fifth toe much longer than third; (13) dorsum brown with darker brown mot- tling; venter dirty cream with brown flecks; poste- rior surfaces of thighs dark brown with cream flecks; (14) SVL in males 21.9-31.4 mm, in fe- males 32.1-37.8 mm. Eleutherodactylus nyctophylax is most similar to E. eugeniae and E. subsigillatus. It differs from E. eugeniae by having a small conical tubercle on the heel and a small tubercle on the upper eyelid; moreover, in E. eugeniae, the posterior surfaces of the thighs are cream with brown reticulations. Structurally, the smaller E. subsigillatus (males 1 9.3-28.5 mm, females 30.0-33.4 mm SVL) is like E. nyctophylax, but in many specimens the dorsum has more small tubercles; furthermore, E. subsigillatus has white spots on the flanks and concealed surfaces of the limbs and may have black marbling associated with those spots (more com- mon in females than in males). Description. — The original description by Lynch ( 1976a) is complete. Coloration in life. — The dorsum is greenish- tan to brown, with or without a middorsal cream stripe; the venter is white to yellow. The eye is distinctive in having orange or red sclera. The variation is evident from the following descrip- tions. KV 110899-905 from Tandapi, Provincia Pichincha: Dorsum creamy tan. brown, or pale greenish-tan with a diffuse pattern of brown and black; some individuals with yellow spots on dor- sum; venter creamy-white to gray; lower flanks, groin, and anterior surfaces of thighs white or yellow with gray reticulations; iris greenish gray, with fine black reticulations; concealed surface of eye orange (JDL, 17 July 1967). KU 110942-53 from Tandapi, Provincia Pichincha: Dorsum creamy tan, brown, greenish brown, or reddish brown with brown or black markings; vertebral stripe cream; venter cream with pink or orange cast, with or without brown flecks; groin cream with brown reticulations; poste- rior surfaces of thighs dark with cream flecks; iris creamy gray with pale green cast and brown reticu- lation (JDL, 23 July 1967). KU 165548-89 from 3.5kmNEMindo, Provincia Pichincha: At night, dorsum dull tan; by day, dull brown with darker brown bars on limbs; venter dull creamy-tan with grayish-brown suffusion; groin mottled black and creamy white; iris pale bronze with fine black reticulation; sclera of eye red (WED, 7 April 1975). Natural history. — At night, this inhabitant of cloud forests perches on vegetation up to 3 m above the ground; individuals were found in deep forest, along forest edge, and near streams. By day, frogs were found in leaf litter, in bromeliads, and in the axils of leaves of elephant ear plants. Although hundreds of individuals were observed at Tandapi in 1967-1978, no vocalizations were traced to this ELEUTHERODACTYLUS IN WESTERN ECUADOR 119 species, and no egg masses were found. A pair in amplexLis was found on 3 July 1968. and small juveniles were abundant in June-August. Distribution. — Eleuthewdactylusnyctophylax occurs at elevations of 1 1 40-2 1 00 m on the western flank of the Andes in provincias Cotopaxi and Pichincha, Ecuador (Fig. 41 ). Nine records of oc- currence are in the humid subtropical regime and two are in the humid temperate regime. Remarks. — To date, Eleuthewdactylus nycto- phyla.x and E. sithsigillatus have allopatric distri- butions, whereas E. nyctophylax and E. eugeniae are sympatric at 6.3 km E of Tandapi, Provincia Pichincha. and their distributions overlap in the Mindo-Tandapi area. Eleuthewdactylus ocellatus Lynch and Burrowes Plate 3 Eleuthewdactylus ocellatus Lynch and Burrowes, 1990:18.— Holotype: IND-AN 1441, adult female, from Reserva Natural La Planada, 1780 m, Municipio Ricaurte. Departamento Narifio. Colombia. Diagnosis. — A member of the Eleuthero- dactyliis {Eleuthewdactylus) cerasinus group hav- ing (1) skin on dorsum shagreen, that on venter smooth; discoidal fold well anteriad of groin; dor- solateral folds absent; (2) tympanic membrane and tympanic annulus prominent, small, its length V^ length of eye; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid lacking prominent tubercles, about as wide as lOD; cranial crests absent; (5) vomerine odontophores triangu- lar in outline; (6) adult males unknown; (7) first finger slightly shorter than second; discs large, emarginate or truncate; (8) fingers bearing lateral fringes; (9) ulnar tubercles small; (10) heel and tarsus bearing small tubercles; inner tarsal fold short; (11) inner metatarsal tubercle elongate, 4x round outer metatarsal tubercle; supernumerary tubercles at bases of Toes II- V; (12) toes lacking lateral fringes; webbing absent; fifth toe longer than third; (13) dorsum brown with black spots having pale centers; venter cream with brown mot- tling; posterior surfaces of thighs dark brown; ( 14) SVL in one female 45.7 mm. Fig. 41. Distribution of Eleutherodactylus nyctophylax. Eleutherodactylus ocellatus is most similar to E. crenunguis but differs from that species in lack- ing a tubercle on the heel, having the first finger shorter than the second (first longer than second in E. crenunguis), and in color pattern. Species in the E. cerasinus group differ in the combination of male secondary sex characters (Lynch et al., 1 994), but adult males of E. ocellatus are not known. Description. — The original description by Lynch and Burrowes ( 1 990) was based on only two specimens. Coloration in life. — Lynch and Burrowes (1990:19) described the coloration of specimens from the type locality as: "In life, E. ocellatus was grayish-tan with brown ocelli surrounding tan tu- bercles; limbs grayish-tan with brown transverse bars; venter creamy tan with brown marbling; brown chevrons were visible on the throats of larger individuals; iris brown with gray blotches." Natural history. — According to Lynch and Burrowes (1990), at La Planada, Departamento Nariiio, Colombia, individuals were on low vegeta- tion (0.5-2.0 m) above stream banks or on branches high in the canopy at night. One Ecuadorian speci- men (USNM 286293) was obtained from the canopy of a felled tree by day. 120 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Distribution. — Limited distributit)nal data in- dicate that EleHthewdactyliis occllatus inhabits cloud forests at elevations of 1255-1780 m on the Pacific versant of the Andes in the departmentos Cauca and Nariiio, Colombia, and extreme north- ern Provincia Carchi, Ecuador. Three of the Ecua- dorian localities are at elevations of 1200-1500 in the humid subtropical regime; an additional local- ity is at 2560 m in the humid temperate regime in Provincia Imbabura (Fig. 39). Remarks. — The scarcity of specimens and ob- servations of this species in the canopy suggest that this species inhabits a stratum of forest not ad- equately sampled by collectors. A juvenile female (USNM 286293) with a SVL of 21.7 mm from Provincia Carchi, Ecuador, agrees in all details with the type specimen. Two specimens (ECO 168-69) from Cuellaje, Provincia Imbabura. are juvenile females (SVL 26.6 and 22. 1 mm. respec- tively) but appear to have tubercles on the upper eyelids and have relatively large tubercles on the heels. Furthermore, these two frogs have bold black-and-white bars on the undersides of the shanks, anterior surfaces of the thighs, posterior surfaces of the flanks, and in the groin. As in typical E. oceJkitus (and those from Departamento Cauca, Colombia), the throat has a pattern of dark, reverse chevrons. The differences in tuberculation might be a reflection of immaturity, but the color pattern on the throat suggests that these specimens are E. ocellatus. Until the species is collected in larger numbers (including adult males), there will remain some doubt as to its characters with respect to E. crenimguis. Eleutherodactylus ornatissimus (Despax) Plate 1 Hylodes ornatissimus Despax, 1911:91. — Holotype: MNHN 6264. from "Ecuador." Eleiitherodcunius oruatissiimis — Peters. 1 955:350. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) uuistrigatus group having ( 1 ) skin on dorsum shagreen, that on venter smooth; discoidal fold poorly defined posteriorly; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus evident, round, its length '/i length of eye; (3) snout long, acuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, slightly narrower than lOD; cranial crests absent; (5) vomerine odontophores low; (6) males having vocal slits and nuptial pads; (7) first finger shorter than second; discs large, round; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles and folds; (11) inner metatarsal tubercle oval, 4-5x round outer metatarsal tubercle; supernumerary plantar tubercles numerous, indistinct; (12) toes bearing lateral fringes; webbing absent; discs large; fifth toe much longer than third; (13) dorsum and venter pale cream (green in life) with black markings dorsally tending to form lines on body and limbs; posterior surfaces of thighs with a few pale spots; (14) SVL in males 21 .6-27.0 mm, in two females 37.8 and 38.1 mm. In life, this pale green frog with black lines and spots on the dorsum is so distinctive as to be impossible to confuse it with any other species of neotropical frog. It is sufficiently "pretty" that some persons have doubted that it is an Eleutherodactylus. Among frogs of the subgenus Eleutherodactylus, it is relatively distinctive in having a long snout and in lacking tubercles on the dorsum. Description. — Lynch's (1971a) description is adequate, but he did not distinguish between adult and juvenile females and failed to notice the nuptial pads of adult males. Coloration in life. — The ornate coloration of this frog consists of a green dorsum and paler green venter and posterior surfaces of the thighs; the stripes and spots on the dorsum are black. The vocal sac is yellow, and the hands and feet are tinted with orange. The iris is green with a bronze tint and fine black reticulations. However, one individual was described as having a yellow venter, greenish- yellow limbs, orange toes, and greenish-gold iris (RWM, 7 January 1979). Natural history. — In 1968, JDLcollected many juveniles in a small woods (probably mostly sec- ondary forest) on the outskirts of Santo Domingo de los Colorados, Provincia Pichincha, but other- wise usually only one to three individuals have been collected at a given locality. However, Luis A. Coloma (pers. comm.) noted the existence of many ELEUTHERODACTYLUS IN WESTERN ECUADOR 121 specimens in the QCAZ collection from San Fran- cisco de Las Pampas, Provincia Cotopaxi. Those collected in 1968 were found mostly on the nearly vertical leaves of large elephant ear plants at night. During a longer stay at Santo Domingo de los Colorados in 1977. three biologists failed to find a single specimen off. ornatissimus, although they sampled a variety of forested (primary and second- ary) habitats. At other sites, individuals have been found in the axils of leaves of large elephant ear plants by day. Thus, it seems that there is a close association between this distinctive frog and these large plants. Distribution. — Eleiitherodactyhis ornatissimus has been collected at 15 localities at elevations of 400-1800 m on the western flank of the Andes in Ecuador (Fig. 42); only four of the localities are above 1000 m. Two localities are in the lower part of the humid temperate regime, whereas 1 3 are in the humid subtropical regime. Remarks. — Lynch ( 1 97 1 a) reported that males are "19.5 to 25.0 mm SVL" and did not mention females. The smallest adult male available (KU 141968) has a SVL of 2 1 .6 mm and has vocal slits and nuptial pads, but a male (KU 1 19748) with a SVL of 19.5 mm lacks both. Only two adult fe- males are known; a juvenile female (KU 141969) has a SVL of 34.0 mm. Eleutherodactyliis ornatissimus does not seem to be closely related to any other species known to us. Its peculiarity stems mostly from its being structurally undistinguished and having a stunning coloration. At present, it can be assigned to the E. unistrigatus group. Eleutherodactyius parvillus Lynch Plate 7 Eleutherodactylus parvilhis Lynch, 1976a: 19. — Holo- type: KU 1 II 345. adult female, from Tandapi, 1460 m, Provincia Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum finely shagreen, that on venter areolate; discoidal fold prominent; dorsolat- eral folds absent; (2) tympanic membrane and tympanic annulus evident, round, its length V4--/5 n 80 0 20 40 60 80 100 Kilometers • E. ornatissimus] Fig. 42. Distribution of Eleutherodactylus ornatissimus. length of eye; (3) snout rounded to subacuminate in dorsal view, rounded in profile; (4) upper eyelid lacking tubercles, usually narrower than lOD; cra- nial crests absent; (5 ) vomerine odontophores oval, not prominent; (6) males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs broad; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) heel bearing small tubercles; outer edge of tarsus bearing minute tu- bercles; inner edge of tarsus lacking tubercles and folds; (11) inner metatarsal tubercle oval, 4x round outer metatarsal tubercle; supernumerary tubercles at bases of Toes II-IV; (12) toes bearing narrow lateral fringes; webbing absent; fifth toe much longer than third; (13) dorsum cream to gray with brown markings; venter cream finely stippled with brown; lower flank, anterior and posterior surfaces of thighs, and concealed surfaces of shanks unpigmented (lemon yellow in life); ( 14) SVL in males 15.5-19.4 mm, in females 18.4-25.9 mm. Eleutherodactylus parvillus is most similar to E. palmeri (Boulenger) and E. ridens (Cope) among named species. The concealed surfaces of the flanks and hind limbs are red in E. ridens and yellow in the other two species. Eleutherodactylus palmeri has nuptial pads in males (absent in the other two species) and small tubercles on the upper eyelid 122 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 and heel, whereas E. ridciis has a tubercle on the heel but none on the eyelid: minute tubercles are present on the eyelids and heels o^ E. parvilln.s. but these are not always evident in preserved speci- mens. Description. — The original description by Lynch ( 1976a) is complete. Coloration in life. — At least in the vicinity of Santo Domingo de los Colorados, Provincia Pichincha. young individuals have orange flash colors, whereas these are yellow in adults. The variation is documented in the following descrip- tion. KU 111345-51 from Tondapi, Provincia Pichincha: Dorsum brown, pale orange-tan, or greenish brown becoming rusty laterally, or in female, grayish tan. with brown spots, broad orangish-brown dorsolateral stripe, or broad brown vertebral mark and black interorbital bar; limbs orange-tan to rusty-brown with dark brown bars (brown with black bars in female); venter gray to nearly black; lower flank, groin, anterior and pos- terior surfaces of thighs, and concealed surfaces of shanks orange-yellow to lemon-yellow; cream spots below eyes in some individuals; iris grayish brown to reddish brown (JDL, 17-23 July 1967). Natural history. — This small species seems to be about equally abundant in cloud forest and lowland tropical rainforest. By day, individuals have been found beneath leaf litter and in the axils of elephant ear plants, whereas at night the frog is active on low (<3m) vegetation at the edge of, and within, forests. However, the observation by R. W. McDiarmid of an individual (USNM 285795) in a small bromeliad about 30 m up in a tree that was felled indicates that this species also invades the canopy of the rainforest. Distribution. — Eleutherodactylus parvillus occurs at elevations of 220-2000 m in the lowlands and on the western slopes of the Andes in Ecuador (Fig. 43). The only localities for the species at elevations above 1000 m are in the drainage of the Rio Blanco in Provincia Pichincha. Four sites are in the humid tropical regime, seven in the humid subtropical, and three in the humid temperate. Remarks. — The report of this species from Colombia by Lynch and Ruiz (1983) applies to individuals now identified as Eleutherodactylus Fig. 43. Distribution of four species of Eleuthero- dactylus in western Ecuador. palnieri ( Boulenger). That Colombian species also has yellow flash markings in the groin and on the concealed surfaces of the hind limbs. It is not known to which species Savage (1981) referred when he reported that E. ridens ranged as far south as Ecuador; he may have meant E. panillus. Eleutherodactylus phoxocephalus Lynch Plate 5 Eleutherodactylus phoxocephalus Lynch. I979a:29. — Holotype: adult male, from Pilalo, 2340 m, Provincia Cotopaxi, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) unistrigatus group ELEUTHERODACTYLUS IN WESTERN ECUADOR 123 having ( 1 ) skin on dorsum shagreen, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus evident, round, its length '/<-'/: length of eye; (3) snout rounded in dorsal view, pointed in profile, bearing terminal keel; (4) upper eyelid lacking tubercles, narrower than lOD; cranial crests ab- sent; (5 ) vomerine odontophores oblique; (6) males having vocal slits and nuptial pads; (7) first finger shorter than second; discs broad; (8) fingers bear- ing lateral keels; (9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles and folds; (11) inner metatarsal tubercle oval, 4-6x round outer metatarsal tubercle; supernumerary tubercles at bases of Toes II-IV; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; (13) dorsum gray to brown with few or no darker markings; venter cream; lower flank, groin, and concealed surfaces of limbs white with brown or black reticulations; (14) SVL in males 22.3-29.9 mm, in females 29.6-38.4 mm. Eleutherodactylus phoxocephalus superficially resembles E. subsigillatus in having the flanks and concealed surfaces of the hind limbs reticulated with brown or black, but it is distinguished easily by the presence of a papilla or vertical keel on the snout; also, E. phoxocephalus lacks tubercles on the upper eyelid, heel, and tarsus. Description. — The original description by Lynch ( 1979a) is complete. Coloration in life. — The dorsal coloration is highly variable, as is evident from the following descriptions. KU 13 1404-79 from Pilalo, Pwvincia Cotopaxi: Dorsum pale greenish brown, pale yellow, rusty- brown, pale brown, or dark brown with brown (black middorsally) mottling or fine reticulations; axilla, groin, posterior surfaces of thigh, and con- cealed surface of shank yellow with brown reticu- lations (faint in juveniles, dark in adults); venter cream with green tint and fine brown reticulations in adults; throat dusky in females, pale yellow in males; iris pale bronze with fine black reticulation and median, horizontal, brown streak; some indi- viduals with cream spots (with or without black borders) on dorsum (JDL, 3 July 1970). KV 142075-101 from Pilalo, Provincia Cotopaxi: Dorsum brown to reddish brown; some individuals with olive-green longitudinal streaks, others with yellow spots outlined with dark brown; groin and posterior surfaces of thighs yellow with black mottling; belly white; throat yellow; iris copper (WED, 7 July 1971). KU 165556-58 from 5 km E of Chirihoga, Provincia Pichincha: Dorsum pale green with dark or reddish-brown markings, or yellowish tan with green middorsal stripe bordered by black; limbs orange-tan with pale green cross bars; groin and posterior surfaces of thighs yellow and dull brown; belly white to creamy yellow; other ventral sur- faces pale pinkish gray to dull yellowish-tan; iris bronze to coppery brown with fine black reticula- tions (WED, 4 April, 1 1 May 1975). Natural history. — Although E. phoxocephalus is widely distributed, most observations on the natural history of the species were made in the vicinity of the type locality, where the frogs are abundant in forest-edge situations, but scarce in the interior of the forest. By day, individuals were in the axils of elephant ear plants and in terrestrial and arboreal bromeliads up to 13 m above the ground. At night, individuals were perched on vegetation. Distribution. — In western Ecuador (Provincia Pichincha southward). E. phoxocephalus occurs in upper humid montane forest at subparamo at eleva- tions of 1800-3100 m (Fig. 44); two localities are in the humid subtropical regime, 12 in the humid temperate, and one in the humid subtemperate. In southern Ecuador, it occurs in subparamo at eleva- tions of 2010-2960 m (Lynch, 1979a). The species also is known from humid montane forests at elevations of 1850-2770 m in the Cordillera de Huancabamba in northern Peru (Duellman and Wild. 1993). Remarks. — In spite of lacking tubercles on the eyelid, heel, and tarsus, we think that E. phoxocephalus is most closely related to E. eugeniae, nyctophylax, and subsigillatus. The grouping of these four allopatric species admit- tedly is phenetic. Additionally, there is an assort- ment of phenetically similar species in Colombia, among which are E. boulengeri Lynch, a species widespread in subparamo habitats in the Cordillera Central, and E. tayromi Lynch and Ruiz, a species inhabiting cloud forests in the Sierra Nevada de Santa Marta. Until these species have been studied 124 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 • E. phoxocephaliis ▲ E. scolodiscus ■ E. simonbolivarl Fig. 44. Distribution of three species of Elenthero- dactylus in western Ecuador. more closely, most of the species loosely referred to as forming the "E. lacrimosus assembly" (Lynch, 1980(1) should be considered as potentially closely allied to E. eugeniae, nyctophylax, pho.xocephalus, and suhsigiUatus. Eleutherodactyluspteridophilus sp. nov. Plate 6 Holotype. — KU 1791 06, adult female, one of a series collected at La Delicia, 27 1 0 m (= 00°22' N, 78°25'W), Cordillera delntacProvincialmbabura, Ecuador, on 3 June 1977 by David C. Cannatella and John D. Lynch. Paratypes.— KU 179107-39, USNM 286043, 239848. (See Appendix I for localities.) Diagnosis. — A member of the Elcuthero- dactylus (ElcHtlwrodactylus) iinistrigatus group having ( 1 ) skin on dorsum smooth in males, bear- ing scattered tubercles in females, that on venter areolate; discoidal fold present; dorsolateral folds absent; (2) tympanic membrane absent; tympanic annulus present, its length '74-'/; length of eye; (3) snout short, subacuminate in dorsal view, rounded in profile, bearing terminal keel; (4) upper eyelid bearing warts, narrower than lOD; cranial crests absent; (5) vomerine odontophores oval; (6) males lacking vocal slits and nuptial pads; (7) first hnger shorter than second; Fingers II-IV bearing large, round discs; ( 8 ) fingers bearing fleshy lateral fringes; (9) ulnar tubercles small, conical; (10) heel and outer edge of tarsus bearing small tubercles; inner edge of tarsus lacking tubercle or fold; (11) inner metatarsal tubercle oval, 4-6x subconical outer metatarsal tubercle; supernumerary tubercles at bases of toes; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; (13) dorsum pale to medium brown with darker markings outlined with cream; venter cream with brown flecks forming loose reticulation; posterior surfaces of thighs brown with cream mottling; ( 14) SVLinmales 17.6-25.1 mm(.r = 22.4±0.3,n = 32) mm, in females 31.9-33.9 mm (.? = 33.1, // = 3). Eleuthewdactylus pteridophilus is distinctive among the Eleuthewdactylus in western Ecuador in lacking secondary sex characters and in having an extremely long fifth toe. In pattern and some aspects of tuberculation, it resembles the smaller £. verecundus. Eleuthenxkictylus siopelus also is simi- lar to E. pteridophilus. but it lacks a tympanic annulus. Description. — (n = 26; 23 for proportions of males, 3 for females) Head slightly narrower than body, as wide as long; HW in males 34.6-39.0% ( v = 36.5 ± 0.2) SVL, in females 37.3-40.0% (v = 38.9)% SVL; E-N in males 65.8-86.2% (j = 77.2 ± 1.2) length of eye, in females 76.2-83.7% (x = 81.2)% length of eye; snout about as long as eye, deep (not flattened) (Fig. 45); nostrils weakly pro- tuberant, directed dorsolaterally; canthus rostralis moderately angular, concave; loreal region flat, sloping abruptly to lips; lips weakly flared posteri- orly; upper eyelid 77.8-108.7% (.r = 90.3 ± 1.5) lOD, in females 84.2-100.0% {x = 90.6) lOD; ELEUTHERODACTYLUS IN WESTERN ECUADOR 125 Fig. 45. Eleutherodactyhis pteiidophihis. O', KU 179113. Scale bar = 2 mm. interorbital space flat; upper eyelid bearing low tubercles; supratympanic fold indistinct; tympanic annulus round, indistinct externally, prominent only upon drying, its length 23.7-37.9% (f = 31.3) length of eye, separated from eye by distance slightly greater than diameter of tympanic annulus; postrictal tubercles indistinct; tongue longer than wide, its posterior border not or feebly notched, posterior '/!-% not adherent to floor of mouth; choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores posteromedian to choanae, separated medially by a distance equal to width of one odontophore in males to one half width of odontophore in females, each three times size of choana in females, each bearing two to four teeth in transverse row; males lack vocal slits. In males, skin on dorsum of body smooth except for a few small warts on upper flanks, in females with small warts on occiput and posterior part of dorsum and larger warts on flanks; skin on venter areolate; discoidal fold well anteriad to groin; cloa- cal sheath absent; pair of warts ventral to cloacal opening. Palmar tubercle bifid (inner lobe largest), much larger than oval thenar tubercle; supernumer- ary palmar tubercles numerous, all smaller and less pungent than round, subconical subarticular tu- bercles; fleshy fringe along outer edge of palm and Finger IV; disc on thumb scarcely expanded; those on Fingers II-IV 1 '/2-2 times width of digit proxi- mal to disc; fingers long, slender. When hind limbs flexed peipendicular to body axis, heels touching or barely overlapping; shank 47.0-56.4% (x = 50.8, // = 24) SVL; inner margin of tarsus lacking tubercles or fold; inner metatarsal tubercle not compressed; subarticular tubercles round, not coni- cal; supernumerary plantar tubercles at bases of toes; discs on toes as large as those of outer fingers; tip of Toe V extending to distal edge of distal subarticular tubercle of Toe IV; tip of Toe III extending to distal edge of penultimate subarticular tubercle of Toe IV. In preservative, dorsum pale to medium brown with dark markings consisting of interorbital bar, W-shaped occipital mark, sacral chevron, suprainguinal spots, canthal-supratympanic stripe, and labial bars; dorsal markings continue as diago- nal bars onto flanks; darks bars on limbs feebly oblique, those on flanks narrower than interspaces. In most specimens dark markings on dorsum out- lined with cream; several individuals with tan areas along midline between dorsal spots; most individu- als with unpigmented tympanic region. Belly and throat cream with brown flecks forming loose re- ticulation; cloacal triangle dark brown; posterior surfaces of thighs brown with some cream mot- tling. Measurements of holotype: SVL 33.9. shank 16.5, HW 13.4, head length 11.9, upper eyelid width 3.2, lOD 3.8, tympanic annulus length 1.2, eye length 4.3. E-N 3.6. Coloration in life. — The dorsum is green, tan, yellowish brown, or reddish brown with reddish- brown to brown markings; the side of the head, flanks, and concealed surfaces of the hind limbs are 126 UNIV. KANSAS NAT. HIST. MUS. SPHC. PUBL. NO. 23 washed with yellow. The throat is llesh colored, and the belly is yellowish white; the iris is reddish copper with black flecks. Natural history. — At the type locality in June 1 977, the holotype was found along a trail in cloud forest, whereas 1 0 males were on ferns at the edge of the road at night. In January 1978, JDL found all individuals on ferns along the edge of a road or along edges of pastures. Height of perch site for 23 individuals was 0.2-2.5 m (f = 0.96) above the ground. The use of ferns by this species stands out dramatically in comparison to the low frequencies with which most other Andean Eleuthemclactyliis perch on ferns. No evidence of reproductive activity was noted in June 1977, when the only female that was found was gravid. The collection made in January 1978 contained 20 males having swollen testes, one adult female, and two juvenile females (SVL 15.9 and 16. 1 mm). The adult female is not gravid and did not appear to have laid eggs recently. On 16 January 1978, males were calling; the call is a single note, "whoop." Distribution. — This species is known from only five localities at elevations of 1500-2710 m in the humid tropical regime (2 localities) and the humid temperate regime (3 localities) on the western flank of the Andes in provincias Imbabura and Pichincha, Ecuador (Fig. 46). Etymology. — The specific epithet is an adjec- tive derived from the Greek noun, pteris meaning a "kind of fern," and the Greek adjective, pliilos meaning "loving." Thus, the name means "fern loving" and is used in allusion to the microhabitat selection evident in this species. Remarks. — There is some pattern polymor- phism in Eleuthewdactylus pteridophilus. Five specimens (KU 132629, 179129, 179136-38) dif- fer from the above description in having a broad, pale, middorsal raphe edged in black. Also, pale, black-edged patches occur on the heels of these individuals. In another individual (KU 1791 16), a darker raphe begins in the scapular region and broadens at the level of the sacrum Another speci- men (KU 179108) has a white stripe along the canthus and outer edge of the eyelid continuing as a dorsolateral stripe to the groin. The remaining 3 1 specimens have simple patterns of dark spots, but 0- • E. pteridophilus A E. sohetes Fig. 46. Distribution of two species of Eleuthew- dactylus in western Ecuador. the spots vary in intensity; in some individuals, they are present only as a concentration of dark flecks that provide only a hint of the more common distinct pattern of spots. Although JDL cannot point to evidence, he thinks that E. pteridophilus is a northern replace- ment of the much larger E. rhyiuolopsoides; no other species seems to group with this pair of species. Eleuthewdactylus pyrrhomerus Lynch Plate 8 Eleiitherodactylus pyrrhomerus Lynch, 1976c:310. — Holotype: KU 131606, adult female, from east edge of Pilalo, 2580 m, Provincia Cotopaxi, Ecuador. Diagnosis. — A member of the Eleuthero- dactyhis (Eleutherodactyhis) myersi group having ( 1 ) skin on dorsum minutely tuberculate with low ridges and W-shaped occipital fold, that on venter areolate; discoidal fold absent; dorsolateral folds absent; (2) tympanic membrane and tympanic an- nulus prominent, round, its length '/i-'/s length of eye; (3) snout rounded, acuminate in dorsal view, rounded in profile; (4) upper eyelid bearing one conical tubercles, narrower than lOD; cranial crests absent; (5) vomerine odontophores low, oval; (6) ELEUTHERODACTYLUS IN WESTERN ECUADOR 127 males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs narrow; (8) fingers lacking lateral fringes; (9) ulnar tubercles indistinct; (10) heel lacking small tubercle; outer edge of tarsus bearing small tubercles; inner edge of tarsus bearing elongate tubercle; (11) inner meta- tarsal tubercle elongate, 4x round outer metatarsal tubercle; supernumerary plantar tubercles numer- ous; (12) toes lacking lateral fringes; webbing absent; fifth toe slightly longer than third; (13) dorsum brown with darker brown spots; venter cream with brown reticulation; colorless areas (bright red in life) in axilla, groin, and on anterior and posterior surfaces of thighs; ( 1 4) SVL in males 16.2-18.9 mm, in females 19.8-24.0 mm. Eleutherodactylus pyrrhomerus is most similar to two other members of the E. myersi group — E. floridiis and E. leoni in sharing (or expected to share in the case of E. floridus) red flash colors in the groin and on the concealed surfaces of the thighs. These three species also are alike in their small size, head shape, and absence of dorsolateral folds. Eleutherodactylus floridus differs by having larger digital discs (barely expanded in E. leoni and E. pyrrhomerus), and E. floridus and E. pyrrhomerus are larger and have white testes in contrast to the smaller E. leoni with black testes. Description. — The original description by Lynch ( 1976c) is complete. Coloration in life. — Field notes on living speci- mens from the vicinity of the type locality reveal that the dorsum is brown to grayish brown with darker brown markings; some individuals have a middorsal stripe that is cream with green cast and edged in black. The venter is gray with brown mottling or black flecks; a cream bar across the mental region was noted in some individuals. The axilla, groin, and concealed surfaces of the thighs and shank are red to salmon-red in adults; these flash colors are less evident in smaller individuals and absent in the smallest juveniles. The iris is creamy green with a median, horizontal, reddish- brown streak. Natural history. — In the vicinity of Pilalo, Provincia Pichincha, this species was most com- monly found under stones, logs, and wood chips along roadcuts and in pastures by day. Only two individuals, both adults, were found at night; they were perched on broad leaves 0.5 and 1 .0 m above the ground in cloud forest. Distribution. — Eleutherodactylus pyrrhomerus seems to be restricted to the upper humid montane forest — humid temperate regime (7 localities) and humid subtemperate regime ( 1 locality) at eleva- tions of 2075-3000 m on the western flank of the Andes in provincias Cotopaxi and Bolivar, Ecua- dor (Fig. 43). Remarks. — Inexplicably, males of E. pyrrho- merus are rarely found; the same is true for E. floridus and E. leoni. Eleutherodcictylus quinquagesimus Lynch and Trueb Plate 4 Eleutherodactylus quinquagesimus Lynch and Trueb, 1980:392.— Holotype: KU 167859, adult female, from Quebrada Zapadores, 2010 m. 5 km ESE Chiriboga. Provincia Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) devil lei group having ( 1 ) skin on dorsum smooth anteriorly and shagreen posteriorly, that on venter areolate; discoidal fold prominent; dorsolateral folds on anterior half of body; interocular fold present; (2) tympanic mem- brane and tympanic annulus evident, small, its length '/6-'/4 length of eye; (3) snout subacuminate to rounded in dorsal view, rounded in profile, papilla at tip; (4) upper eyelid lacking tubercles, broader than lOD; cranial crests present; (5) vomer- ine odontophores round in outline; (6) males lack- ing vocal slits and nuptial pads; (7) first finger shorter than second; discs broad; (8) fingers bear- ing lateral fringes; (9) ulnar tubercles conical; ( 10) heel bearing prominent calcar; outer edge of tarsus bearing row of conical tubercles; inner edge of tarsus bearing nonconical tubercle; (II) inner meta- tarsal tubercle oval, 6-8x subconical outer metatar- sal tubercle; supernumerary tubercles only at bases of toes; (12) toes bearing lateral fringes; webbing absent; fifth toe longer than third; (13) dorsum brown with darker brown markings; venter tan with brown mottling; posterior surfaces of thighs chocolate-brown with white flecks; (14) SVL in Ecuadorian males 27.8-30.8 mm, in females 33.6- 40.1 mm. 128 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Eleiitherodactyhis ijniiu/u(ii^c'.simus i.s easily rec- ognized by the presence of an interocular fold. Even if that is not noticed, the species is distin- guished by a prominent calcar on the heel. Some caution must be exercised in identifying this spe- cies by color pattern, because there is geographic variation in color pattern even across the relatively restricted geographic distribution of the species (Lynch and Trueb. 1980). Description. — The original description by Lynch and Trueb (1980) is complete except for variation in coloration (described below) and in adult size. (See Remarks.) Coloration in life. — The individual and geo- graphic variation in this predominately brown frog are documented in the following descriptions, which are arranged from north to south. KU 202625 from 5 km WofLa Gruel, Provincia Carchi: Dorsum black with white marks on flanks and thighs; labial stripe yellow; venter black with white mottling; iris dark reddish-brown (WED, 26 February 1984). KU 132656 and 132777-78 from La Delicia, Provincia Imbabura: Dorsum dark brown (two with large white blotches on head and back); venter black with or without white flecks; one with flanks and anterior surfaces of thighs with black and white bars(SRE, 11 August 1970). KU 179365-68 from La Delicia, Provincia Imbabura: Dorsum brown with black markings and bronze canthal stripe and dorsolateral fold; flanks brown with white spots; throat and belly black with cream and white mottling on belly; posterior surfaces of thighs brown; iris copper with brown flecks (JDL. 3 June 1977). KU 167881-86 from 9 km SE Tamlayapa, Provincia Pichincha: At night, dorsum reddish tan to yellowish tan; throat and anterior part of belly dull yellowish-green with brown mottling; other ventral surfaces, groin. Fingers I and H, and Toes I- III dull orange; iris dark reddish-copper with black flecks (WED, 9 April 1975). KU 167852-69 from Quebrada Zapadores, Provincia Pichincha: At night, dorsum tan to yel- lowish tan with faintly darker markings; by day, dorsum dark brown to dull olive with or without bronze-tan upper lip; venter dull olive-tan with or without dark brown mottling; iris dark reddish- brown; one individual (KU 167852) with dull olive flanks with creamy-yellow spots, dull orange groin, and Fingers I and II and toes I-III orange brown (WED, 3, 4 April 1975). KU 179376-77 from Quebrada Zapadores, Provincia Pichincha: Dorsum brown with touch of green on eyelids and snout and white labial stripe in one; nearly black with pale whitish bronze areas on face in another; white flecks on flanks, upper arms, and face in one individual and white area in groin in another; throat brown, with dark brown mottling in one individual; belly brown or dirty-yellow with brown flecks and spots; undersides of legs pale orange with brown spots; iris deep chocolate-brown (JDL, 8, 9 July 1977). KU 179378-81 from 1 km SW San Ignacio, Provincia Pichincha: Dorsum pale brown to dark brown with copper marks on face; white areas on upper arms, flanks, and concealed surfaces of limbs; posterior surfaces of thighs brown; belly brown with bluish white mottling; iris deep chocolate- brown (JDL, 10 July 1977). Natural history. — All individuals have been found in cloud forest at night. Most were perched on leaves and branches at heights to 2.5 m above the ground, but one was sitting amidst dead bamboo leaves on the ground. No reproductive activity has been noted. However, males with swollen testes and females with mature eggs have been found in January, April, May, June, and July, thereby sug- gesting that reproduction may be aseasonal. In Ecuador, juvenile females (straight oviducts) are less than 27 mm SVL. Distribution. — Eleutherodactylus ipiinqua- gesimus inhabits cloud forests at elevations of 1 4 1 0-27 1 0 m on the western slopes of the Andes in Ecuador and extreme southern Colombia (Fig. 47). Most (8 ) of the localities are in the humid temperate regime; only one is in the humid subtropical re- gime. Remarks. — Specimens of this species from La Planada, Departamento Nariiio, Colombia, are smaller than those from Ecuador; SVLs of adult males from La Planada are only 20.3-24.6 (v = 23.2 ± 0.2, // = 1 7) and of adult females 30.5-34.0 (.f = 32.3±0.4, /z = 9). Eleutherodactylus quinquagesimus is highly distinctive from other species in western Ecuador ELEUTHERODACTYLUS IN WESTERN ECUADOR 129 81" 80° 79^ 78° -1° 0 20 40 60 80 100 Kilometers /^ 30()m ( ^B >4()()()ni sniiw ^ -0° / ^ 4-^ / / r f 1 ,) /• -\ r^ > / /// ^f ;-' -. i°- 81° \ 80° \ \ 1 J ■-" o ■ - 0 ]• E. c/iiinqiiai;esii>uis\ ^9 78° Fig. 47. Distribution of Eleiitherodactyliis quinqitagesimus in Ecuador. but has the fifth toe longer than the third, as in frogs of several species groups. Here we associated E. quinqiiagesimiis with the E. devillei group not because we have pressing evidence for doing so, but because it is not apparent where else the species might belong. We do not consider E. quinqua- gesimus to be closely related to the cis-Andean E. devillei or to E. appendiciilatus, tniebae, or vertebral is on the Pacific versant. Eleutherodactylus quinquagesimus is less easily placed in any of the other species groups in the E. conspiciUatus series. The trivial name, proposed on the fiftieth birth- day of WED, is dated, but there is no provision in the Code of Zoological Nomenclature for keeping up with aging; besides, 65 in Latin is less euphoni- ous. Eleutherodactylus rosadoi Flores Plate 7 Eleutherodactylus roseus — Lynch. 1980b: 180-82. Eleutherodactylus rosadoi Flores, 1988b:34. — Holo- type: MCZ 92937, adult female, from the Rio San Miguel, about 1 km above the Rfo Cayapas, Provincia Esmeraldas, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having ( I ) skin on dorsum bearing many small warts, that on venter areolate; discoidal fold promi- nent; dorsolateral folds absent; (2) tympanic mem- brane and tympanic annulus prominent, its length -A length of eye; (3) snout subacuminate in dorsal view, rounded in profile, with small papilla at tip; (4) upper eyelid bearing one or two conical tu- bercles, as wide as or broader than lOD; cranial crests absent; (5) vomerine odontophores oval in outline, not prominent; (6) males lacking vocal slits; nuptial pads present; testes black; (7) first finger shorter than second; discs broad, rounded; (8 ) fingers bearing weakly crenulated lateral fringes; (9) ulnar tubercles nonconical; ( 1 0) heel and outer edges of tarsus bearing small tubercles; inner sur- face of tarsus bearing tubercle; (11) inner metatar- sal tubercle oval, 4x round outer metatarsal tu- bercle; supernumerary palmar tubercles low; (12) toes bearing narrow lateral fringes; webbing ab- sent; fifth toe much longer than third; (13) dorsum brown with darker brown markings, including large postaxillary blotch; venter cream with brown flecks forming spots and/or reticulum; posterior surfaces of thighs cream with brown mottling; posterior part of flank, adjacent anterior surface of thigh, and concealed surfaces of shank and tarsus unpig- mented;(14)SVLintwomales 16.5 and 16.7 mm, in three females 23.4-25.7 mm. The combination of conical tubercles on the eyelid and heel in such a small frog enables distinc- tion from other species in Ecuador. A few species (e.g., E. latidiscus and E. muricatus) share this combination of characters, but these species are not sexually mature at the size off. rosadoi. Addition- ally, the black mesorchium in E. rosadoi distin- guishes inales from most other Eleutherodactylus. Eleutherodactylus roseus, an equally small frog in which males have black mesorchia, differs from E. rosadoi in having a single tubercle on the upper eyelid, a shagreen dorsum lacking folds, and in lacking a tympanic annulus and nuptial pads. Eleutherodactylus rosadoi also is similar to the equally small E. colonial. These two species are separated readily by snout shape (truncate in pro- file in E. colomai), heel anatomy (large, conical tubercle in E. colomai), and testis color (white in E. colonial). 130 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Description. — The original description by Flores ( 1988b) has been augmented by additional material. (See Remarks.) Coloration in life. — Examination of color trans- parencies taken by Luis A. Coloma of two speci- mens (KU 2 1905 1 and QCAZ 7862) from the Rio Canoi. 4 km S of Pedro Vicente Maldonado, Provincia Pichincha, reveals that in the former the dorsum of the body and limbs is tan with dark brown markings consisting of an irregular interorbital bar. irregular chevron in scapular re- gion, spots in the sacral region, broad canthal stripe, large blotch in the postaxillary region, and transverse bars on the limbs: the lower flanks are yellowish tan. In the latter specimen, the dorsum is reddish tan with a nairow cream middorsal stripe bordered by broader, iiregular dark brown marks; other markings are dark brown and consist of labial bars, broad canthal stripe, large postaxillary blotch, and transverse bars on the limbs: the heels are tan. In both individuals, the iris is pale bronze with small black flecks. Natural iiistory. — The holotype and paratype were in amplexus (Flores, 1988b), as was the pair collected on Isia Gorgona, Colombia. All individu- als have been found at night on low vegetation in lowland tropical rainforest. Distribution. — This species is known from low elevations (<600 m) in northwestern Ecuador and on Isla Gorgona, Colombia (Fig. 43). One of the Ecuadorian localities is in the humid tropical re- gime, and three are in the humid subtropical re- gime. Remarks. — Eleutherodactylus rosadoi remains inexplicably rare in collections. The measurements for QCAZ 7862 and KU 2 1 805 1 . an adult male and an adult female, respectively, are: SVL 16.7, 25.7; shank 8.4, 12.3: HW 7.0, 10.4: upper eyelid width 1.9, 3.1: lOD 1.9, 2.8: tympanic annulus I.l, 1.5: eye length 2.7, 3.3: E-N 2.3, 3.2. Each individual has a minute heel tubercle. The male has no obvious eyelid tubercles, but the female has well-developed tubercles on the upper eyelids. The male has a pale middorsal stripe extending from the level of the anterior edges of the orbits nearly to the level of the sacrum. Initially, this species was confused with E. roseus (Boulenger), known only from lowlands in north- ern Chocoan Colombia (Lynch, 1 980b). Eleuthero- (hictyliis rosadoi is likely to remain poorly known until collectors intensify their efforts on small eleutherodactylines in leaf litter by day and on low vegetation at night. Eleutherodactylus ruidus Lynch Eleutherodactylus riiidits Lynch, I979a:40. — Holotype: AMNH 17390. adult female, from Molleturo, 2317 m, Provincia Azuay, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum warty with W-shaped occipital folds, that on venter areolate: discoidal fold relatively prominent: dorsolateral folds ab- sent: (2) tympanic membrane and tympanic annu- lus absent: (3) snout short, rounded in dorsal view and in profile: (4) upper eyelid bearing flattened warts, as wide as lOD: cranial crests low on poste- rior parts of frontoparietals: (5) vomerine odontophores triangular in outline: (6) males lack- ing vocal slits: nuptial pads present: (7) first finger shorter than second: discs broad: (8) fingers bear- ing lateral fringes: (9) ulnar tubercles absent: ( 10) heel bearing small tubercles: outer edge of tarsus lacking tubercles: inner edge of tarsus bearing fold: (11) inner metatarsal tubercle oval, 2x round outer metatarsal tubercle: supernumerary plantar tu- bercles at bases of toes: (12) toes having lateral fringes: webbing absent: fifth toe much longer than third: (13) dorsum brown with darker markings: venter cream with diffuse brown spots on throat: posterior surfaces of thighs brown: (14) SVL in males 25.8-3 1. 1 mm, in females 37.1-39.8 mm. Eleutherodactylus ruidus is the only species in western Ecuador having the combination of char- acters, tympanic annulus absent and fifth toe mark- edly longer than third. In other species lacking an ear (£. duellmani, gentryi, hamiotae, siopelus, sobetes, and surdus), the fifth toe is only slightly longer than the third. Description. — The original description by Lynch ( 1 979a) is complete and based on all known specimens. Coloration in life. — Unknown. Natural history. — Little is known about this frog. The species has been collected only once from ELEUTHERODACTYLUS IN WESTERN ECUADOR 131 beneath stones in pastureland and scrubby vegeta- tion in an area having little forest in June 1922. At that time, G. H. Tate collected 16 specimens — seven adult males having nuptial pads, seven gravid females, and two juvenile females, the largest of which (SVL 29.0 mm) shows initial signs of devel- oping oviductal convolutions. Distribution. — Eleuthewdactyhis ritidus is known only from the type locality at 23 1 7 m in the humid temperate regime on the Pacific slopes of the Cordillera Occidental in Ecuador (Fig. 43). Remarks. — Lynch ( 1 979a) suggested relation- ships of E. niidus with E. halionotus and E. riveti, species that inhabit paramo and subparamo in south- ern Ecuador. Eleutherodactylus scolodiscus Lynch and BuiTOwes Plate 8 Eleutherodactylus scolodiscus Lynch and Burrowes, 1990:20.— Holotype: IND-AN 1416, adult female, from Reserva La Planada, 1780 m, Municipio de Ricaurte, Departamento Nariiio, Colombia. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) diastema group hav- ing ( 1 ) skin on dorsum shagreen, that on venter areolate; discoidal fold present: dorsolateral folds absent; (2) tympanic membrane absent but tym- panic annulus evident beneath skin, its length about '/? length of eye; (3) snout subacuminate in dorsal view, rounded in profile, with papilla at tip; (4) upper eyelid bearing small tubercles, narrower than lOD; cranial crests absent; (5) vomerine odontophores absent or only barely evident; (6) males having vocal slits; nuptial pads absent; (7) first finger shorter than second; discs broad, with papilla at tip of disc on Finger IIL (8) fingers bearing fleshy lateral keels; (9) ulnar tubercles absent; (10) heel bearing minute tubercle; tarsus lacking tubercles or folds; (11) inner metatarsal tubercle oval, 5x round outer metatarsal tubercle; supernumerary tubercles numerous; ( 1 2) toes bear- ing lateral fringes; webbing absent; discs broad, bearing distal papillae of Toes II-IV; fifth toe much longer than third; (13) dorsum cream with fine black flecks providing hint of markings; venter cream with fewer black flecks; axilla, groin, and concealed surfaces of thighs unpigmenlcd (orange in life); (14) SVL in males 17.9-20.4 mm, in females 18.4-22.3 mm. By having papillae on the tips of some digits, E. scolodiscus can be confused with E. chalceus and E. gularis. Living individuals ofE. scolodiscus are recognizable immediately by the blue iris (black in E. chalceus and E. gularis). Eleutherodactylus chalceus is a larger frog (SVL in males 17.5-26.9 mm, in females 27.7-31.2 mm) and has areolate skin on all surfaces, whereas E. gularis is about the same size as E. scolodiscus but has smooth skin on the dorsum. The larger E. ccdcarulatus has a blue- gray iris, but it differs in having a prominent tym- panic annulus and heel tubercle, and lacking papil- lae on the tips of the digits. Description. — The original description by Lynch and Burrowes (1990) is complete. Coloration in life. — According to Lynch and Burrowes (1990), in living specimens from La Planada. Departamento Narino, Colombia, the dor- sum was pale tan with a pink or orange suffusion; some individuals also had brown flecks on the dorsum. The limbs were tan with brown flecks forming transverse bars in some individuals. The groin and posterior surfaces of the thighs were orange, and the venter was uniform white. The iris was pale bright-blue with fine black reticulations. Description of two Ecuadorian specimens are, as follow. KU 1 77651 from Maldonado, Provincia Carchi: Dorsum pinkish orange; venter somewhat yellow; iris blue (JDL, 17 May 1977). USNM 286292 from Chicai Provincia Carchi: Groin and anterior surfaces of thighs yellow; dor- sum of body and hind limbs with rosey-red (E. W. Schupp, 25 September 1979). Natural history. — Little is known about this recently described species. At the type locality, individuals were found at night on leaves and mossy branches in dense cloud forest and along closed-canopy streams (Lynch and Burrowes, 1990). One Ecuadorian specimen was on vegeta- tion below a waterfall at night, and another came from the canopy of a felled tree by day. Distribution. — Eleutherodactylus scolodiscus is known from few localities at elevations of 1 200- 132 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 1780 m ill tlie luiniid subtropical regime on the Pacific .slopesof the Andes in northern Ecuador and adjacent Colombia (Fig. 44). Remarks. — Specimens from Ecuador (3 males 17.6-19.0 mm SVL; 1 female 20.1 mm SVL) are about the same size as those from the type locality. At present. E. scolodiscus seems to be the nearest relative of the larger E. chalceus, and this pair of species is related to other members of the E. di- astema group {E. diastema, gidaris, hylaeformis, and vocator). The digital papillae of E. chalceus and E. scolodiscus are larger than those of E. gidaris, but are smaller than those in undescribed species in northwestern Colombia and eastern Panama. Eleuthewdcictylus simonholivari Wiens and Coloma Plate 8 Elentliewdactylus siiuonholivari Wiens and Coloma. 1992:197.— Holotype: QCAZ 1459, adult female, from Bosque Protector Cashca Totoras, 3200 m, 10 km SE of Santiago on road to Santa Rosa de Totoras, Provincia Bolivar, Ecuador. Diagnosis. — A member of the Eleuthero- dactyliisiEleittherodactylus) orestes group having ( 1 ) skin on dorsum bearing low warts of inegular sizes (appears more or less smooth), that on venter areolate; discoidal fold evident; dorsolateral folds absent; (2) tympanic annulus not well defined ex- ternally, round, its length V^-A length of eye; (3) snout rounded in dorsal view and in profile; (4) upper eyelid lacking tubercles, narrower than lOD; cranial crests absent; (5) vomerine odontophores low; (6) males having vocal slits; nuptial pads absent; testes white; (7) first finger shorter than second; discs narrow; (8) fingers lacking lateral fringes; (9) ulnar tubercles indistinct or absent; (10) heel bearing small tubercles; tarsus lacking tubercles and fold; (11) inner metatarsal tubercle oval, 1.5x round outer metatarsal tubercle; super- numerary plantar tubercles indistinct or absent; (12) toes lacking lateral fringes; discs narrow, as large as those on fingers; webbing absent; fifth toe longer than third; (13) dorsum brown with darker brown markings; venter orange (in life) in males, brown with pale spots in females; white spots in axilla and groin and on concealed surfaces of limbs; (14) SVL in males 16.0-19.2 mm, in fe- males 18.5-22.0 mm. Eleuthewdactylus simonholivari is most simi- lar to E. orestes Lynch in the Andes of southern Ecuador. The skin on the dorsum in E. simonholivari is smooth or nearly so, but usually it is areolate (rarely smooth with scattered tubercles) in E. orestes. Living males off. simonholivari are reddish brown above and orange below, whereas those oiE. orestes are brown above and gray below. In E. orestes, the groin and concealed surfaces of the limbs are black with large white spots, whereas E. simonholivari has smaller white spots on a black background and appears to have a black venter covered with white flecks (pale areas coixesponding to individual wails). Description. — The original description by Wiens and Coloma ( 1 992) is based on all known specimens. Coloration in life. — As reported by Wiens and Coloma (1992), sexual dimorphism is evident in coloration. They described males as having a red- dish-brown dorsum with darker spots; labial bars brown; venter and digits orange; ventral surfaces of thighs gray; white spots present in axilla and groin and on concealed surfaces of shanks and tarsi. In contrast, females were described as having a dark brown dorsum with or without reddish tint; labial bars brown with white borders; flanks nearly black with minute white spots; venter dark brown to nearly black with paler spots corresponding to centers of pustules; axilla, groin, and concealed surfaces of shank and tarsus black with white spots. The iris is gray with a median, horizontal, gray streak. Natural history. — At the type locality, all indi- viduals were in the vicinity of a spring in upper humid montane forest. The frogs, all adults, were in leaf litter and under rotting logs and under moss on logs by day. Distribution. — This species is known only from elevations of 3000-3300 m in the vicinity of the type locality in the humid subtemperate regime in the Cordillera Occidental in Provincia Bolivar, Ecuador (Fig. 44). Remarks. — Wiens and Coloma (1992) used the results of analysis of molecular data to argue that E. simonholivari was the sister species of E. ELEUTHERODACTYLUS IN WESTERN ECUADOR 133 orestes. They also postulated that these two species were allied to E. trepidotiis and E. ocreatus, but those two species have much shorter fifth toes than do E. orestes, simonholivah, and vidua. Eleuthew- dactylus ocreatus and E. trepidotus share the same toe length state as do the species placed in the E. myersi subgroup (£. gladiator leoni. myersi, pyrrhonierus. and repens) by Lynch ( 1984a). Eleutherodactylus siopelus Lynch and Burrowes EleiitJwrodactylus siopelus Lynch and Burrowes, 1990:22.— Holotype: IND-AN 1563. adult female. from Reserva Natural La Planada, 1780 m, Municipio de Ricaurte. Departamento Naritio, Colombia. Diagnosis. — A member of the Eleuthero- dactylus { Eleutherodactylus ) devil lei group having ( 1 ) skin on dorsum shagreen, that on venter ar- eolate; discoidal fold evident posteriorly; dorsolat- eral folds short, extending to level of sacrum; (2) tympanic membrane and tympanic annulus absent; (3) snout subacuminate in dorsal view, rounded in profile; (4) upper eyelid bearing small tubercle, narrower than lOD; cranial crests present, low; (5) vomerine odontophores triangular in outline; (6) males lacking vocal slits and nuptial pads; (7) first finger shorter than second; discs large; (8) fingers bearing lateral fringes; (9) ulnar tubercles conical; (10) heel bearing calcar; outer edge of tarsus bear- ing row of conical tubercles; inner edge of tarsus bearing low fold on distal quarter; (11) inner meta- tarsal tubercle oval, 3x flat outer metatarsal tu- bercle; supernumerary plantar tubercles numer- ous; (12) toes bearing fleshy lateral fringes; web- bing absent; fifth toe longer than third; (13) dorsum brown with darker brown markings; venter white with brown flecks; lower flanks with cream flecks on darker background; posterior surfaces of thighs brown with cream flecks; (14) SVLin males 18.6- 27.3 mm, in females 34.3^0.0 mm. By virtue of having cranial crests and a calcar on the heel, E. siopelus is distinguished easily from most species of Eleutherodactylus. Those features and the absence of a tympanic annulus distinguish it from all Eleutherodactylus known to us. Eleutherodactylus quinquagesiiuus also has cra- nial crests and calcars, but it has an evident (albeit small) tympanic annulus, and an interocular fold. Description. — The original description by Lynch and Burrowes ( 1990) is based on all known specimens. Coloration in life. — According to Lynch and Bunowes ( 1990), the dorsum is uniformly choco- late-brown or tan with brown chevrons and limb bars; the flanks are brown with varying amounts of white mottling. The venter is white with brown blotches. The posterior surfaces of the thighs are brown in males and brown with small white flecks in females. The iris is golden-yellow. Natural history. — All individuals were perched on vegetation overhanging streams by 0.5-2.0 m in cloud forest at night. Distribution. — Eleutherodactylus siopelus is known only from the type locality in cloud forest on the western slopes of the Andes in extreme south- ern Colombia. In spite of the absence of records from Ecuador, we suspect that the species will be found in Ecuador, at least in northern Provincia Carchi. Remarks. — A single juvenile female (SVL 31.1 mm) is known. The relationships of E. siopelus remain obscure. The initial impression of JDL is that E. siopelus is allied closely to E. acatallelus found at higher elevations (2000-2600 m) in the Cordillera Occidental in departamentos Cauca and Valle del Cauca, Colombia. Our association of E. siopelus {and E. acatallelus) in the E. devil lei group is made on the basis of negative, rather than posi- tive, evidence; in other words, this group is one of convenience. Eleutherodactylus sobetes Lynch Plate 4 Eleutherodactylus sobetes Lynch, I980c:334. — Holo- type: KU 179389, adult female, from Quebrada Zapadores, 2010 m. 5 km ESE Chiriboga, Provincia Pichincha, Ecuador. Diagnosis. — A member of the Eleuthero- dactylus (Eleutherodactylus) surdus group having (1) skin on dorsum smooth with many small pus- tules, that on venter areolate; discoidal fold well anteriad to groin; dorsolateral folds thin; (2) tym- panic membrane and tympanic annulus absent; (3) snout short, subacuminate in dorsal view, rounded 134 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 in profile; (4) upper eyelid lacking tubercles, as wide as lOD: cranial crests present: (5) vomerine odontophores massive, oval; (6) males unknown; (7) first finger shorter than second; discs broad; (8) fingers bearing thick lateral fringes; (9) ulnar tu- bercles absent; ( 10) heel bearing conical tubercle; outer edge of tarsus bearing thin fold with small tubercles; inner edge of tarsus bearing indistinct tubercle; (11) inner metatarsal tubercle oval, 5-6x subconical outer metatarsal tubercle; supernumer- ary tubercles low, at bases of toes; ( 1 2) toes bearing lateral fringes; webbing absent; fifth toe longer than third; (13) dorsum brown with bold brown markings; dorsolateral folds cream; venter cream with brown spots peripherally; posterior surfaces of thighs brown; (14) SVL in one adult female 41.3 mm. Eleiithewdactylus sobetes is distinguished readily from other species in the E. surdus group because it has distinct pustules on the skin of the dorsum and a conical tubercle on the heel. Further- more, the digits of E. sobetes are longer and more slender than those in other species of the group. (See Fig. 1 inLynch, 1980c.) Living individuals are readily identified by having an orange iris. Eleiithewdactylus degener also has an orange iris but is a much smaller frog having smooth skin on the dorsum and a much longer fifth toe. Description. — The original description by Lynch ( 1980c) is based on all known specimens. Coloration in life.— KU 179289-90 from Quebrada Zapadores, Provincia Pichincha: Dor- sum dark brown and dull olive-yellow; groin and posterior surfaces of thighs pale violet; venter dirty-yellow with brown markings; iris bright or- ange (JDL, 9 July 1977). Natural history. — The two known specimens were found in the same ravine ( Quebrada Zapadores ) in cloud forest at night in 1 977; one was on a branch 1 m above the forest floor, and the other was on low vegetation beside a stream. Although the locality was searched extensively in 1977 and has been visited many times by other biologists, no addi- tional specimens have been found. There was noth- ing peculiar about the habitat or behavior of the frogs in 1977 to suggest why the species should seem to be so rare. The second specimen is a juvenile female with a SVL of 34.4 mm. Distribution. — This species is known only from the type locality at an elevation of 2010 m in the humid temperate regime on the western flank of the Andes in Ecuador (Fig. 46). Remarks. — Lynch (198()c) suggested that E. sobetes was allied to E. duellmani and E. surdus, a position with which we have no disagreement. However, Lynch's ( 1980c) hypothesis concerning several Andean species is weak and rejected here. That hypothesis placed too much emphasis on the absence of a tympanic annulus and ignored several species described subsequently from Colombia. Furthermore, if we are coiTect in interpreting the variation in the length of the toes as a systematic character, then E. banecuus and E. ruidus have no relationship with the other frogs that Lynch in- cluded in the E. surdus group. Eleiitherodactyliis subsigillatiis (Boulenger) Plate 6 Hylodes subsigillatus Boulenger, 1902:52. — Holotype: BM 1947.2.17.1, adult female, from Salidero, Provincia Esmeraldas, Ecuador Eleutherodoctylus subsigillatus — Peters, 1 955:339. Diagnosis. — A member of the Eleuthero- dactylus {Eleutherodactylus) unistrigatus group having ( 1 ) skin on dorsum smooth to finely shagreen, that on venter areolate; discoidal fold prominent; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus evident, round, its length '/4- -/? length of eye; (3) snout subacuminate in dorsal view, rounded in profile, bearing papilla at tip; (4) upper eyelid usually lacking tubercles, (occasion- ally 1 minute tubercle), slightly narrower than lOD; cranial crests absent; (5) vomerine odontophores oval to subtriangular in outline; (6) males having vocal slits and white nuptial pads; (7) first finger shorter than second; outer fingers bear- ing broad discs (larger than tympanic annulus); (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles low, if visible; (10) heel bearing small conical tubercle; outer edge oftarsus bearing vague tubercles; inner edge of tarsus bearing tubercle; (11) inner metatarsal tubercle oval, 5-6x round outer metatarsal tubercle; supernumerary plantar tubercles numerous; (12) toes bearing lateral fringes; webbing absent; fifth toe much longer than third; ELEUTHERODACTYLUS IN WESTERN ECUADOR 135 ( 1 3) dorsum pale tan with brown markings, becom- ing darker on flanks; venter white to cream with scattered dark flecks or loose reticulum; lower flanks and anterior surfaces of thighs with dark brown or black bars or reticulum enclosing white spots; ( 14) SVL in males 19.3-28.5 mm, in females 30.0-33.4 mm. Eleuthewdactylus subsigiUatus is most similar to£. eugenicie, nyctophyUi.x, vindphoxocepluilus. It is most easily distinguished from those species by the pattern on the lower flanks and anterior surfaces of the thighs — barred in E. suhsigillatus, cream or white with brown reticulations in E. eugeniae and E. phoxocephaliis and brown with cream flecks in E. nyctophylax. Of these four species, only E. eugeniae and E. subsigillatiis have a terminal pa- pillae on the snout; E. phoxocephalus has a vertical keel, and the snout in E. nyctophylax is unadorned. The tubercles on the upper eyelid and heels of E. suhsigillatus are small and easily overlooked; of the four species discussed here, the only other having tubercles on the eyelids and heels is E. nyctophylax. Description. — The species was redescribed in detail by Lynch (1980b). Coloration in life. — The general appearance of E. suhsigillatus is tan, brown, or dull green; varia- tion is documented in the following descriptions: KU 131566-70 from Balsapamha, Provincia Bolivar: Dorsum tan to dark brown with green and white flecks; throat dusky; rest of venter dirty cream with brown flecks or reticulations; posterior surfaces of thighs black with white flecks; anterior surfaces of thighs dirty gray and black with cream to pale yellow spots; iris chalk-white with black reticulations and soft bronze, median, horizontal streak; concealed edge of eye pale orange to yeflow (JDL, 17 July 1970)^ KU 177839^3 from Santo Domingo de los Colorados, Provincia Pichincha: Dorsum various shades of green and brown (one with decidedly orange cast on posterior part of body and hind limbs); throat pale green, yellow, or gray; rest of venter usually pale yellow (white in 1 'm4(IU(lm f ■ ,: Maldonado ©v ' ^ [ZJ Permanent k .,-■, ■ LaDelicia*^ \' -l' -0° Sto. 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"3 "^ •js Q a Q a a o ^ ^ u = 11 ^ a to 00 s>. -ii S O "iJ K -.^ -S S S ? a 00 t;; § c s ^ cos 2 ^^ Sec ki tjj Uj Uj t^ I-d Uj Uj I4J IjJ IjJ Ijj Ijj (jj Ijj Ly Ijj Lq Uj Uj Lij Ltj Uj (jj Lq Uj [jg 152 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 O > X o tLi < o o U c« — 'G (U \D o. S. 00 3 « H I I I I I I I I I I I I I I I > I I I I I I I I I + I I I I + I I I I I I I I + I + I I I I "+ I II + lll + l + lllllll -+ + +^^l++l + l I + I + I + I + 1^;. + + I + + + + + + +"+ + lllllllll + ll + l + l 4.+ I I + + + + + + + + + + + I + + + +I I I I I + I++I+ + I 1 I I I I I I I I I I I I I + I++I + I I l + l I I++I l + l + l 1 I I I I+ + +I I l + l l + l + l I I I I++I + 1 + Ill + I + I IIII++IIII1II + I + + +IIIIIIIIIII + I I + IIIIIIIIIIII+ + + +IIIIII + I + 1III + I + IIIIII + I + IIII + 00 ri-] ir] rn — • ^ O — ' ^ "^' f^' <^j ^ '~~~ '^ "~; rn c-i r^! <^. n r^i ri ^ <^, ^ r^. ^ C-l ^' '^ <"< --H o fN -^ m o r^J •— ^ ^ 2 5 5 ;~; Co -^ s . , o ex Ci, a, Ci. IS 00 2 ^ ^ uj!jjuji^[jjtjjkikikjtjjt±iuikit^tti'^ J= CI. _aj "S 0 c 0 c« '0 C/D c (U C 13 £ S ELEUTHERODACTYLUS IN WESTERN ECUADOR 153 4. Tandapi, Provincia Pichincha. 1460 m, {)0°24' S, 78°47' W: Humid subtropical regime. Mean annual rainfall 1 800 mm; mean annual tempera- ture 15°C. Disturbed cloud forest and clearings with a fast flowing river and small streams. Twelve species. 5. Maldonado, Provincia Carchi, 1 4 1 0 m. 00°54' N, 78°06' W: Humid subtropical regime. Mean annual rainfall 1000 mm; mean annual tempera- ture 16°C. Disturbed cloud forest with some pastures and three small streams. Eight species. 6. Mindo, Provincia Pichincha, 1410 m, 00°02' S, 78°46' W: Mean annual rainfall 3466 mm; mean annual temperature 16.4°C. Humid subtropical regime. Steep slopes with cloud forest and small streams. Sixteen species. 7. Quebrada Zapadores, Provincia Pichincha, 20 1 0 m, 00° 15' S, 78°43' W: Humid temperate re- gime. Rainfall and temperature unknown, but at Chiriboga, 5 km W of Quebrada Zapadores at 1960 m, mean annual rainfall 2003 mm; mean annual temperature 16.2°C. Deep ravine with cloud forest and high-gradient stream. Ten spe- cies. 8. Pilalo, Provincia Cotopaxi, 2400 m, 00°56' S, 58°59' W: Humid temperate regime. Mean an- nual rainfall 1490 m; mean annual temperature 12.6°C. Disturbed cloud forest with many bro- meliads; adjacent clearings and small streams. Six species. 9. La Delicia, Provincia Imbabura, 2710 m, 00°22' N, 78° 25' W: Humid temperate regime. Rainfall and temperature unknown. Steep slopes sup- porting cloud forest; many terrestrial bromeli- ads (along roadside) and one small stream. Ten species. In the following paragraphs, we address abun- dance, body sizes, food, and microhabitat of the species found at these nine localities. Species diversity. — The numbers of species at the nine sites range from six to fourteen. Compari- son of species occurrence among sites was accom- plished by calculating the coefficient of biogeo- graphic resemblance (CBR; Duellman. 1990). The highest CBR (0.92) is between the two lowland sites, Rio Palenque and Santo Domingo de los Colorados, which has all 1 1 species known from Rio Palenque (Table 7). Only two to four species (CBR = 0.1 7-0.32 ) are shared between the lowland sites and three of the sites in cloud forest in the humid subtropical regime. However, five lowland species (E. achatinus, cholceus, miiricatus, pan'illus, and wcdkeh) present at both Rio Palenque and Santo Domingo de los Colorados and also E. necerus at Santo Domingo de los Colorados are among the 14 species at the lowest site in cloud forest, Rio Faisanes (1380 m). Three of the four sites in cloud forest in the humid subtropical regime also have high CBRs (0.64-0.77). The three sites (Rio Faisanes, Mindo, and Tandapi ) closely approximate one another geo- graphically and altitudinally, whereas the fourth Table 7. Occurrence of species of Eleutherodactylus in nine communities in western Ecuador. Abbreviations in headings to columns correspond to sites in first column. The number of species in each community is shown in boldface in the common cell; the numbers of species that are shared by two communities are shown in the upper right, and the coefficients of bigeographic resemblance are in italics in the lower left. Site PAL SDC RFA TAN MAL MIN QZA PIL LDE Rio Palenque (PAL) 11 11 5 2 2 3 0 0 0 Santo Domingo (SDC) 0.92 13 6 3 3 4 0 0 0 Rfo Faisanes (RFA) 0.40 0.44 14 10 4 10 T 0 2 Tandapi (TAN) 0.17 0.32 0.77 12 3 9 3 1 3 Maldonado (MAL) 0.21 0.30 0.36 0.30 8 4 2 0 2 Mindo (MIN) 0.22 0.28 0.67 0.64 0.33 16 T 1 3 Qda. Zapadores (QZA) 0.00 0.00 0.17 0.27 0.22 0.15 10 ■) 5 Pilalo (PIL) 0.00 0.00 0.00 0.11 0.00 0.09 0.25 6 2 La Delicia (LDE) 0.00 0.00 0.17 0.27 0.24 0.24 0.50 0.25 10 154 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 site, Maldonadi), although at the same elevation as Mindo. is about 75 km NE of Mindo and separated from the other sites by the Rio Chota Valley; the CBRs between Maldonado and the other sites are 0.30-0.33. The three sites in cloud forest in the humid temperate regime (Quebrada Zapadores. Pilalo, and La Delicia) share no species with the lowland sites and few species (CBRs = 0.00-0.27) with the sites in cloud forest in the humid subtropical re- gime. Four species (£■. ilitelhmini, quimiiiagesimns, vertehnilis, and w-nignmi ) are shared by Quebrada Zapadores and La Delicia, whereas two species (£. phoxocephalus and E. vertebmlis ) are common to all three high cloud forest sites. The CBRs among these three high sites are 0.25-0.50. No attempt was made to determine actual abun- dance of species. However, during visits at five sites, all individuals observed were collected; these values range from 50 specimens of eight of the 1 1 species at Rio Palenque to 372 specimens of eight of the 12 species at Tandapi. During these visits, selectivity in collecting was minimized; although biases of preferred microhabitat (of frogs and col- lectors) and of collecting schedules versus activity cycles of the frogs exist, they are relatively constant among sites. Other variables also exist — the time spent at each site and the times of the year that sites were visited. Thus, we have utilized these data only to provide crude estimates of relative abundance and rank order of abundance (Table 8). Differences in sampling effort at the five sites resulted in vastly different numbers of individuals at some of the sites. This discrepancy in the data can be overcome by application of Krebs' (1989) rare- faction algorithm, which extrapolates the number of species that can be expected in a given sample size. Based on this algorithm, the numbers of species expected in a sample of 30 specimens at each site are: Rio Palenque 7, Tandapi 6, Quebrada Zapadores 6, Mindo 5, and Pilalo 4. Only one taxonomic comparison can be made with rank abundance data for Rio Palenque with other sites; the forest-edge species E. achatinus is ranked third in abundance at the lowland site (Rio Palenque) and second and fourth at two sites in cloud forest — Tandapi and Mindo, respectively. Among the sites in cloud forest, six of the nine u lij C^ r- < H .S oj in On a, u. N < ON o -- .= a- s < Oh "^ 3 ^ 3 -— 52 ~ ~ c ^ ^ 1^ 2 »^ ^ ^ j^' p* Ci. a is "^ cC uj uj Uj ijj ui <-) <"J ^^ ~ '^ On >: -2 ON ip, 52 •^ I 5^ '0 ■^ .2 ^ I ^Sc S- S ^ S .s y^ .^ Cl. "^ 2? >J 0 1 a: Ci. 2; -s: 3: G- s^ •^ S ^ Ci, ui ui tii ki tij uj tij 2 ^ .2 :- r "^ ^ ^ ri - ^■croo;::2:~^^~=:'o~ g ^ s a -2? ^ ~ a ^ -G- kikjUJtijliitijkitjjUJljq n-, rr-, 00 - f^' ^. _ - - — r- -^ — t^ a s ?• -s;' S s ■t; - « s ^ "s p 5 v^ .' O P^. C n3 ;_ .„ .^ bjujtiiijjiiiljjujtq ■2 — — .2 — — ri c: !** r^ Co 1— >; >3 ^— ' :3 -c -5 ri „ 10 ^^ \D ^-^ ^ r<", trj '■^ i^ to ';::; ^ lo !0 'C ■S ■2 "a 2 to So -2 0 5 -2 uj tij tij tij ki tij Pi Uj Lq -— " n r<-i -^ in NO r-~ 00 On O ELEUTHERODACTYLUS IN WESTERN ECUADOR 155 species at Tandapi are among the 10 species at Mindo, and one species at Tandapi is among the seven species at QuebradaZapadores, which shares one species with Pilalo. Rankings among the six species shared by Tandapi and Mindo are entirely different. Eleutherodactyhis \v-nigrnm is the most common species at Tandapi, but it is ranked fifth at Mindo. where E. calcarulatiis is most abundant (fourth at Tandapi). Likewise, the second-ranked species at Mindo. E. hiteolatendis, is sixth at Tandapi, and the third-ranked species at Mindo. E. crenunguis, is the least abundant of eight species at Tandapi. Eleutherodactyhis w-nigrum is the only species shared by Tandapi and Quebrada Zapadores. and it is the most abundant species at both sites. Eleutherodactyhis phoxocephcdus is the most abun- dant species at Pilalo; this species is shared with Quebrada Zapadores, where it is the least abundant species. In most cases, there is no ready explana- tion for differences in relative abundance, at least among sites in cloud forest. However, the differ- ence in abundance and ranking of £. phoxocephalus at Quebrada Zapadores and Pilalo might relate to the great abundance of arboreal bromeliads, the diurnal retreat of E. phoxocephalus, at Pilalo, as compared with the scarcity of these plants at QuebradaZapadores. Abundance may change with time. The greater abundance of E. luteolateralis at Mindo than at Tandapi ( in 1 967 ) may reflect recency of disturbance at Tandapi; the species was more common there in 1968 and 1970 as regrowth oc- curred (Lynch 1976a). It is obvious from the data presented in Table 9 that great unevenness exists in the abundances of species in all five communities. The ratios of rela- tive abundances of the most abundant : second- most abundant species vary from 1 : 0.35 to 1 : 0.70 (Table 9); in no case do these represent the same pair of species. The ratios of the most abun- dant : least abundant species range from 1 : 0.01 to 1 : 0.05; again the same pairs of species are not involved. Data on biomass are not available, but extrapo- lation of mass from body size reveals no clear relationships between size and relative abundance or rank of abundance, as illustrated for four of the communities (Fig. 55). At Tandapi and Quebrada Table 9. Ratios of relative abundance of four most common species oi Eleutherodactyhis in five commu- nities in western Ecuador. Community Relative abundance Rfo Palenque Tandapi Mindo Quebrada Zapadores Pilalo 0.35 0.30 0.25 0.70 0.66 0.57 0.54 0.36 0.27 0.51 0.24 0.18 0.49 0.13 0.05 Zapadores, the largest species, E. w-nigrum, is the most abundant, whereas at Mindo the second small- est, at Pilalo the second largest, and at Rio Palenque the third largest species are the most abundant. The smallest species are the least abundant at Mindo and Pilalo. Body size. — Hutchinson (1959) suggested a critical size ratio of 1 .3 for coexistence of species in the same trophic level. Subsequently, two ideas regarding size and coexistence promulgated the literature (reviewed in Simberloff and Boecklen, 1981): ( 1 ) There is a minimum size ratio compat- ible with coexistence of ecologically similar spe- cies. (2) Three or more ecologically similar species tend to have constant size ratios between species adjacent in size-ranking, but the constant factor may vary among sites. Ratios (larger SVL/smaller SVL) were com- puted for adjacent-sized species in the nine com- munities oi Eleutherodactyhis in western Ecuador; as examples, six of these are illustrated graphically in Figures 56 and 57. Clearly there is a linear relationship that falls between ratios of 1 .00 and 1 .50. The few exceptions are females of the larger species — E. anomalus at Rio Palenque and E. crenunguis and E. necerus at Tandapi and Mindo. The size ratios between adjacent species is not constant; there tend to be minor differences among smaller species, whereas the differences are greater among larger species. When these data are treated on a log-log scale, the differences are reduced. Food. — No analysis of stomach contents was undertaken, but cursory examination of stomach contents revealed diverse kinds of insects ranging from ants to orthopterans, and also spiders. Among the species that we have examined, there is no ant specialist, such as E. acuminatus in the Amazon Basin (Duellman, 1978). 156 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Tandapi 60 50 40 30 20 • O • o • ° • 80 c > I Quebiada Zapadores O Rio Palenque o s C/5 70 60 50 40 30 20 10 Mindo O J L 2 3 456789 10 12 3 456789 10 Rank order of abundance Fig. 55. Rank order of abundance and body size among Eleulherodactylus in four communities in western Ecuador. Solid dots are males; open circles are females. ELEUTHERODACTYLUS IN WESTERN ECUADOR 157 > Q. on 50 4U 30 20 Rio Palenque / / '^y o / / *o <§^/ / o •^ / *^ / Tandapi / / i _ ^ / - 7 7 / o / /o°v Av Av A/ : . Pilalo / / 40 - 30 20 Ri'o Faisanes / ^/ / O/ /o / / * ° h / // , , ! , , : , / ° Maldonado / / / / o / / /• /• / 10 10 20 30 40 50 60 70 80 90 10 20 30 40 50 60 70 80 90 Larger Species (SVL in mm) Fig. 57. Ratios of body sizes in succeeding larger species of Eleutherodacryliis in two communities in western Ecuador Solid dots are males; open circles are females. communities, three (E. achatinus, actites, and walkeh) inhabit clearings and (with E. phoxo- cephalus and E. pteridophilus) forest-edge situa- tions. Nine other species (E. chalceus, muricatus, nyctophylax, ornatissimus, parvillus, surdus, thymalopsoides, unistrigatus, and w-nignmi) also are found at the edge of the forest but range into the interior of the forest. All other species are exclsuively forest inhabitants. The most obvious microhabitat distinction for nocturnal activity is between the terrestrial and arboreal species. Four large (.v SVL ac 47.0-54.2 mm, 99 69.0-98.6 mm) toadlike species (E. anatipes, anomalus, helonotus, and necenis) are strictly terrestrial and usually found on the banks of streams. Adult females off. w-nignim{x Sy\^59 .^ mm) are terrestrial. Six other species most com- monly are on low vegetation, but some individuals have been observed on the ground; three of these (E. crenunguis, longirostris, and surdus) are al- ways near streams; two others (E. duellmani and E. loustes) usually are in the spray zone of waterfalls, whereas the sixth, E. unistrigatus, is wide ranging. Only four species (E. eremitus, phoxocephalus, scolodiscus, and suhsigillatus) are known to occur on vegetation at heights of 4 m or more; of these, only the latter has been found on low vegetation (< 1 m). Many arboreal species have been found only, or most commonly, on vegetation along and over streams; these include E. actites, calcarulatus, crenunguis, crucifer, dissimulatus, labiosus, longirostris, luteolateralis, muricatus, nyctophylax, sobetes, surdus, and males of w-nigrum. However, the apparent abundance of these species at, or their ELEUTHERODACTYLUS IN WHSTHRN ECUADOR 159 Fig. 58. Nocturnal microhabitats of 1 1 species oi Eleiithewdactylus at the Estacion Biologica Ri'o Palenque, Provincia Los Rios, Ecuador. restriction to, streams may be simply the use of such streams as "trails"" by collectors. Nocturnal observations notwithstanding, our experiences with several species suggest that bro- meliads are a microhabitat utilized for diurnal retreats. Especially noteworthy are E. crucifer in arboreal bromeliads at Tandapi. E. phoxocephahis in arboreal bromeliads at Pilalo, and E. celator in terrestrial bromeliads at La Delicia. Likewise, in dry subtropical areas, E. cajamurcensis is abundant by day in small arboreal bromeliads. The apparent rarity of E. eremitus and, in some places, E. suhsigiUatiis may reflect inadequate sampling of bromeliads, especially where they are inaccessible without special equipment. Discussion Our data on the ecology of Eleuthewdactylus in western Ecuador have severe limitations and are sufficient solely to generalize and point out areas 160 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 59. Nocturnal microhabitats of 13 species of Eleutherodactylns at Santo Domingo de los Colorados, Provincia Pichincha, Ecuador. for future research. Studies need to be carried out throughout the year (and for 2 or more years) at specific sites in various environments (e.g.. low- land tropical rainforest and cloud forest), where different assemblages of species exist. The only such existing studies are those by Duellman ( 1 978 ) at Santa Cecilia, Ecuador; Schluter (1984) at Panguana. Peru; Burrowes (1987) at La Planada, Colombia; Heyer et al . ( 1 990 ) at Boraceia in south- eastern Brazil; and Rodriguez (1992) at Cocha Cashu. Peru. All of these studies included Eleutherodactylns in analyses of the ecology of anuran faunas; only that by Burrowes (1987) in- cludes species that occur in western Ecuador, and only that by Rodriguez ( 1 992) contains real data on abundance. A crude estimate of abundance can be obtained by comparing the numbers of museum specimens that exist for each species (Fig. 63). Specimens of the six most abundant species (in descending or- der— E. achatinus, w-nigrum, walkeri, longirosths, actites, and pho.xocepholus) constitute more than ELEUTHERODACTYLUS IN WESTERN ECUADOR 161 Fig. 60. Nocturnal microhabitats of 12 species of Eleutherodactyhis at Tandapi, Provincia Pichincha. Ecuador. 50% of the total. At the low end of the spectrum, four species (E. babax, colomai, helonotiis, and sobetes) are represented by only two specimens each and three (E. apiciilatiis, degener, and hectiis) are represented by three specimens each. However, this representation incorporates several biases. First, some species simply are more conspicuous than others; this is especially true of larger species (e.g., E. actites and E. w-nigrum) that inhabit the edge of the forest, where they are more easily detected by eyeshine than smaller species in more secluded situations in the interior of the forest. The six species represented by more than 200 specimens inhabit, but are not restricted to, the forest edge. In the case of species determined to be abundant at a given locality, individuals frequently were not col- lected during subsequent visits to that locality; thus, the large numbers of these specimens may under-represent the actual abundance of these spe- cies. The apparent rarity of some species, even at localities that have been worked repeatedly, pre- sumably indicates that the species actually does occur in low densities or that it utilizes a microhabi- tat not usually investigated by collectors. However, differences in densities of some species seem to defy explanation; for example, E. caprifer is abun- dant in the spray zone of a waterfall at the type locality, but rarely is found in similar microhabitats at other sites. In the absence of any evidence for interspecific competition, the disparities in relative abundances and rank-abundances of given species in the five communities analyzed are inexplicable. The appar- ent decline in relative abundance of E. achatinus from the lowland site, Rfo Palenque (mean annual temperature 24.4°C), to the upland sites, Mindo and Tandapi (mean annual temperatures 16.4°C and 15.0°C, respectively), may reflect the species 162 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Quebrada Zapadores, 2010 m Fig. 61. Nocturnal microhabitats of 10 species of Eleiitherodactyhis at Quebrada Zapadores, Provincia Pichincha, Ecuador. having reached the upper limits of its distribution at these upland sites where possibly, it has reached the lower limits of its temperature tolerance. As an effect of low temperatures, the species might re- quire more time to develop mature eggs, thereby reducing the number of times individual females can breed during the year. Similarly, there probably is a negative correlation between temperature and rates of ovarian development (as noted for Eleutherodactylus coqui in Puerto Rico by Townsend and Stewart [ 1 986] ), and growth; thus a longer time would be required for individuals to reach sexual maturity. Because there is an inverse relationship between temperature and metabolic activity, suboptimal temperature regimes would affect growth in a species, as well as its ability to capture prey and escape predators (Miller and Zoghby, 1986; Putnam and Bennett, 1981). Any one of these factors, or a combination thereof, may be responsible for lower abundances oiE. achotinus at higher elevations. A completely different situation apparently ex- ists with E. w-nigrwn, which is the most abundant species at Tandapi and Quebrada Zapadores but fifth in rank-abundance at Mindo; all three sites are in cloud forest. At Quebrada Zapadores, E. w- nigrum is the largest species oi Eleutherodactylus, but at Tandapi and Mindo it is barely exceeded in size by E. crenunguis. Eleutherodactylus cremmguis the least abundant species at Tandapi and third in rank-abundance at Mindo, where E. necerus, ninth in rank-abundance, is a much larger species. Eleutherodactylus necerus and adult females of £. w-nigrum are terrestrial (but the former is exclu- sively streamside); also, individuals off. crenunguis sometimes are active on the ground. Possibly in the terrestrial microhabitat, these three species use the same food resources and compete with one another. ELEUTHERODACTYLUS IN WESTERN ECUADOR 163 Fig. 62. Ecuador. Nocturnal microhabitats of 10 species of Eleiitherodactylus at La Delicia. Provincia Iinbabura, The reversal in relative abundance off. w-nigrwn and E. crenungiiis at Tandapi and Mindo might have resulted from other factors that differentially influence the abundances of these species at those sites. Throughout the lowlands and lower cloud for- ests in western Ecuador, the most obvious correla- tion of body size of Eleutherodactylus is with terrestrial versus arboreal habits. All 10 nocturnal, terrestrial species {E. anatipes, anonuilits, babax, cerastes, helonotiis, longirostris, loitstes, lymani, necerus, and w-nigrum) in these habitats are large (J SVL 99 46.7-98.6 mm, x = 65.1 ). In contrast, most arboreal species have SVLs less than 35 mm; only four arboreal species (E. actites, creminguis, labiosus, and thymalopsoides) have SVLs greater than 50 mm (x SVL 99 50.4-61.9 mm, .r = 56.1). However, the one diurnal species {E. hectus, x SVL 99 20.7 mm) is terrestrial, as are £■. /eon/ (j SVL 99 2 1 .9 mm; E. myersi group) and E. simonboUvari {x SVL 99 20.6 mm; E. orestes group) at high eleva- tions in upper cloud forest and subparamo. Thus, within any given assemblage, most species are arboreal and have SVLs of 1 5-35 mm in males and 20^5 mm in females, and considerable overlap in body sizes exists among these arboreal species within assemblages (Fig. 55). The ratios of body sizes of species making up different communities do not conform to Hutchinson's ( 1959) predictions. The results con- form to those obtained for a community of prima- rily arboreal species of Eleutherodactylus at Santa Cecilia, Ecuador (Duellman, 1978) and for Eleutherodactylus in the southern Andes of Ecua- dor (Lynch. 1979a). The ratios of all three of these studies are less disparate than those for members of 164 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 • ' ! 1 \ 1 • 900 800 - 700 '• ^ 600 C3 3 ■O > - 500 - T3 C <*- ^ 400 _ • i-i o x; - E |300 - • ~ • • 200 • % 100 V 0 10 20 30 40 50 Number of Species 60 Fig. 63. Relative abundance of 61 species of Eleuthewdactylus in western Ecuador based on numbers of specimens collected. the Eleutherodactylus ''fitzingeri group" in the Chocoan region (Lynch and Myers. 1983). This apparent discrepancy probably is because Lynch and Myers compared ratios among species through- out the region and, thereby, included many allopat- ric species; furthermore, they were using data only on a small segment of the entire Eleuthewdactylus fauna in the region. Diets of neotropical frogs are poorly known, and those studies in which analyses of stomach contents were undertaken (e.g., Duellman, 1978) have included identifications of prey only to the ordinal or familial level. Such crude distinctions probably do not reflect accurately prey discrimina- tion by the frogs. With few exceptions (none known in western Ecuador), Eleutherodactylus are insec- tivorous generalists that employ a sit-and-wait feed- ing strategy. Generally, there is a positive correlation be- tween body size and prey size (Toft, 1 985 ), but this generalization has limited application to frogs such as Eleutherodactylus, most of which seem to be feeding generalists. True, the generalization ap- plies to the maximum size of prey that can be captured and ingested, but frogs do not necessarily forage only on maximum-size prey. For example, the large, predaceous frog Ceratophiys cornuta is capable of consuming large orthopterans and small vertebrates; these make up 75% of the volume of prey eaten, but ants constitute 70% of the prey items consumed (Duellman and Lizana, 1994). Small arthropods compose the bulk of diets of many species of Eleutherodactylus. including all large, broad-headed taxa, dissected by JDL. Moreover, juveniles of larger species of Eleutherodactylus grow through size ranges of adults of smaller species; these juveniles would be expected to forage on prey of the same size being eaten by adults of smaller species. The common occurrence of juveniles of some arboreal species (e.g., E. achatmus and E. walkeri) in the leaf litter on the forest floor may be indicative of a greater abundance of appropriate-sized prey (ants, ter- mites, and small beetles) on the ground than on the bushes. Such habitat segregation by age and size would eliminate potential competition for food by juveniles and adults. Within given microhabitats, broadly sympatric species usually occur together. Microhabitat spe- cialists tend to be segregated geographically. For example, three species that inhabit seepages in the upper cloud forest {E. duellmani, hamiotae, and surdus) do not occur sympatrically. Of course, Eleutherodactylus exist in communi- ties containing other species of frogs, and consid- eration of resource utilization must include these species. If we view most of the species of Eleutherodactylus as small members of a noctur- nal-arboreal-insectivore guild, most other mem- bers of that guild in the lowlands and Andean slopes ELEUTHERODACTYLUS IN WESTERN ECUADOR 165 in Ecuador are centrolenids. Duellmaii and Burrowes (1989) identified 20 species of centrolenids on the Pacific versant of Ecuador and southern Colombia. At lower elevations (<15()0 m), small to moderate-sized species of HyUi (e.g.. H. alytolylax and H. camifex ) also are members of that guild. However, because of their breeding habits, the hylids and centrolenids are nearly al- ways collected in the immediate vicinity of streams, a subarea of the forest frequented by many species oi Eleutherodactylus. Because of their similarities in size, structure, and microhabitat utilization, we suggest that sev- eral pairs of species are ecological replacements either altitudinally or latitudinally. The pairs E. labiosus-cremmguis, E. latidiscus-laticlavius, E. nyctophylax-eugeniae, and E. walkeri-litteo- lateralis are examples of apparent low elevation- high elevation replacement. The pairs E. ocellatiis- crenunguis and E. vertehralis-truebae are examples of apparent north-south latitudinal replacement. Not all instances ofecogeographic replacement are self evident. For example, E. vv-nigrnm is a large, conspicuous, and usually abundant species through- out the upper cloud forest. However, within its latitudinal range, the species is absent at Pilalo, Provincia Cotopaxi, where apparently it is replaced by E. actite.s, which is known only from that area. Until far more diverse and precise information is available, most explanations of ecological asso- ciations of Eleutherodactylus in western Ecuador must be considered as highly speculative. Now that the taxonomy of these frogs has been studied thor- oughly and the species can be identified with more confidence than was possible in the past, we hope that detailed ecological studies will be undertaken to address the many fascinating aspects of the ecology oi Eleutherodactylus in western Ecuador. BIOGEOGRAPHY Assuming that most of the species of Eleuthero- dactylus in western Ecuador are now known and that a reasonable perception of their geographic ranges and microhabitats exists, we attempt to synthesize the available data on biogeography of these frogs. Although we are concerned principally with western Ecuador, data from extralimital re- gions, especially Chocoan Colombia, have been included. Patterns of Dlstribution Data used for the biogeographic synthesis in- clude ( 1 ) species groups, as defined in the section Subgenera and Species Groups, (2) body size, (3) microhabitat, (4) macrohabitat (bioclimatic regimes, as defined in the section on Western Ecuador), (5) altitudinal range, (5) latitudinal distribution, and (6) area of geographical distribution (Appendix IV), together with the 681 locality records for 61 species (Appendix I). For purposes of this synthe- sis, we use mean SVL of females for body size. Microhabitat is defined as the usual place of activ- ity; all species except E. hectus are nocturnal. Bioclimatic regimes. — The distribution of spe- cies in the six bioclimatic regimes inhabited by Eleutherodactylus in western Ecuador is highly uneven (Table 1 0). Also, the distribution of collect- ing sites is uneven: the large number of sites in the humid subtropical and humid temperate regimes reflect the accessibility of these sites and collec- tors" proclivities for spending more time at sites where frogs are common (perhaps also the higher comfort levels of doing field work in the subtropics. as opposed to the hotter lowlands and the colder highlands). Only five species are known from the dry tropi- cal regime; E. achatinus is represented by 1 1 records, E. longirostris by two, and the others by one each. No species is restricted to that regime; instead, their occurrence in the dry tropics is peripheral to their distributions in the humid tropics. Three of the species are in the E. unistrigatus group; one is in the E. conspicillatus group, and one in the E.fitzingeri group. The mean body size is 37.0 mm (2 1 .6 mm in E. walkeri to 48.6 mm in E. longirostris). The latter is a terrestrial streamside species; the other four are arboreal and have a mean body size of 34.2 mm (2 1 .6 mm in E. walkeri to 42.3 mm in E. latidiscus). Three of the five species occurring in the dry tropical regime are among the seven species in the dry subtropical regime, which again seems to be a 166 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 peripheral habitat for species that are more wide- spread in other environments. However, in western Ecuador, E. lymani occurs only in the dry subtrop- ics, but elsewhere it also occurs in the dry temper- ate regime. Three of the species are in the E. conspicillatus group, three in the E. imistrigatus group, and one in the E. diastema group. Body size ranges from 2 1 .6 mm in E. walkeh to 60.2 mm in E. lymani {x - 38.2 mm). Only E. lymani is terres- trial. Among the six arboreal species, body size ranges from 2 1 .6 mm in E. walkeri to 59.8 mm in E. w-nigrum (v = 34.6). Rather surprisingly, only 15 species have been recorded from the humid tropical regime; none of these is restricted to the regime. These species belong to eight groups, of which the E. unistrigatiis group is represented by eight species; the E. cerasinus and E. conspicillatus groups have two species each, whereas the E. anomalus, diastema, and fitzingeri groups have one species each. Size ranges from 2 1 .6 mm in E. walkeri to 85.4 mm in E. anomalus (J = 37.4). Only two species are terrestrial — E. anomalus (SVL 85.4 mm) and E. longirostris (SVL 48.2 mm); both are streamside inhabitants. The mean size for the 13 arboreal species is 32.9 mm (21 .6 mm in E. walkeri to 50.4 mm in E. labiosus). By far, the greatest number of species (46, 75.4%) inhabits the humid subtropical regime; 12 species (26.0% of the total) are restricted to this regime. All species groups, except the E. curtipes group, are represented in this assemblage of spe- cies. The range in size is from 20.7 mm in E. hectus and E. scolodiscus to 98.6 mm in E. anatipes (.? = 40.2). Nine species are terrestrial; the mean SVL in these is 61 .3 mm (20.7 mm in the diurnal E. hectus to 98.6 mm in E. anatipes). Five of the terrestrial species are streamside dwellers, and these gener- ally are large frogs; the range in size is 46.7 mm in E. loustes to 98.6 mm in E. anatipes {x = 73.4). Most species (34) are arboreal; these range in size from 20.7 mm in E. scolodiscus to 61 .9 mm in E. crenunguis (x = 35.0). Three arboreal species — -E. caprifer (42.7 mm), E. duellmani (41.8 mm), and E. surdus (45.8 mm) — seem to be restricted to spray zones of waterfalls. The second largest assemblage of species (33, 54. 1 %) is in the humid temperate regime, but only seven species groups are represented — cerasinus ( 1 species), conspicillatus (3 ), curtipes ( 1 ), devil lei (4), myersi (3), surdus (5) and unistrigatus (17). The total range in size is from 21.6 mm in E. pyrrhomerus to 59.8 mm in E. w-nigrum ( x = 34.5 ). Three species are tenestrial and have sizes of 21.9 (E. leoni) to 36.0 mm {E. hamiotae); the latter is a streamside species. The mean body size of 30 arboreal species is 35.5 mm (21.6 mm in E. pyrrhomerus to 59.8 mm in E. \\'-nigrum)\ two of these species — E. duellmani (41.8 mm) and E. surdus (45.8 mm) — occur only in the spray zones of waterfalls. Only eight species inhabit the cold humid sub- temperate regime. Six species groups are repre- sented— curtipes (1 species), devillei (1), myersi (2), orestes (1), surdus (1), and unistrigatus (2). Body size ranges from 20.6 mm in E simonholivari to 45.8 mm in E. surdus ( v = 3 1 .0). Three terrestrial species have sizes of 20.6 mm in E. simonholivari Table 10. Distribution of species of Eleutherodactylus in bioclimatic regimes in western Ecuador. Bioclimatic Regime Dry Dry Humid Humid Humid Humid Parameter tropical subtropical tropical subtropical temperate subtemperate Total Locality records 16 28 87 363 167 20 681 Percent of records 2.3 4.1 12.8 53.3 24.5 3.0 100 Number of species 5 7 15 46 33 8 61 Percent of species 8.2 11.5 24.5 75.4 54.1 13.1 — Endemic species 0 1 0 12 7 1 — Percent endemic 0.0 14.3 0.0 26.0 21.2 12.5 — ELEUTHERODACTYLUS IN WESTERN ECUADOR 167 to 33.5 mm in E. gentryi {x = 25.3). Body sizes are noticeably larger in the five arboreal species, in which the mean SVL is 34.4 mm (21.6 mm in E. pyrrliomerus to 45.8 mm in E. siinlus). Some patterns are evident with respect to spe- cies groups. Of the three species groups of the subgenus Craugastor, the E. anomalns and E. hufonifonuis groups are restricted to the humid tropical and humid subtropical regimes, whereas the sole member of the E. fitzingeri group in the region (E. longirostris) also ventures into riparian situations in the dry tropical regime. Of the 12 species groups in the subgenus Eleuthewdactylus, two {E. conspicillotus and E. unistrigatus ) encom- pass all six regimes. The E. cerasinus and E. diastema groups are essentially restricted to the humid tropical and humid subtropical regimes; one record of a member of the E. diastema group {E. gulans)is, from the dry subtropical regime, and one record for the E. cerasinus group is from the humid temperate regime. All members of the E. dolops, loustes, and sidcatus groups are confined to the humid subtropical regime. Of the other five groups, the E. devillei, myersi, and surdiis groups are pri- marily inhabitants of the humid temperate and humid subtemperate regimes, but some members of all three groups enter the humid tropical regime. The E. curtipes and E. orestes groups are distrib- uted primarily in the humid subtemperate regime. In general, specific fidelity to bioclimatic re- gimes is rather low; the highest amount of ende- mism (26.0%) is in the humid subtropical regime, followed by the humid temperate regime (21.2%), dry subtropical regime (14.3%), and the humid subtemperate regime (12.5%); the dry tropical and humid tropical regimes lack endemics. The great- est number of shared species (22) is between the humid subtn)pical and humid temperate regimes, and 1 4 species are shared by the humid tropical and humid subtropical regimes. Although fewer num- bers of species are shared by other regimes, the highest coefficient of biogeographic resemblance is between the dry tropical and dry subtropical regimes (Table 1 1 ). In five of the regimes, arboreal species account for more than 80% of the eleutherodactylines in the regime; the exception is the humid subtemperate regime, where only 62.5% of the species are arbo- real, and most of these are most commonly found by day under stones. Only one species, E. longirostris, seems to be associated with streams in the dry regimes, and only one, E. surdus, does so in the humid subtemperate regime. Terrestrial species associated with streams and arboreal species asso- ciated with spray zones of waterfalls are most numerous in the humid tropical, subtropical, and temperate regimes, where they account for 20.0%, 17.4%, and 9.6% of the species, respectively. Little variation in mean female body size is evident among arboreal species in the six regimes. Terrestrial species generally are larger than arbo- real species in each regime, except in the humid temperate and subtemperate regimes, where terres- trial species are smaller. This is owing to the fact that tenestrial species in these regimes are mem- bers of the E. myersi group, all of which are small frogs. Variation in body size is discussed more fully in a following section on altitudinal distribution. Table 11. Distribution of species of Eleutherodactylus in six bioclimatic regimes in western Ecuador. Abbreviations in headings to columns correspond to regimes in first column. The number of species in each regime is shown in boldface in the common cell; the numbers of species that are in common to two regimes are shown in the upper right, and the coefficient of biogeographic resemblance are in italics in the lower left. Regime DTR DST HTR HST HTE HSE Dry tropical (DTR) 5 4 5 6 1 0 Dry subtropical (DST) 0.67 7 5 5 1 0 Humid tropical (HTR) 0.50 0.45 15 14 2 0 Humid subtropical (HST) 0.24 0.19 0.46 46 22 2 Humid temperate (HTE) 0.05 0.05 0.08 0.56 33 6 Humid subtemperate (HSE) 0.00 0.00 0.00 0.07 0.31 8 168 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Latitudinal distribution. — The eleulhero- daclyline fauna in western Eeuador represents the southern terminus of the rieh fauna on the Pacirte lowlands and the western slopes of the Cordillera Occidental in Colombia (hereinafter referred to as the Chocoan fauna). The latitudinal gradient from humid tropical forest to dry tropical forest in the lowlands of western Ecuador and the lower, drier mountains in the Huancabamba Depression in south- ern Ecuador are reflected in the diminution of species south of 1 °S Lat. in the lowlands and in the highlands (Fig. 64). Among the species oi Eleiitherodactylus inhab- iting the Pacific versant of South America, none has a distribution that includes both Colombia and Peru. Three species in western Ecuador {E. cajamarcensis. lynuiiii, and pho.xocephalns) range into northern Peru. Thirty-two species are endemic to western Ecuador. Another 24 species of Eleuthewdactylus reach the southern limits of their distributions in western Ecuador. Of these, two species primarily distributed at elevations less than 1000 m (E. achatimts and E. kmgirosths) range northward into Panama. Fourteen other species range for varying distances northward into Colom- bia; eight of these (£. anatipes, anonialus, caprifer, chalceiis, gularis, labiosus, latidiscus, and subsigillatus) are strictly Chocoan and have their major distributions on the lowlands and lower (<1000 m) slopes of the Cordillera Occidental of Colombia. Three other species, E. babax, cerastes, and ocellotiis, are primarily Chocoan at intermedi- ate elevations (1200-1800 m), and E. diiellmani is Chocoan at higher elevations (1500-2700 m). Ten species {E. apiculotiis, appendiculatus, colomai, degener, eremitus, hectiis, laticlavius, loustes, quinquagesimus, and scolodiscus) known from in- termediate elevations in Ecuador barely range into Colombia; they are known there only from La Planada ( 1 ° 1 0' N, 1 780 m) and/or slightly lower elevations on the Pacific slopes of the Cordillera Occidental. Two other species, £■. itnistrigatiis and E. w-nigriim, range northward at elevations of more than 2000 m into various ranges of the Andes of Colombia. One other high-Andean species, E. leoni, is not yet known from the western slopes of the Andes in Colombia, but this species also occurs in the Cordillera Oriental of Ecuador and Colombia (Lynch and Duellman, 1 980). Among the 32 species endemic to western Ecua- dor, 25 are restricted to the area within one degree of the equator; however, except for La Planada, the frog fauna of Departamento Narino in southwest- ern Colombia has been poorly sampled. One of these species, £". rosadoi, also occurs on IslaGorgona off the coast of Colombia, but it is unknown from the mainland of Colombia. Of these 27 species, nine (e.g., E. niuricatus, ornatissiimis, and tenebrionis) inhabit, but are not restricted to, low- lands below 1000 m. Of the 18 species that are distributed solely above 1000 m, nine are at eleva- tions above 1500 m, and three of those do not descend below 2000 m. Of the other seven species endemic to western Ecuador, only E. walkeri is restricted to the lowlands, where it ranges to 3°43' S Lat. Two other species, E. crucifer and E. pon'illus, are distributed primarily at intermediate eleva- tions; the former ranges to I ° 1 6' S Lat. and the latter to 2°34' S Lat. The other four species (E. pyrrhomerus, ruidus, siinonbolivari, and tniebae) occur only at elevations above 2000 m south of the Equator. Altitudinal distribution. — The great amount of physical relief in western Ecuador provides opportunities for considerable altitudinal distribu- tion (Figs. 2, 3). Records for species of Eleuthewdactylus in western Ecuador range in elevation from 20 to 3400 m. Eleuthewdactylus w- nigrum has the greatest elevational range (800- 3200 = 2400 m), whereas five species are known from a single elevation (Fig. 65). Fifteen species that have their lowest limits below 500 m have elevational distributions of 520-1 780 m(.r= 1067), and nine species that have their highest limits above 3000 m have elevational distributions of 300-2400 m (v = 1095); even by removing the widespread E. \v- nigrum from the latter group, the distributions are 300-1640 m (r = 932). Twenty-three inhabitants of cloud forest having the limits of their distributions between 1000 m and 3000 m have elevational distri- butions of 20-1560 m ( J = 8 1 6); four other species in this elevational range are known only from one locality. Thus, it seems that species that range into the lower elevations have greater altitudinal distribu- tions than those at the highest elevations, which in turn have greater elevational distributions than those species restricted to intermediate elevations. ELEUTHERODACTYLUS IN WESTERN ECUADOR 169 .r40' rio' rio' 1 ° I y 1 10' \'\y 2'30' 6'45' 10' N- rio' 3°50' 2°40' 7° 12' r20'N- 6°45' 5°20' 4° 10' rio' no' I'lO'N- 8°05' 6°45' 6° 15' .VOO'N- 9°00' 7° 12' i. anatipes E. loustes E. scolodiscus ■ E. colomai " £. hectus "■ £. degeiier ^^^^^^^ £. ocellatiis ^^^^^""■■i^" £. cerastes ^^■^^^^^^^^ £. rosadoi ' £. helonntiis ^■^■^^^^^^^^" £. celdtor ' £. hamiolae . , ^^" £. (ipiciilatiis , I ^^" £ /('^'/)/ . I ' £. sobetes \ ^^^ £. laticlovius | ^^^" £. caprifer \ | ■^^^ £ illoliis I I ^^^■■" £ (luellmani \ ^^^^ £ hiteolateralis \ ^^^^^ £ haha.x ' \ ^^^"^ E. pterklophilits \ ^^^^■" £. crenwigitis \ ^^" £ eiigeniae i ^^ £ dissinmlatus i ^^^^^ £ imistrigatus i ■^^^^" £ surdus\ I ^^^^^■■^ £. anomalus i ^^^^^^" £ latidiscus i ^^^■^^^ £. labiosus i ^^^^^^^ £ tenehrionis i ■^^ £ muhcatits i ^^^^^^^^ £ appcndiculatus ■^^^^^^^ £ ereniitus ^^i^^^^^" £ verecumius i ■^^^^^^" £. necerus i ^■^^^^^ £ nycfophyla.x i ^^^^^^■^^ £ calccirulatiis i ^^^^^^^■■« E. floridiis ^^^■■^^^^ £ cpiic/uagesimus ^am^^^a^^^^^ £. onuitissimus ^^■^^^^^^^^" £. vertebralis ' £. actites i ' £. gentryi ' £ thymalopsoides ^^^■■^^^^^^^"■^^ £ cruclfer ^^^^^^^^^" £. pyrrlwmenis ' E. siirionbolirari ^■■^■■^^^^^^^^^^■^^^^■^^^^^^ £ longirostris ^^^^mi^^mmmmmmmmmmmmmmmmmmmmmm^^^^ E. chalceus mmmmmmmmmmmmmmmmmmmmmmmmmmmm^^^^mm^ £. guUiris ' ' ' ^^^^^"^^^ £ truebae ^■i^^^^^"^^^^^^^^^^^^^^^^^^^^^^^ £. pel nil I us ^^m^mmm^^^m^^^^^^^^^^^^^^^m^^^^^^^ £ siibsigillatlis • £ ruidits E. phoxoceplialiis E. cajanmrrcnsis E. Ixnuini E. mhatituis E. w-nigriini ■ £ walkeri •5 23' S •6 3()'S ■S 34'S FN rs 2^s Decrees Latitude 3°S 4°S Fig. 64. Latitudinal distribution of species of EleutheiodactyUis in western Ecuador. Shaded bars indicate major discontinuities in ranges. 170 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Upon examining altitiidinal distributions by lOO- m increments, the greatest concentration otspecies is between 1200 m and 2200 m, and 28 species occur at elevations of 1800-1 900 m (Fig. 66). Only 10 species occur below 200 m. and an equal num- ber occur at, or above, 3000 m. Mean body sizes are highest between 500 m and 1500 m; this is diminished somewhat if the four species of the subgenus CraKgastorare eliminated (Table 12). In that case, the range of mean body sizes of frogs of the subgenus Eleuthewdactylus in 500-m elevational increments varies only from 34.5-37.3 mm. Thus, the somewhat larger mean body sizes at lower elevations are dictated by the presence of large species of the subgenus Craugastor. Throughout the elevational range of the genus in western Ecuador, most species are arboreal (Table 1 3 ). However, only 50% of the species at elevations above 3000 m are arboreal; the highest percentage of terrestrial species (37.5%) is in the high Andes. Only five species are terrestrial streamside inhabit- ants; three of these {E. anatipes, cmomalus, and longirostris) are members of the subgenus Craugastor and occur only at elevations below 1500 m. Other terrestrial streamside species in- clude E. loHstes (E. hastes group) that is known from 1200-1410 m and E. hamiotae {E. siirdus group) that is known only from 2140 m. Of the three species that are arboreal in the spray zones of waterfalls. E. caprifer (E. conspicillatus group) is confined to elevations below 1000 m, whereas the other two species (E. duellmani and E. siirdiis) are members of the E. surdus group and do not descend below 1500 m. Different patterns of altitudinal distribution are evident among species groups of Eleutherodactylus in western Ecuador, although members of the E. unistrigatus group ( 27 species ) and E. conspicillatus group ( 6 species ) extend throughout the elevational range (Table 14). The four members of the subge- nus Craugastor {E. anomalus, bufomformis, and fitzingeri groups) are distributed primarily below 1500 m, but E. necerus (E. bufomformis group) ascends to 1540 m. Likewise, members of the E. cerasinus and E. diastema groups primarily occur at elevations of less than 2000 m. However, one member of the E. cerasinus group (£". crenunguis) is restricted to intermediate elevations of 800- 1 640 m. whereas another member (E. ocellatus) occurs at elevations of 1255-2560 m. The two species in the E. sulcatus group occur at lower intermediate elevations — E. cerastes at 500-1200 m and E. helonotus at 1410 m. Each member of the E. dolops and E. loustes groups is restricted to intermediate elevations — E. babaxai 1 550- 1 780 m and E. loustes at 1200-1410 m, respectively. Five species groups are restricted to higher elevations. Of these. E. gentryi (E. curtipes group) is distributed at 2850- 3380 m. Of the four species in the E. devillei group, only two descend to elevations below 1500 — E. appendiculatus to 1460 m and E. quinquagesimus to 1410 m. Two species in the E. myersi group are restricted to intermediate elevations, whereas the other two species occur primarily at elevations in excess of 2000 m. The four members of the E. surdus group range from 1550 to 3190 m. and the sole member of the E. orestes group (E. simon- bolivari) occurs at elevations of 3000-3300 m. Trans-Andean distributions. — Three species that occur on the western slopes of the Andes in Ecuador also inhabit the Amazonian slopes in Ec- uador and Colombia. Eleutherodactylus leoni oc- curs at elevations of 1960-3400 m on the western slopes and at elevations of 2540-2700 m on the Amazonian slopes (Lynch and Duellman, 1980). The ubiquitous E. w-nigrum occurs at elevations of 800-3200 m on the western slopes and at eleva- tions of 1 100-2540 m on the Amazonian slopes of Ecuador and also ranges far northward into Colom- bia. Eleutherodactylus unistrigatus primarily in- habits inter-Andean valleys, but apparently iso- lated populations exist at elevations as low as 2400 m on the western slopes of the Andes and at 2700 m on the Amazonian slopes, where also a contigu- ous inter-Andean population descends the Pastaza Trench to an elevation of 1 800 m. Size of distributions. — Determination of areas of distribution was accomplished by measuring the area of the distribution of a species on a map by closely encircling peripheral localities. Although this method assumes a continuous distribution, it is a repeatable method. For species known from only one locality, the area was calculated as 10 km-; for those known from only two localities, the distance between the localities was measured, and a breadth ELEUTHERODACTYLUS IN WESTERN ECUADOR 171 I i.tiyM/txfuoiiiis- 'J ^ sapiosdojiyuixiii -^ srju^iuoifjjXd -jj snpin.t -^j m saiaqos -^ sNinputiisyip -^ snjvifdajoxoiid -^ sisuaD-ivuivfuj -^ Sl]D.iq9l.lJ.\ 'J , snjnin.iidn z] .loivj^-^ '3 I ,inuiry^n,j ■^ snpyjns "^ luniuj/^np ^7 stipifdopi.it)id -^ xuquq J s;)ijDpiJipioddv -Tj ^^.^^ sniuisBSnnbmnb tj ^^^ snjouoj^i/ ■;j sniAVjDiinj ■^ smopi 3 smopajo ■^ snuaif 21 srijsipojojs -^ s9isnoi ^7 snujjn.iDJjvj -Tj XVl\Lldi)J.y\U 3 sijD.i^wio3jnj 2 snputiJMSA 3 wnj8iu-M ■■^ .I3U3E3P ^7 I IDUIOJO.l ■^ 1 xinh'iimiMJ \5 iinmixj :j . S3dntmv -^ sdiaii.i,^.') J snumsumuo 3 snw^utnu '3 srisoiqnj :/ snioi.icjiiu^i 7 smViifSisqns' :/ snjmuoim j , jafi.tdv.i "7 mt^^^ lopnso.i '7 SmiflDljJD -J snajpiif.y j sunjuii yj snjsipijnj '7 Sl.llSO.II^'llOI '■] (LU) U01JBA913 172 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Western Slopes M eten Eastern Slopes Andes and lowlands Restricted to ranges east of Andes 4()()() 3500 I I I I I M I I I ! I I M M I I 25 20 15 10 5 Number of Species 0 15 20 25 Number of Species Fig. 66. Numbers of species of Eleutherodactylus at 1 00-m increments on the Pacific lowlands and western slopes of the Andes in Ecuador (including La Planada in extreme southwestern Colombia) compared with species at the same elevations in the Amazon Basin and on the eastern slopes of the Andes in Ecuador. Mountain ranges east of the Andes include the Cordillera del Condor, Cordillera de Cutucu. and Volcan Sumaco. ELEUrHERODACTYLUS IN WESTERN ECUADOR 173 tiJ — o o o O "O — O o o m o — O o o o in — O o o >n o — (N — o o o O in 58 >n o o o o d oo ON in I «n <^ I ^d O J- ^ as m O) 00 00 00 I ^ d 00 __ ir-, p <"' J- S (N in ^ ON d J- "^ X) f2 r3 ^ s c S s 3 C V 0£ =: Qi 174 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Table 13. Microhabitats utilized by EU'iitlwiodactylus at different elevations in western Ecuador. Numbers are percents within given elevations. Microhabitat Elevation Arboreal Terrestrial (meters) n Arboreal (spray zone) Terrestrial (streamside) 0-500 15 80.0 6.7 0.0 13.3 501-1000 23 69.6 4.3 8.7 17.4 1001-1500 29 75.9 0.0 13.8 10.2 1501-2000 35 80.8 5.7 11.4 2.9 2001-2500 24 83.3 8.3 4.2 4.2 2501-3000 21 81.0 9.5 9.5 0.0 3001-3500 8 50.0 12.5 37.5 0.0 of 2 km was assigned. In two cases in which single outliers of a broad distribution exist (£. chalceus and E. gularis). the outlier was not included in the area. With a few exceptions (£. cajamarcensis. lymani, Knistrigatiis. and w-nigrum), the most striking as- pect of distributional area is the correspondence with altitudinal distribution. Eleven species (E. achatifius, anomalus, caprifer, chalceus, gukiris, labiosus, latidiscus, longirostris, rosadoi, subsigillatus, and wolkeri) have the lower limits of their altitudinal distributions at, or below 150 m; for these species, areas are 3277-39,687 km- (r - 16,963). In contrast, 20 species {E. apiculatus, appendiculatus, babax, calcarulatus, celator, cru- cifer, duellmani, eremitus, eugeniae. Hiatus, laticlavius, luteolateralis, nyctophylax, ocellatus, phoxocephalus, ptehdophilus, quinquagesimus, scolodiscus, surdus, and vertebral is) known from three or more localities and having their lower limits of distribution at 1000-2000 m have areas of 354-8707 km- (.r = 2306). Likewise, species dis- tributed at elevations greater than 2000 m have even smaller areas; five species {E. gentryi. leoni, pyrrhomerus, simonbolivari, and truebae) have areas of 50-2287 km- (.v = 1032). Size of area of distribution is determined not only by longitudinal distribution, but also by altitu- dinal distribution. The lowland species ranging from Panama to southern Ecuador have the largest areas — E. achatinus (39.687 km-) and E. longirostris (31,1 83 km-) (Fig. 67). A few species having long latitudinal distributions have rather narrow altitudinal ranges on the Andean slopes and, consequently, have much smaller areas of distribution; examples are E. babax (4262 km-) and E. cerastes (4194 km-) (Fig. 68). As expected, the 27 species endemic to the Andean slopes between 1 °N Lat. and 1 °S Lat. have rather small distributional areas. The sizes of their areas of distribution are 10^783 km- (J = 1438). Eight of these species are known from only one or two localities; the areas of distribution of the 19 species known from three or more localities are 50- 4783 km- (J = 2030). The exceptions to the general pattern of area of distribution and altitudinal distribution include two species that are distributed primarily in the Huancabamba Depression in northern Peru and southern Ecuador and that reach the northern limits of their distributions in western Ecuador. Both E. cajamarcensis and E. lymani inhabit locally humid areas within this region and have fragmented distri- butions within the region. Consequently, although the calculated areas of distribution (20,532 km- for E. cajamarcensis and 23,559 km- for E. lymani) encompass the outermost known ranges of the species, the actual areas of distribution are greatly exaggerated. The ubiquitous E. w-nigrum in most areas of cloud forest at elevations of 800-3200 in the Andes north of the Huancabamba Depression has an esti- mated area of distribution of 27,266 km-. With a few exceptions (e.g., vicinity of Pilalo, Provincia Cotopaxi, Ecuador), this large species is distrib- uted on all slopes of the Andes north of the depres- ELEUTHERODACTYLUS IN WESTERN ECUADOR 175 w ttl ti c e* o c« _u « > ex "S !/3 m 03 ^ a_i o c3 c« b^ f/5 lU S j> 'T CJ — 3 c« H o U I o — o O lO O m I O — ' o o o lO in I o — o O IT) O CN I o — o o o I/-) — I o o o a- 3 O a o o o o >n V) "1 o o o V-) >o o d d d d 04 r4 (N d d d (N (N d o o o o r^ o o <0 o o o CM O O o OOOOOmoOr<-jOOO>/-jOO OOOmmO\0000000 rJ r<-j O "O lAi >/-i r~- — [^ m IT) VD J^ o o On o r^ r«-) O M2 ON On o NO o o r^ ^ o o o d o o d o o ^ d O o6 O n 00 [^ lO O r^ in NO 1 o ^_ ^ O r<-i r*-! O o o O O d o d r^ — ' U-) O m r<-) O O O (N m d d d ^ rj o o o ^ n r^ O t^ rn r<-i NO d NO rn m O O O C ^ in O O ro — in (-<-) oor~;OOOooo odNodddddd ooooooooo m CN-^-^TtND — '^(N — ^ ^ (N "* r- s ^ 05 to •2? ^ ^ ^ 'H ^ a C to o tn ^ to ^j T3 .jo 1 3 -2 s s; 0 S to c s a -C) ^ u o u "a 176 UNIV. KANSAS NAT. HIST, MUS. SPEC. PUBL. NO. 23 Fig. 67. Distribution of Eleutherodocryius achatimis as an example of a broadly distributed species in the Chocoan lowlands. Note the disjunct populations in the lower Rio Cauca Valley in northern Colombia and along the Pacific coast of western Ecuador. Fig. 68. Distribution of Eleutherodactylus cerastes as an example of a widely distributed species on the slopes of the Cordillera Occidental in Colombia and Ecuador. The hiatus in the range is at the Rio Patfa Valley. sion (Fig. 69). Eleutherodactylus uuistrigatus also is a widely distributed Andean species with an area of distribution of 1 6.524 km- mostly encompassing inter-Andean valleys but also extending onto the western and eastern slopes of the Andes in Ecuador and southern Colombia (Fig. 70). There is no clear correlation between body size and area of distribution. Excluding E. cajamarcensis and E. lymani for the reasons given above and using mean S VL of females as a measure of size, the 20th- ranked species in size (largest to smallest) is E. achatimis with a SVLof 41 .2 mm; it has the largest area of distribution (39.687 km-). The second larg- est area of distribution (31,183 km-) is that of E. longirostris, which is the lOth-ranked species in size (SVL 48.2 mm). Nonetheless, the 10 largest species have areas of distribution of 1 0-3 1.183 kin- ( X = 8757 ) in contrast to 20- 1 9. 1 26 km- ( x = 4060) for the 10 smallest species. Also, there is no significant difference between the areas of distribution of ten'estrial and arboreal species. With the eliinination oi E. cajamarcensis and E. lymani, the areas of 1 2 terrestrial species are 1 0-3 1,183 km- ( x = 5845 ) compared to 1 0-39.687 km- (J = 5214) for 47 arboreal species. Patterns of Speciation Although the phylogenetic relationships are not known for all of the species of Eleutherodactylus in western Ecuador, ceitain groups and pairs of sister species have been identified; among these taxa ELEUTHERODACTYLUS IN WESTERN ECUADOR 177 E. unisthgatus Fig. 69. Distribution of Eleiitherodactyhis w- nigrum in cloud forests in the northern Andes. This is, by far, the most widespread distribution of a species of Eleiitherodactyhis inhabiting cloud forest. there are repeated biogeographic patterns. How- ever, precise knowledge of relationships (species level cladograms for three or more species) is not generally available. Phylogenetic hypotheses in- volving only pairs of species cannot be used to suppoit some general hypothesis, because two- species cladograms cannot be combined into a general cladogram ( Nelson and Platnick, 1 98 1 ). Of all of the species of Eleiitherodactyhis in western Ecuador, only one (£. loustes) is pail of a published three-taxon statement (Lynch, 1992a). The ab- sence of explicit phylogenetic hypotheses means that we cannot examine modes of speciation as discussed by Lynch ( 1 989b) and are limited here to discussing possibilities. Nevertheless, the general pattern is that nearest relatives (albeit in two-taxon statements) are allo- Fig. 70. Distribution of Eleiitherodactyliis imistri- gatus as an example of a widespread distribution in the Andes of Ecuador. patric and have relatively large geographic ranges (and estimated populations). This pattern supports Lynch's (1989b) contention that speciation by pe- ripheral isolation is rare in the speciose genus Eleiitherodactyhis in western Ecuador. Eleiithero- dactylus gentryi is the only potential example of speciation by peripheral isolation we see in this fauna. Chocoan region. — The northern part of the Chocoan lowlands has an eleutherodactyline fauna that essentially is part of that in lower Central America. For example, the region is inhabited by E. hiifonifonuis, i^aigei, and ridens. all of which range northward at least into Costa Rica; these species and some endemics (e.g., E. zygodactylus) do not range south of the Rio San Juan. However, two species, E. achatiniis and E. longirostris, range throughout the Chocoan lowlands from Panama to 178 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 southern Ecuador. The Chocoan lowlands to the south of the Rio San Juan are inhabited by many taxa(e.g.,£. lahiosus, latidiscus, and scolodiscus). The lowlands in Departamento Narifio in south- western Colombia have not been collected inten- sively. It is likely that several species known now only from the Pacific lowlands of Ecuador will be found there; these include E. colomai, E. degener, and E. rosadoi. Among the patterns that are evident is that displayed by the E.fitzingeri group of the subgenus Craugastor. Of the eight species in this group, which has five species endemic to Central America, two species (£. fitzingeh and E. raniformis) are distributed in lower Central America and the north- em part of the Choco, whereas E. longirostris ranges from Panama to southern Ecuador (Lynch and Myers, 1983). Among other groups in the subgenus Craugastor, patterns of speciation are evident among sister taxa. For example, sister species in the E. anomalus group exhibit a north- south pattern of differentiation in the Chocoan lowlands in E. anatipes and E. zygodactylus — the former to the north of the Rio San Juan and the latter to the south of the river (Fig. 71). Another pattern is exhibited by the lowland E. anomalus and its upland sister species, E. cheiroplethus (Lynch, 1990). In each case, the distributions of sister species are adjacent. Members of the E. bufoniformis group of the subgenus Craugastor also have a north-south vicariance pattern, but in this case E. bufoniformis is restricted to lowlands north of Rio San Juan northward into southeastern Costa Rica, whereas its sister species, E. necerus is restricted to eleva- tions of 600-1540 m on the Andean slopes in Ecuador (Fig. 72). The distributions of this pair of species are dichopatric. Other patterns involve partial sympatry of re- lated species. For example, of the three recognized members of the Eleutherodactylus diastema group in the Chocoan region, E. gularis is coastal, E. chalceus partially overlaps E. gularis inland and completely overlaps E. scolodiscus on the lower Andean slopes (Fig. 73). Likewise, in the E. cerasinus group, E. labiosus is widespread in the Chocoan lowlands south of the Rio San Juan; in the southeastern part of its range it is sympatric with its sister species, E. tenebrionis (Fig. 74), but other species in the group have allopatric distributions. Similarly, E. subsigillatus is widespread in the southern Chocoan lowlands, and it is sympatric with its putative sister species, E. degener in Provincia Esmeraldas, Ecuador; the apparent clos- est relatives of this pair of species are E. eremitus and E. phoxocephalus on the Andean slopes of Ecuador. The pattern of altitudinal replacement of related species is evident in other groups. For example, the E. loustes group consists of three species; E. loustes on the Andean slopes of Ecuador in adjacent Colombia is the sister species of E. jaimei on the Andean slopes of Departamento Cauca, Colombia, and E. hybotragus in the Chocoan low- lands (Lynch, 1992a). In western Ecuador, this pattern is repeated in the presumptive species pairs, E. walkeri in the lowlands and E. luteolateralis on the Andean slopes, and in E. latidiscus in the lowlands and E. laticlavius on the slopes (Figs. 75, 76). In each of these cases, the species in the lowlands have much greater areas of distribution than their putative highland sister species; for ex- ample the areas of distribution of the lowland E. walkeri and E. latidiscus are 1 9, 1 26 km- and 15,930 km-, respectively, in contrast to their respective putative upland sister species, E. luteolateralis (606 km-) and E. laticlavius (1900 km-). On the Pacific slopes of the Cordillera Occiden- tal of the Andes in Colombia and Ecuador, there seems to be a gradual latitudinal displacement of related species. However, the accumulating distri- butional data indicate the possibility of two major breaks. The first of these is in northern Departamento Valle, Colombia; this may be a reflection of greater antiquity of the northern part of the Cordillera Occidental (Galvis and Mojica, 1994), where there seems to be considerable endemism in Eleuthero- dactylus (e.g., E. bellona and half a dozen undescribed species). A second major break seems to exist between the area of La Munchique in Departamento Cauca, Colombia, and La Planada in southern Departamento Nariiio, Colombia. This disruption probably reflects the effects of the low and dry Rio Patia Valley, a barrier that is both climatological and a physiographic hiatus in the humid uplands of the Cordillera Occidental. This barrier creates disjunct distributions to the north ELEUTHERODACTYLUS IN WESTERN ECUADOR 179 Fig. 7 1 . Latitudinal replacement of sister species of Eleutherodactylus in the Chocoan lowlands. The disjunction in the range of E. anatipes is at the Rio Patia Valley. Fig. 72. Latitudinal and altitudinal replacement of sister species in the Chocoan region. Eleuthero-dactylus bufonifonnis is restricted to the lowlands to the north and E. necerus to middle elevations to the south. and south of the Rio Patia in several species (e.g., E. babax, cerastes, and duellmani). To the south in Ecuador, the upper drainage of the Rio Blanco seems to be a major region for speciation, which includes upland vicariants of lowland species, such as E. walkeri-E. luteolateralis and E. latidiscus- laticlavius. Latitudinal replacement of sister species at middle elevations is evident on the slopes of the Cordillera Occidental. This vicariance pattern is exemplified by E. ocellatus to the north and E. crenunguis to the south (Fig. 77) and by E. illotus to the south and E. thectoptemus to the north (Fig. 78). Trans-Andean patterns. — Members of some lineages have representatives on both sides of the Andes. For example, the E. cerasinus group (Lynch et al., 1994) consists of E. cerasinus in lower Central America, £. crenunguis, labiosus, ocellatus, orpacobates, and tenebrionis on the Pacific versant in Colombia and Ecuador (Figs. 74, 77) and E. rubicundus in cloud forests on the eastern slopes of the Andes in Ecuador. Similarly, the two species comprising the E. dolops group are distributed in cloud forests on opposite sides of the Andes; E. dolops occurs on the eastern slopes in northern Ecuador and southern Colombia, whereas E. babax has a lengthy distribution on the Pacific slopes in Colombia and Ecuador (Fig. 79), although the formerexample also demonstrates altitudinal shifts. The Eleutherodactylus sulcatus group has a more complex distribution in the Amazon lowlands and on various slopes of the Andes (Lynch, 1 986b). Eleutherodactylus sulcatus is widely distributed in 180 UNIV. KANSAS NAT. HI.ST. MUS. SPEC. PUBL. NO. 23 Fig. 73. Geographic replacement of members of the Eleutherodactylus diastema group in the Chocoan region. Fig. 74. Distributions of sister species in the Eleuthewdactyhts cenisinus group in the Chocoan region. the Amazon Ba.sin in Ecuador, northern Peru, and western Brazil; this species seems to be the sister taxon to the other members of the group, all of which are distributed in the Andes. Of these. E. comufiis, with a moderately broad latitudinal dis- tribution at elevations of 1150-1800 m on the Amazon slopes of the Andes in Ecuador and south- ern Colombia, seems to be the sister species of E. ingeri (eastern slopes of the Cordillera Oriental in Colombia) and E. cadenai (northern part of the Cordillera Occidental in Colombia. All of the other species now recognized occur on the Pacihc slopes of the Cordillera Occidental; E. cerastes has a broad latitudinal range in Colombia and Ecuador, whereas E. hekmotus, niizi, and seriuii have re- stricted distributions there. High Andes. — At elevations above 1500 m on the slopes of the Andes, the pattern of latitudinal replacement of related species is confounded by the complex topography. For example, of the four species comprising the E. siirdus group, E. diteUmcmi has a comparatively extensive latitudinal range of about 150 km at elevations of 1550-2710 m. Its range encompasses those of E. luimiotae and E. sobetes (each known from one, but different, lo- calities in Ecuador). The fourth member of the group has a latitudinal distribution of about 75 km at elevations generally higher than those of the other species in the group (Fig. 80). Likewise, in the E. devillei group, E. truehae replaces E. vertebndis to the south, but in this case the closest relative, E. devillei, is on the eastern slopes of the Andes (Fig. 81). In the cloud forests at higher elevations (>20()0 m), species tend to have small ranges and related species tend to replace one another geographically. ELEUTHERODACTYLUS IN WESTERN ECUADOR 181 Fig. 75. Altitudinal replacement of sister species of Eleittherodactylus in western Ecuador. This is exemplified by the nine species in the E. myersi group (Fig. 82). Eleiitherodactylus myersi has the most extensive distribution in the Cordil- lera Central in Colombia and the Nudo de Pasto in southern Colombia and adjacent Ecuador; three species (E. gladiator, repens, and trepidotiis) are restricted to the Amazonian slopes of the Andes in Ecuador and southern Colombia, and four species {E. floridus, hectus, ocreatiis, and pyrrhomerus) are restricted to the Pacific slopes in Ecuador and extreme southern Colombia. Two of these species seem to be lower-elevation vicariants of this prima- rily highland group; E. floridus is known from elevations of 700-2000 m, and E. hectus, from elevations of 1200-1750 m. One species in this group, E. leoni occurs on both Pacific and Amazo- nian slopes of the Andes. The species in the Eleiitherodactylus orestes group are distributed at high elevations in the Fig. 76. Altitudinal replacement of sister species of Eleiitherodactylus in the Chocoan region. Andes of southern Ecuador (Fig. 83). The range of E. vidua is encompassed within the distribution of E. orestes in the Cordillera Oriental, whereas E. simonholivari is restricted to the Cordillera Occi- dental. Frogs of the E. curt i pes group inhabit paramo and subparamo in the high Andes (2700-4400 m) of Ecuador and southern Colombia (Fig. 84). The ranges of E. buckleyi and E. curtipes overlap in southern Colombia and northern Ecuador; other- wise the two species seem to replace one another latitudinally. Eleiitherodactylus cryophilius is dis- junct to the south, and E. gentryi seems to be a peripheral isolate on the high Pacific slopes of the Cordillera Occidental in central Ecuador. The Co- lombian E. satagius and E. xestus have small, disjunct distributions on high peaks in the northern part of the Cordillera Occidental. 182 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 77. Latitudinal replacement of sister species of Eleutherodactylus on the Pacific slopes of the Andes in Ecuador and Colombia. Fig. 78. Latitudinal replacement of sister species of Eleutherodactylus in the Andes of Colombia and Ecuador. Historical Biogeography Although the Andes were upUfted moderately in the Cretaceous, some regions, such as the northern part of the Cordillera Occidental in Colombia and an island arc paralleling the Cordillera Occidental on the west, were uplifted in the early Tertiary, the major orogeny of the Andes north of the Huancabamba Depression commenced in the Pliocene (Galvis and Mojica, 1994; Sauer, 1971; Zeil, 1 979). The resulting high mountain chain was affected by dramatic climatic changes during the Pleistocene (Simpson, 1975; van der Hammen, 1974), and climatic fluctuations (mostly oscilla- tions in rainfall) had a marked effect on the adjacent lowlands (Haffer, 1974). (See foregoing section, Western Ecuador, for details.) Speciation models of Andean anurans based on vicariance events resulting from alternating cli- matic depression and elevation were proposed by Duellman (1983a) and Lynch (1986c). Climatic conditions seem to be extremely important in lim- iting distributions of anurans. Usually in the case of frogs, such as Eleutherodactylus, that are primarily inhabitants of humid rainforest or cloud forest, environmental moisture is viewed as an important environmental variable in limiting distributions. This is evident in the southern termination of the distributions of many species of Eleutherodactylus on the Pacific lowlands of Ecuador, where in a distance of a few score kilometers rainfall dimin- ishes from about 3000 mm to 600 mm or less annually. Moisture also seems to play an important role in the dry Rio Patia Valley in southern Colom- bia resulting in disjunctions in the distributions of some species (e.g., E. babax and E. cerastes), as do ELEUTHERODACTYLUS IN WESTERN ECUADOR 183 Fig. 79. Distributions of species in the Eleuthero- dactylus dolops group as an example of trans-Andean disjunction. The hiatus in the distribution off. babax is at the Rio Patia Valley. the dry valleys in the Huancabamba Depression in distributions of E. cajamarcensis and E. lymani. However, temperature also may play an important role in distributions. As concluded by Rome et al. ( 1 992), thermal adaptation of tolerance limits com- monly is evident in interspecific analyses and is likely to play an important role in setting temporal and distributional limits on amphibians. One as- pect of thermal tolerances that usually is not con- sidered in amphibian biogeography is their repro- ductive biology, but the results of investigations by Moore ( 1 939) and Zweifel ( 1 968) show that devel- opment of eggs and embryos is tied closely with temperature. Thus, changes in temperature, as well as humidity, most likely have played an important role in the isolation and subsequent differentiation of populations of Eleuthewdactylus as the Andes Fig. 80. Distributions of members of the f/^Mr/j^ra- dactylus surdus group as an example of latitudinal and altitudinal replacement on the upper Andean slopes. The hiatus in the distribution of £. duellmani includes the Rio Pati'a Valley. were uplifted to higher (and cooler) elevations and the temperatures and rainfall fluctuated during the Pleistocene. The complex topographic and climatic history since the Cretaceous and culminating in dramatic climatic changes during the Quaternary must have had a profound effect on speciation and distribution of organisms in western Ecuador and adjacent regions. By combining data on orogenic history of. and climatic vicissitudes in, western Ecuador, we propose a speciation model that incorporates the history of the region with possible vicariance events in the biota that are applicable to organisms that were inhabiting the region that is now western Ecuador prior to major uplift of the Andes. Our graphic model and attendant cladograms 184 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Fig. 81. Distiibutionsof species in the E/t-z/r/u^m- dactylus devillei group as an example of latitudinal replacement on the Pacific slopes with a disjunct spe- cies on the Amazonian slopes of the Andes. Fig. 82. Distributions of species in the Eleuthero- dactylus myersi group in the high Andes with species on eastern and western slopes of the Andes. Note that E. leoni occurs on both slopes. (Figs. 85, 86) are not based on any known cla- dograms or examples, but instead represent our expectations of what will be found (combining and extending our previous suggestions [Duellman, 1983a; Lynch, 1986c]). We envision the primary cause of speciation as being adaptation to climates (elevated regions vs. lowland regions) coupled with fragmentation of the once contiguous low- lands. As uplift continued, the impact was greatest on upland species because they would be more subject to vicariance events, such as erosional fragmentation of elevational zones and/or deposi- tion of discharge resulting in unsuitable habitat. The impacts of glacial stages would further frag- ment lower elevational zones, whereas interglacial stages would have the effect of generating addi- tional vertical slicing of distributions on Andean slopes. This model emphasizes climatic compres- sion during glacial times, which possibly could result in a mixture of species from the uplands and lowlands, as well as those from dry and humid environments. Possibly during interglacials. there was a resortment of taxa, but our ability to detect purported ecological compression will be limited by the resolution available in cladograms yet to be generated. This graphic model attempts to incorporate what we think we know of the patterns of distribution among Andean Eleuthewdactyhis — i.e., distribu- tions are most extensive in the topographically simple lowlands and near the upper limits of distri- butions (paramo and subparamo), whereas on the Andean slopes, distributions are narrow (limited altitudinal extent) and latitudinally elongated (but ELEUTHERODACTYLUS IN WESTERN ECUADOR 185 Fig. 83. Distributions of species in the Eleuthew- dactyliis orestes group in the Andes. not so elongated as in the lowlands). Our expecta- tion is that as additional species-level cladograms become available for Eleuthewdactylus in western Ecuador, this model will be .seen as a general one, at least for those groups limited by moisture but enjoying substantial vertical distributions. Comparisons with Other Regions An obvious comparison to be made is that between the eleutherodactyline faunas on the op- posite slopes of the Andes in Ecuador. Data were summarized from the comprehensive accounts of the Eleuthewdactylus of the Amazonian slopes of the Andes (Lynch and Duellman, 1 980), the south- ern Andes of Ecuador (Lynch, 1979a), and the Amazon Basin (Lynch, 1980a). Taxa that have been added subsequently to that region include E. ernesti (Flores, 1987), E. katoptroides (Flores, Fig. 84. Distributions of species in the Eleuthew- dactylus curtipes group in the Andes. 1988b), E. gauonotus and E. pecki (Duellman and Lynch, 1988), and E. libnirius (Flores and Vigle, 1994). Herein. E. kirkkmdi (Flores. 1985) is con- sidered to be a junior synonym of E. inusitatus Lynch and Duellman, 1980. The eastern versant in Ecuador differs strikingly from the western versant. The eastern lowlands are entirely covered by humid tropical rainforest re- ceiving up to 5000 mm of rainfall annually and displaying little or no seasonality. Furthermore, on the Andean slopes, cloud forest is nearly continu- ous throughout the length of the country. Unlike the western versant, the Cordillera Oriental is inter- rupted by the deep Pastaza Trench. Also, high elevations (>2000 m) exist in isolated ranges (Cor- dillera del Condor, Cordillera deCutucii, and Volcan Sumaco) to the east of the Cordillera Oriental. Thus, although there is greater latitudinal continu- ity of humid environments and a much greater 186 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Moderate Uplift Continued Uplift A C B ErosionA^olcanic Discharge D B Fig. 85. Speciation model for taxa with respect to uplift and erosion of the Cordillera Occidental of the Andes. Changes in the Andes are shown in the diagrams to the right; phylogenetic history is shown in the cladograms to the left, and distributions of lineages are indicated by letters on the diagrams corresponding to letters in cladograms. Lineage C is a vicariant on the eastern slopes after continued uplift results in uninhabitable elevations that fragment the range of Lineage B. ELEUTHERODACTYLUS IN WESTERN ECUADOR 1 87 Maximum Glacial A C D Maximum Interglacial A F C D E B Present Fig. 86. Speciation model continued into the Quaternary. In Maximum Glacial. Lineage A is restricted to humid area in the lowlands; depression distributions of Lineages B and D results in their dispersal into the areas previously occupied by only one of the lineages. In the Maximum Interglacial. Lineages C and E disperse (indicated by angled arrows) across the crest of the mountains. In the Present. Species H has differentiated on the eastern slopes, whereas Species C is distributed on eastern and western slopes. Contemporaneous species (a- h, lower case identifiers) are products of lineages (identified by upper case letters in cladograms and diagrams representing earlier times). 188 UNIV. KANSAS NAT. HIST. MUS. SPFiC. PUBL. NO. 23 expanse of lowland rainforesl, the topography is more complex than in western Ecuador. In contrast to 6 1 species oi'Eleutlu'rodactyliis in western Ecuador, 71 species are known from the eastern slopes of the Andes and the Amazon Basin. The lowlands are inhabited by numerous species. As many as 16 species are known from Santa Cecilia at an elevation of 340 m (Duellman. 1978). and 15 species have been reported from Jatun Sacha at an elevation of 450 m at the base of the Andes (Flores and Vigle. 1994). Many of these species are widespread latitudinally in the upper Amazon Basin from southern Colombia and Ecua- dor southward to northern Peru (e.g., E. croceoingiiinis, hmthcmites. nigrovittatus), central Peru (e.g.. £. martiae, sulcatiis, and variabilis), and even southern Peru (e.g.. E. altaniazoiiiciis, penivianus, and ventrimannoratiis)(Lync\\, 1980a, Duellman and Salas, 1991). Of the species restricted to the Andean slopes, 20 occur to the north of the Pastaza Trench, and six of these enter the trench; 15 species occur south of the Pastaza Trench, and five of those enter the trench (Lynch and Duellman, 1980). In addition to these, E. emesti is restricted to high elevations on Volcan Sumaco (Flores, 1987), and E. condor and E. pecki are known only from the Cordillera de Cutucii and Cordillera del Condor (Duellman and Lynch, 1988). The most striking contrast is in altitudinal distri- butions. Accumulated numbers of species at 1 00-m increments throughout the latitudinal expanse of the country reveals that the greatest numbers of species occur at different elevations on the eastern and western slopes of the Andes (Fig. 66). On the western slopes, more species are found at eleva- tions of 1 400-2 1 00 m than at other elevations, and as many as 28 species occur at elevations of 1 800- 1900 m. In contrast, on the eastern slopes, no more than 20 species are found within any 100-m incre- ment, and these are at elevations of 2400-2700 m. These altitudinal differences are apparent in the numbers of species in communities on the two slopes; on the eastern slopes, the largest commu- nity is 1 1 species at 1 740 m, and five other commu- nities contain six or seven species (Lynch and Duellman, 1980). In contrast, .seven communities on the western slopes of the Andes have six to thirteen species (r = 10.7). On the other hand, communities in lowland rainforests contain more species in the eastern lowlands ( 1 6 at Santa Cecilia; 1 5 at Jatun Sacha) than those on the western low- lands ( 1 ?> at Santo Domingo de los Colorados; 1 2 at Rio Palenque). Comparisons with Other Taxa Only two other groups of frogs are represented by many species on the Pacitic versant of Ecuador, but neither approaches the number of species of Eleutherodactylus. Seventeen species of centro- lenids are known from the Pacitic versant of Ecua- dor (Duellman and Burrowes, 1989). Four species (Centrolene prosoblepon, Cochrauella spinosa, Hyalinohatrachiumfleisclunanni, and H. valerioi) have extensive distributions in the lowlands and range at least from lower Central America through the humid tropical regime in western Ecuador; no centrolenids are known from the lowlands south of Quevedo, Provincia Los Rios, Ecuador. Centrolene buckleyi occurs in the high Andes of Colombia to extreme northern Peru (Duellman and Wild, 1993), whereas the other species are restricted to the slopes of the Andes in southern Colombia and Ecuador; of these, Centrolene gemmatunu heloderniatuni lynchi, peristictum, scirtetes, and Cochranella ocellifera are known only from eleva- tions of 800-2000 m between 1 °N Lat. and 1 °S Lat. Thus, like Eleutherodact}'lus, centrolenids show a diminution of Chocoan species southward on the Pacitic lowlands but a significant number of endemics in the cloud forests on the Pacific slopes of the Andes in Ecuador. In contrast, dendrobatid frogs of the genus Colostetlms exhibit a pattern of north-south geo- graphic replacement, both in the lowlands and in the highlands. Fifteen species of Colostetlms are known from the Pacific versant of Ecuador ( Coloma, 1995). Colostetlms talamancae is the only species that ranges from Central America to northwestern Ecuador. Colostetlms chocoensis and C lehinanni extend from southern Chocoan Colombia into north- western Ecuador, where these northern species broadly overlap C. awa and C. toachi in the low- lands and are replaced to the south by C breviquartus and C whymperi on the lowlands and lower slopes ELEUTHERODACTYLUS IN WESTERN ECUADOR 189 of the Andes. In contrast to patterns in Hleittliero- (hictylus, which seem to be restricted to more humid environments. C. nuicluililUi occurs in dry subtropical regime in the Cordillera de la Costa southward to about 3°S. Lat.. and C. infrcii^iitfcitus inhabits dry subtropical habitats from about 1°S Lat. to at least 5°S Lat. in northern Peru. Four species (C delatorreae, jcicohitspetcrsi. imiqui- puciimi, and vertehniUs) inhabit western high- lands, mostly above 2000 m. There is no prolifera- tion of species in the cloud forests in the Andes between 1 °N Lat. and 1 °S Lat. With the exception of Gastrothecu, hylid frogs are represented by few species on the Pacific ver- sant of Ecuador. Members of the Gastwtheca coniiita group exhibit altitudinal replacement on the Pacific versant (Duellman. 1983b), whereas most members of the G. phimhea group show latitudinal replacement in the high Andes, but the sister species, G. plwnbea and G. owphyla.x are on western and eastern slopes, respectively (Duellman andHillis, 1987). Bufonids are reasonably well represented on the Pacific versant of Ecuador. Again, the common pattern of diminution of Chocoan species (e.g., Bufo blomhergi, coccifer, and haenuititicus) exhts. However, in Atelopiis, four species (A. arthuri, coynei, longimstris, and mindoensis) are endemic to cloud forests on the Pacific versant in Ecuador. One species, A. elegans, in the Chocoan lowlands reaches the southern limits of its distribution in northwestern Ecuador; one species, A. halios, is known only from elevations of 350-650 m in the dry subtropical regime in southwestern Ecuador, and A. ignescens, prior to its great decline in the past decade, was widespread throughout the paramos of Ecuador and southern Colombia (Miyata, 1980b; Peters, 1973). As in Eleiithew- dactyliis, species of Atelopus on the western slopes of the Andes have relatives on the eastern slopes of the Andes. The three species of Andinopliryne exhibit latitudinal (A. atelopoides and A. olallai) and altitudinal {A. colomai) replacement on the western slopes of the Andes in southern Colombia and Ecuador (Hoogmoed, 1985). Among reptiles, only three genera of lizards are speciose in the region under consideration. At least 21 species of Anolis are known from the Pacific versant of Ecuador (Peters and Donoso-Barros, 1970); this region is the southern terminus of the genus in western South America. Five of these {A. auratus, hiponciliis, chloris, latifrons, and twpidogaster) range from Central America to Ec- uador; three others (A. enlaenuis, fniseri, and gnmuUceps) occur on the Pacific versant in Co- lombia and Ecuador. Twelve species are endemic to the humid lowlands and cloud forests of the Pacific versant of Ecuador, and one species {A. nigroUneatiis) is the most southern species on the Pacific lowlands, where it occurs in the dry sub- tropical regime. Gymnophthalmid lizards of the genus Procto- ponis are speciose in the cloud forests of Ecuador, where nine species are endemic to the Pacific slopes ( Kizirian. 1 994); six of these are known only between 1°N Lat. and 1°S Lat. Gymnophthalmids of the genus Pholidobolus are represented by five species in Andean Ecuador; these lizards display a general pattern of latitudinal replacement in the high Andes throughout the length of the country (Montanucci. 1973; Hillis. 1985). Thus, the same patterns exhibited by Eleuthew- dactylus are evident in other groups of frogs and lizards. Although some species are widespread throughout the Chocoan lowlands (and northward into at least lower Central America) and reach the southern limits of their distributions in the humid lowland forests of western Ecuador, latitudinal replacement of species is common in the Chocoan lowlands and in the Andes, especially at higher elevations. Also, taxa inhabiting the upper western slopes of the Andes commonly have related species on the Amazonian slopes of the Andes. However, no other genera of amphibians or reptiles exhibit an amount of speciation on the Pacific slopes of the Andes between 1°N Lat. and 1°S Lat. comparable to that in Elciitherodactyliis, in which 27 species are endemic. Woody plants (Gentw, 1982) and birds (Cracraft. 1 985 ) show a gradual diminution of Chtxxxm species in the lowlands luid on the Pacific slopes of the Andes. Tliere is no great burst of endemic birds on the Pacific slopes of Ecuador, but there aie mimy endemic species of plants both in the lowlands and on the slopes, which have not been thoroughly studied floristically. 190 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 FUTURE RESEARCH Assuming that we have encountered most of the species ofeleutherodactyUne frogs in western Ec- uador, we consider the most urgent priority for future research is investigation of the putative dechne in amphibian diversity and abundance dur- ing the past decade or so. Only a few biologists made the collections in the 1960s and 1970s that established western Ecuador as a region of species diversity; those collectors primarily were Kenneth Miyata and ourselves. It is incontrovertible that habitat destruction has occurred at an accelerated rate in recent years (Dodson and Gentry, 1 99 1 ; Parker and Carr, 1 992; Mejia et al., 1995). and it seems plausible that habitat destruction would bring about coincident disappearance of apparently endemic species (es- pecially the diversity "bubble" between 1°N and 1°S Lat.), but such disappearances are inferred based on tenuous evidence. Several investigators have worked in western Ecuador in the past decade (1985-1995), but those investigators may have far different field skills (search images and work hab- its) than did those who worked there earlier ( 1 966- 1980) when frog diversity seemed to be so great. Before we agree that Eleutherodactylus siirdiis is on the verge of extinction, as claimed by Flores ( 1 988a, 1 993 ), we advocate a current estimate of its distribution and abundance by persons having field experience in its habitats; neither of us has collected at sites where E. siirdus is known since 1978, when the species was abundant. Observ- ing that no new records are available when no one has searched for the species in proper habi- tats at known sites of occurrence provides the illusion of a species (and biota) in peril. Herein, we provide data on abundance of spe- cies of Eleutherodactylus at nine specific sites in western Ecuador; these data were accumulated in the period 1966-1978. By spending an equivalent number of collecting days (nights) at these sites, contemporary collectors should be able to gather sufficient data to determine the occurrence and abundance of the same (or other) taxa at these sites and thereby determine if changes in species com- position and abundance have occurred at either local or regional scales. This is a unique opportu- nity and challenge to carry out nine ecological and biogeographic experiments and to provide poten- tially meaningful data pertaining to postulated de- clines in abundance of amphibian populations. Although we have indicated that there is an area of endemicity over two degrees of latitude ( 1°N- 1°S) on the western slopes of the Andes, we ac- knowledge that this assertion may be an illusion of past collecting efforts that focused on accessible and interesting sites rather than attempt to sample the fauna throughout the entire region. Aside from Patricia Burrowes's intensive fieldwork at La Planada. Departamento Nariiio, Colombia (Lynch and Burrowes, 1990), little serious fieldwork has been carried out in western Nariiio, Colombia. Thus, the postulated limit at 1 °N Lat. is based more on failure to sample the herpetofauna immediately to the north than on the existence of latitudinal replacement of species. Several species (e.g., E. crucifer, ornatissimus, and tenebrionis) were collected recently in north- western Ecuador near the Colombian border; in each case the known distributions of these species were extended more than one degree of latitude to the north. It is unlikely that those distributions were (or are) disjunct; the gaps exist because little or no collecting has been done in the intervening area. Now that descriptions, a key, and color illustra- tions are available for most of the eleutherodactyline frogs of western Ecuador, we anticipate an explo- sion of discoveries of distributional extensions that will render our stated distributions to be severe underestimates. However, we predict that our gen- eral conclusions concerning communities in altitu- dinal zones and relatively small distributional areas will be reinforced by additional sampling. Some of the new species named herein {E. dissimulatus, gentryi, and truebae) have been known to us for nearly 30 years but are being described only now. Others (£. coloniai and E. degener) have been discov- ered within the past 5 years ; previously, we had no inkling of their existence. These species, as well as two undescribed taxa (see below ) and an undescribed species of Phyllonastes (L. Coloma and JDL, in prep.), attest the fact that we are still in the discovery phase of document- ing the eleuthero-dactyline fauna of westem Ecuador. ELEUTHERODACTYLIJS IN WESTERN ECUADOR 191 We are aware of yet two more undescribed species of Eleiithewdactylus in western Ecuador. One is a minute species from Provincia Esmeraldas; it is known from a single female found brooding an egg clutch in 1993 by Martha L. Crump. The other is represented by three specimens (an adult male and a juvenile female in an enclave of cloud forest west of Piiias, Provincia El Oro by WED in 1975, and another immature female from Piiias collected by H. Vargas in December 1 994). Neither sample is judged to be adequate for a description, and we defer describing these species until additional ma- terial is available.' The presence of these undescribed species is evidence of the incomplete- ness of the present endeavor and suggests that still other species may remain hidden in forest remnants in western Ecuador. We are concerned that small, isolated, enclaves of forest in southern Ecuador have not received sufficient attention. This is illus- trated by the fact that three species obtained there by non-herpetologists either represent the only sample of a species {E. ruidus) or the most southern (and disjunct) records for the species (£". chalceus and E. gularis). Although, in general, frogs of the genus Eleutherodactylus are sensitive to moisture, it should not be presumed that drier regions in southwestern Ecuador lack eleutherodactylines. Samples obtained on isolated hills along the coast by Ana Almendariz, John Carr, and Alwin Gentry (Parker and Carr, 1992) surprised us, given our limited previous sampling in Provincia Manabi and our expecta- tions that the region was too dry to support eleutherodactyline frogs. Little did we image three decades ago that the 20th Century would be waning before our endeav- ors on the Eleutherodactylus in western Ecuador would come to fruition. In the intervening years we have learned much about these frogs and have become increasingly aware that there is so much yet to be known. We leave this challenge to our successors, to whom we bid: /Que les vaya bieri! LITERATURE CITED AcosTA-SoLis, M. 1 984. Los Paramos Andinos del Ecua- dor Quito, Ecuador: Publ. Cien. MAS. Andersson, L. G. 1945. 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The Andes: A Geological Review. Berlin: Gebriider Borntraeger ZwEiFEL, R. G. 1968. Reproductive biology of anurans of the arid southwest, with emphasis on adaptation of embryos to temperature. Bull. Am. Mus. Nat. Hist. 140:3-64. APPENDIX I Specimens Examined The 5360 specimens identified to species from the Pacific versant of Ecuador are documented below, alphabetically by species. Localities are arranged alphabetically by provinces, which also are arranged alphabetically. Within a locality, speci- mens are arranged chronologically by catalogue number following museum abbreviations, which Eleutherodactylus achatimts (923 specimens) Ecuador: Prov. Azuay: 0.9 km W Luz Mari'a, 1770 m, QCAZ 7803; 7.7 km W Luz Maria, 1 300 m, QCAZ 7804-08; 1 1 .2 km W Luz Maria, 930 m, QCAZ 7809- 11; 12.9 km W Luz Maria, 740 m, QCAZ 7812-13; 13.5 km W Luz Maria, 740 m, QCAZ 7814-15. Prov. Bolivar: Balsapamba, 800 m, KU 130387-453. Prov. Canar: Chimbo, BM 98.3.1.31-32. Prov. Carchi: Maldonado. 1410 m, KU 177597-611; Rio Baboso, near Lita, MECN-LAC 30, 35, 37-40, 43-47, 49, 62, 65, 78, 80-81, 107, 120-22, 126. 144-46, 150-51; 2 km NE Rio Blanco, 930 m, USNM 204640-42. Prov. Chimborazo: Pagma Forest, ANSP 1 8244 (holotype of HylodespagtnaeYMoY'Q^C'ddo, 1600ft.AMNH 17546- 47, 17550, 17634-35. Prov. Cotopa.xi: Las Pampas. QCAZ 295, 543-48, 562; 3 km E Macuchi, 1500 m, USNM 204636-37; 18.6 km W Pilalo, 930 m, KU 141751; 20.3 km W Pilalo, 830 m, 141769. Prov. El are listed alphabetically. Localities that we believe to be incorrect are listed at the end of the records for a given species as erroneous locality. Only speci- mens from the Pacific versant of Ecuador are listed here; specimens from southern Ecuador listed by Lynch ( 1979) are not included. For detailed infor- mation on localities, see Appendix II. Oro: 7 km NE Buena Vista, 1 00 ft, USNM 204643-49; Gualtaco, USNM 204651-57; 7 km ESE Machala, 10 m, USNM 204650; 3 km E Pasaje, 30 m, AMNH 89738-42; 1 8 km W Piiias, 780 m. KU 1 65088-92; 4.4 km NW Pihas. 1100 m, KU 141770-71. Prov. E.smeraldas: Bulum, UMMZ 58907; Cachabe, BM 1947.2.15.69 (holotype); 1 km SW Cachabe. 20 m, USNM 204628-29; El Cristal. 1200 m. ECO 72-76; Pambelar, BM 1901.6.27.13; 2-4 km W Placer. 360- 390 m, USNM 204624-27; 10.5 km N Quininde. 130 m. KU 141 747; region of Ri'oCaoni, sector de Lagartera, UIMNH 53425, 53428-30; Rio San Miguel, ca. 1 km upstream from Rio Cayapas, MCZ 92931-32, 92940, 92947-49; Rfo Cupa, EPN 1422-26; Salidero, 350 ft, BM 1901.8.3.13; San Javier, AMNH 10708, UIMNH 55720; San Miguel, EPN 1105, 1 120. MCZ 92942-46; 38 km NW Santo Domingo de los Colorados, 1000 ft, USNM 204609-15, 204617-21; 31 km NNW Santo Domingo de los Colorados, 1000 ft, USNM 204616. 198 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 204622-23. Prov. Guayas: Bucay, 900 ft. AMNH 16250, 16987-88; 20 km W Guayas. 300 ft. USNM 204660; Naranjal. AMNH 17631-32; near junction of Rio Chimbo and Rfo del Oro, 2000 ft, CAS-SU 9434; Rfo Frio. UMMZ 123899. Prov. Imhabura: Lita, 520 m, KU 132605-09. MECN-LAC 226, 228, USNM 204658; Paramba. BM 98.3. 1 .30; Rfo Parambas, Ibarra- Lita road, EPN 1112. Prov. Los Rios: Echeandia. EPN 1462, 1468; Estacion Biologica Rfo Palenque, 220 m, AMNH 89730-32, KU 146049-55, 147557-60, 152572, 165082-87, 180638, MCZ 88427-30, 8990 1 - 04. 89918, 89920-26. 89928-38. 89940-47. 92114. 93456-62, 93464, 93468-72, 93474-75, 94800-06, 118035 (hatchlings); Quevedo, USNM 204630; 1 km NQuevedo, 300 ft, USNM 20463 1-35; 1 mi N Quevedo (Finca Playa Grande), UIMNH 93149-52; 4 km N Quevedo, 140 m, KU 130365-86, 135326-38; 3 km E Quevedo, MCZ 89913-15. Prov. Manabi: 2 km W Desvio, 250 m, USNM 204659; 38 km NW EI Carmen, 330 m, QCAZ 7816; 50 km WSW El Carmen, 400 m, MCZ 92014-18; 23 km N Manglaralto, 120 m, MCZ 92038^7; 25 km N Manglaralto, 60 km, MCZ 920 1 9- 20; Rfo Ayampe, 25 km N, 3.7 km E Montaiiita, 70 m, QCAZ 7817-23; Rfo Cuaque, E of Pedernales, 190 m, QCAZ 7824-25. Prov. Pichincha: 1 km N Buena Fe, MCZ 89905-12, 93320-419; 25.8 km W Chiriboga, 1720 m, KU 141748; Dos Rios, 1270 m, KU 135491- 93; 4 km E Dos Rfos, 1140 m, KU 165093-111. 202285-86; 3 km W Dos Rfos. 1050 m, KU 141765- 67; Finca La Esperanza, NW of Santo Domingo de los Colorados, EPN 1380, 1384, 1417; Guatea, 2900 ft, BM 1920.2.9.8-9; La Concordia, Bosque Protector La Perla, 190 m, KU 217808, QCAZ 7826-32; 5.3 km W La Concordia, 190 m, QCAZ 783-35; 5 km NW La Florida, 860 m, QCAZ 7836; La Palma, 920 m, KU 177618-21 ; 5 km E La Palma, 900 m, KU 165 112-19; 12.6 km E La Palma, MCZ 92847: 14.4 km E La Palma, 1 380 m, MCZ 9 1 986; 1 6 km E La Palma, 1 500 m, MCZ 89917; 22 km E La Palma, 1770 m, MCZ 89916; Llambo, USNM 204606-08; Miligali, USNM 204599- 600; Mindo, 4000 ft, UMMZ 55518; 18 km N Mindo, USNM 204584; 3.5 km NE Mindo, 1540 m, KU 165120-27, 165504; I km E Mindo, USNM 204573- 83; 4 km N Nanegal Chico, UMMZ 132918 (4); 1 .5 km SW Nanegal Chico, USNM 204594-95; 5 km NW Nanegal Chico, USNM 204588-90; 1 0 km NW Nanegal Chico, USNM 204591-93; Nanegalito. USNM 204585-87; Pachijal, USNM 204601; Pacto, USNM 204603; below Pacto, USNM 204602; Playa Rica, QCAZ 865-67; Puerto de Ila, USNM 204604; 8 km ESE Puerto Quito, 530 m, KU 165128; 46 km N Quevedo (Hda.CerroChivo), MCZ 9 1987-9201 3; Rfo Baba, 1 0 km S, 4 km E Santo Domingo de los Colorados, 400 m. KU 141752-64. 146056-59; Rfo Baba, 24 km S Santo Domingo de los Colorados, UIMNH 93148; Rfo Canoi, 4 km S Vicente Maldonado, 570 m, QCAZ m7837-38; Rfo Faisanes, 13.5 km E La Palma, 1380 m, KU 180639. MCZ 94470, 94917-18; Rfo Sapayo, 450 ft. BM 1902.7.29.25; RfoToachi, USNM 204596- 98; RfoToachi, 8 km E Alluriqufn, 800 m, KU 141768; San Miguel de los Colorados, USNM 204605; Santo Domingo de los Colorados, 500-660 m, FMNH 173070-71, KU 117778-79, 119465-71, 119472 (C&S), 141749-50, 109060, 177612-16, USNM 204549-53; I km N, 2 km E Santo Domingo de los Colorados, 620 m, KU 1 776 1 7; 2 km E Santo Domingo de los Colorados, 670 m, USNM 204566-68; 6 km E Santo Domingo de los Colorados, USNM 204569-73; 16 km E Santo Domingo de los Colorados, MCZ 93318-19; 35 km E Santo Domingo de los Colorados (Hda. Lelia), CAS-SU 10607-09, 10614; 8 km SE Santo Domingo de los Colorados, UMMZ 1 32916 (4); 0.5 km S Santo Domingo de los Colorados, 670 m, USNM 204554; 24 km S Santo Domingo de los Colorados (4 km E Rfo Baba bridge), UIMNH 93153- 54; 5 km W Santo Domingo de los Colorados, USNM 204555; 9 km W Santo Domingo de los Colorados (Hda. Espinosa), CAS 94853-54, CAS-SU 10481-92, 10495; 6 mi W Santo Domingo de los Colorados, CAS 85180; 18 km W Santo Domingo de los Colorados, USNM 204556-65; Tandapi, 1500 m, KU 1 1 1306-08 (C&S), 111278-305, 111309^4, 120256-60, 135463- 68, 135470, 135483-84, 135487-88, MCZ 75172-74; 2.1 km E Tandapi, 1500 m, MCZ 92839; Tinalandia, 700 m, KU 202278-84, MCZ 88420, 88426, 89883- 900; I km E Vicente Maldonado, 670 m, KU 218226, QCAZ 7839-42. Prov. Undetermined: Km 94, Guayaquil-Cuenca road, CAS 94787. Erroneous Lo- cality: Prov. Cotopaxi: region of Sigchos, USNM 204638-39. Prov. Pichincha: 10 km E Chiriboga, 7000 ft, USNM 204548. No data: EPN 1442, MECN 175, UIMNH 90297-98. Eleutherodactylus actites (250 specimens) Ecuador: Prov. Cotopaxi: La Esperanza, 1500 m, KU 131261-62; Pilalo, 2100-2800 m, KU 120111 (holotype), 120112-24. 131210-60, 141776-93, 141794-832, 177622-24, 202287-302, 202616, MCZ 103977; 2 km W Pilalo, 1820 m, UIMNH 90299-302, USNM 239616-25; 3 km W Pilalo, 1760 m, USNM 239557-615. Erroneous Locality: Prov. Cotopaxi: Latacunga, 2770 m, MCZ 104025. ELEUTHERODACTYLUS IN WESTERN ECUADOR 199 Eleutherodactylus anatipes (4 specimens) Ecuador: Prow Ccirchi: Maldonado, 1410 m, KU 177625, 177626 (holotype). Prow Esmeraldas: 2 km S junction of Rio Lita and Rio Mira, 520 m, USNM 233092-93. Eleutherodactylus anomalus (56 specimens) Ecuador: Prov. Esmeraldas: Cachabe, BM 1947. 16.8-10 (syntypes); Lagarto, Reserva Mayronga, 100 m. QCAZ 4729-30, 4319, 4437-39; Rio Cupa, CAS- SU 1 1455, USNM 204714; San Miguel-Telembi, EPN 1359; 38 km NW Santo Domingo de los Colorados, 1 000 ft, USNM 2047 1 9-20. Prow Esmeraldas: Estacion Biologica Rio Palenque, 220 m, KU 1 65 1 29-32, MCZ 93420-22, UMMZ 127891 (4); USNM 285345-47. Prow Pichincha: Rio Toachi, USNM 204718; Santo Domingo de los Colorados, 580 m, FMNH 174027, KU 177627-29; 1 km N, 2 km E Santo Domingo de los Colorados, 620 m, KU 177630-32; 6 km E Santo Domingo de los Colorados, USNM 204715-16; 35 km E Santo Domingo de los Colorados (Hda. Lelia), CAS- SU 10606; 8 km SE Santo Domingo de los Colorados, UMMZ 132815; 0.5 km S Santo Domingo de los Colorados, 670 m, USNM 204717; 9 km W Santo Domingo de los Colorados (Hda. La Espinosa), CAS- SU 10467-80; Tinalandia, 700 m, USNM 285960. Eleutherodactylus apiculatus (3 specimens) Ecuador: Prov. Pichincha: 4 km W Chiriboga, 2120 m, KU 142165-66; 6.2 km E Tandapi, Rio Corazon, 1750 m, MCZ 98078. Eleutherodactylus appendiculatus (51 specimens) Ecuador: Prov. Cotopaxi: Las Pampas, QCAZ 233, 587-89; Prov. Pichincha: 14 km W Chiriboga, 1960 m, KU 165142; 25.7 km from La Palma on Hwy 28 (old road to Quito), 1820 m, MCZ 91884-85, 94815-22; Las Maquinas, 3 km NW San Ignacio, AMNH 20143; Las Palmeras, QCAZ 853; Lloa, QCAZ 7968; Quebrada Zapadores, 5 km ESE Chiriboga, 20 1 0 m, KU 1 65 1 33- 41, 166230 (lot of young), 166264 (C&S), 177634-37, MCZ 98030, USNM 285925; Rio Blanco, USNM 204713; Rio Faisanes, QCAZ 599; Tandapi, 1460 m, QCAZ 129, 164, 600; 6.2 km E Tandapi, 1750 m, MCZ 9 1 886; Tandayapa, 2300 m, CAS-SU 1 1 460-6 1 , USNM 204712. Prow undetermined: "Ecuador," NHM 16507 (holotype). Erroneous Locality: Prov. Pastaza: Mera, Rio Pastaza, AMNH 52841^2. Eleutherodactylus hahax (2 specimens) Ecuador: Prov. Pichincha: 2 km E Tandapi, Quebrada La Plata. 1550 m, USNM 285970; 6.2 km E Tandapi, 1750 m, MCZ 92031. Eleutherodactylus cajamarcensis (8 specimens) Ecuador: Prov. Azuay: Luz Maria, 1800 m, KU 217853-56. QCAZ 7889-92. Eleutherodactylus calcarulatus (221 specimens) Ecuador: Prov. Carchi: 14 km SE Maldonado, 2500 m., KU 179273. Prow Cotopaxi: Reserva Otonga, 2100 m, QCAZ 4545. Prow Imbabura: 23 km W Apuela, 2 1 90 m, KU 1 79385-86; above Cuellaje, 2560 m, ECO 251, QCAZ 8050-59, 8063-71; La Delicia, 2700 m,KU 132754, 132758-66, 132768-69, 13277 1- 76, 177652-57, 179384. Prow Pichincha: 14 km W Chiriboga. 1960 m, KU 165160-61, 165869-70; 4 km NE Dos Rios, 1140 m, KU 165159; 14.4 km E La Palma, 1380 m. MCZ 91892; 25.7 km E La Palma, 1820 m, MCZ 94857-58, 94945; Mindo (Hda. San Vicente), 1465-1620 m, USNM 284352; 3.5 km NE Mindo, 1540 m, KU 165162-89, 166266 (C&S); Reserva Floristica-Ecologica Rio Guajalito, QCAZ 6434. 6558; Rio Faisanes, 13.5 km E La Palma, 1280 m. MCZ 92086. 92106, 94814, 95498, 93423, USNM 285873; 1 km SW San Ignacio, 1920 m, KU 177662, 179097-105; Tandapi, 1460 m, AMNH 114500-05, BM 1969.659-61, KU 111216-17, 111218 (holotype), 111219-40, 111242-75, 1 1 1276-79 (C&S), 117774- 75, 120255, 120270-76, 135455-57, 177659-61. MCZ 75 1 70-7 1, 75 1 75-77; 2.1 km E Tandapi, 1500 m, MCZ 94718, 94863, 95496-97, 95609-11, 98032, USNM 285972-73; 6.2 km E Tandapi, 1750 m, MCZ 9 1893- 94, 98076, USNM 285940. 2 km E Tandayapa, USNM 233094; 1 km SW Tandayapa, 1640 m, MCZ 90336, USNM 285949; 4.5 km W Tandayapa, QCAZ 4789. Eleutherodactylus caprifer (52 specimens) Ecuador: Prov. Carchi: Rio Baboso. near Lita, MECN-LAC 50-5 1 , 56, 87-88, 1 37. Prow E.smeraldas: Cachabe, BM 98.3.1 .29. Prow Pichincha: 5 km NW La Florida, QCAZ 577-79, 581; La Palma, 920 m, KU 131589 (holotype), 131590-602, 177663-82; 1.1 km E La Palma. MCZ 95630-35. Erroneous locality: Prov. Cotopaxi: Las Pampas, QCAZ 500. 200 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Eleuthcrodactylus ccUitor (68 specimens) Ecuador: Frov. Carclil: 14 km SE Maldonado, 2500 m, AMNH 114527-31, KU 177684 (C&S). 177689 (C&S), 177690-725. Prov. Imbahura: 22 km E Apuela. 2860 m. QCAZ 2842-43; La Delicia, 2700 m. KU 131573 (holotype), 131574-88. 1 77726-29. Pmi'. Pichincha: Lloa, QCAZ 7972; 9.5 km NW Nono. 2530 m. KU 165200-01. EleutherodactylHS cerastes ( 16 specimens) Ecuador: Prow Imhahuni: Lila, USNM 195786- 88. 195795. Prov. Picliinclta: Pachijal. USNM 195789- 93, MCZ 89462; road from Pacto to Guayllabamba, USNM 195797; Palma Real, USNM 195785 (holo- type), 195794; Rio Pitsara. USNM 195796 (3). EleutherodactylHS chalceus ( 140 specimens) Ecuador: Prov. Carchi: Maldonado. 1410 m. KU 1 77638. Prov EsmeraUlas: Alto Tambo. 830 m, QCAZ 1259. 1291-93; Cachabe. BM 1947.2.15.38 (syntype ot" Syrrhophus areolatiis): 10.5 km N Quininde. KU 141774-75; Rio San Miguel, ca 1 km upstream from Rio Cayapas, MCZ 92972-76, 93016; San Miguel. MCZ 92977-80. QCAZ 166-67; 38 km NW Santo Domingo de los Colorados. 1000 ft. USNM 204730- 32. Prov. Imhabura: Lita, 570 m, QCAZ 1309. Prov. Guayas: Chimbo, BM 1947.2.15.39 (syntype of Syrrhophus areolatiis). Prov. Esmeraldas: Estacion Biologica Rfo Palenque, 220 m, MCZ 88415-16, 89948-53. 91890. 92836. Prov. Pichincha: Centinela, 1 4. 1 km ESE Patricia Pilar, 550-600 m, USNM 285563- 80; 4 km E Dos Rios. 1 120 m, KU 203322; 14.4 km E La Palma, 1 380 m, MCZ 9 1 887-89; 3.5 km NE Mindo, 1540 m, KU 165143-49; 5 km NW Nanegal Chico, USNM 204721; Puerto Quito, MHNG 18879; Km 12 Quito-Puerto Quito road, 750 m, MECN ( 1 ); Rio Baba, 10 km S, 4 km E Santo Domingo de los Colorados, 400 m. KU 141773; Rio Baba, 19 km S, 5 km E Santo Domingo de los Colorados, UIMNH 77409; Rio Faisanes, 13.5 km E La Palma, 1380 m, MCZ 94471- 73, 948 1 3, 95628, USNM 285874-75; Santo Domingo de los Colorados, 600 m. KU 117487-91, 118129 (C&S), 1 19474-500, 1 19501 (C&S), 120252. 177639- 48. MCZ 884 1 3- 1 4. WC AB 44396-400; 2 km E, 1 km S Santo Domingo de los Colorados, 620 m, KU 1 77649- 50; 6 km E Santo Domingo de los Colorados. USNM 204724-28; 1 8 km E Santo Domingo de los Colorados. USNM 204729; 5 km W Santo Domingo de los Colorados. USNM 204722-23; Tinalandia. MCZ 88412,91891. EleutherodactylHS colonial (3 specimens) Ecuador: Prov. Esmeraldas, Alto Tambo, 830 m. QCAZ 1289 (holotype). 1296; ElCristal. 1 200 m. ECO 217. Eleuthcrodactylus crenimguis (99 specimens) Ecuador: Prov. Los Rios: Estacion Biologica Rfo Palenque. QCAZ 172-73. Prov. Pichincha: 5 km NW La Florida. 860 m, QCAZ 7843-45; 1.1 km E La Palma. MCZ 95636-38, 12.6 km E La Palma. MCZ 90005. 94726; 16 km E La Palma, 1500 m, MCZ 90007; Mindo (Hda. San Vicente), 1465-1620 m, USNM 284355; 3.5 km NE Mindo, 1540 m, KU 165223-34; Nanegalito, USNM 239847; Pachijal, USNM 239851; Palma Real. USNM 239855-57; Quebrada La Plata, 3 km SE Tandapi, 1550 m. MCZ 89973. 98133-34, USNM 285974; Rfo Faisanes, 13.5 km E La Palma. 1380 m. MCZ 93424-27. 94465. 94849-56. 94902-05. 95499-500. 97516-25. USNM 285877-83; Rfo Mulaule. trib. Rfo Blanco, USNM 239850; Rfo Pitsara, USNM 239853-54; 1 km SW Tandayapa, MCZ 97877, USNM 285950; Tandapi, 1460 m, KU 111213-15, 120125. 120126 (holotype). 120127-31. 131605. 177730-32. MCZ 105473; 2.1 km E Tandapi, 1 550 m, MCZ 92840 95493-95, 94823; Tinalandia, MCZ 92083-84, 92107-08, 94713-16. USNM 217413. EleutherodactylHS crucifer (48 specimens) Ecuador: Prov. Bolivar: Porvenir, 1760 m. BM 1947.2.16.91 (holotype). Prov. Carchi: Rfo Baboso. near Lita, MECN-LAC 42. Prov. Cotopaxi: Las Pam- pas, QCAZ 591, 604-05. Prov. Imbahura: "above" Lita, 1200 m. ECO 087, 123. Prov. Pichincha: 18 km E La Palma. 1600 m. MCZ 90013; 3.5 km NE Mindo. 1540 m. KU 165235; ReservaFlorfstica-EcologicaRfo Guajalito. near Las Palmeras. 1800 m. QCAZ 2627; Rfo Faisanes. 13.5 km E La Palma. 1380 m. MCZ 94474-75; RfoToachi. USNM 21 1574; Tandapi. 1460 m. KU 111206-12. 120132-50, 131667-71, 177733, QCAZ 603; 1.6 km W Tandapi. 1400 m. KU 177734. Erroneous locality: Prov. Pastaza: 1 km W Puyo. MCZ 90012. Eleuthcrodactylus degener (3 specimens) Ecuador: Prov. Esmeraldas: Alto Tambo. 830 m. QCAZ 1297. Prov. Esmeraldas: El Cristal. 1200 m. ECO 77, 122. ELEUTHERODACTYLUS IN WESTERN ECUADOR 201 Ek'utherodcutylus (lisslninkitus ( 13 specimens) Ecuador: Prow Ficliiucha: Quebrada Zapadores, 5 km ESE Chiriboga. 2020 m. KU 165923-25, 179087. 179088 (holotype). 179089-95; 1 km SW San Ignacio. 1920 m. KU 179096. NW Nono. MCZ 90320-24; Rfo Blanco, near mouth of Rio Yambi, 700 m. USNM 239683; Rio Lelia, tributary ofRioToachi. USNM 239676-79; Rio Peripa headwa- ters, San Miguel dc Congoma. USNM 239684; Rio Toachi (Km 100-110), USNM 239685; Tandayapa, USNM 239681-82. Eleuthewdactylus duellmani (139 specimens) Ecuador: Pwv. Carclii: 5 km W La Gruel, 2340 m, KU 202403-04; 26.9-27.3 km E Maldonado. 2420 m. KU 2 1 7997-98, QCAZ 3084-89; 1 4 km SE Maldonado. 2500 m, KU 179251-67. 179268 (C&S), 179269-72. 179274-95. Prov. Imbahura: La Delicia. 2700 m, KU 179296-3 15. Prav. Pichincha: 14.8kmESEChiriboga. 2410 m, KU 179338-46; 14 km WChiriboga, 1960 m, KU 165910-12; Quebrada Zapadores, 5 km ESE Chiriboga, 2010 m, KU 165913-21, 179316-24, 179325 (holotype), 179326-37; 2.1 km E Tandapi, 1 550 m, MCZ 947 1 9-20, 956 1 5-27, 9748 1 ; 9 km SE Tandayapa. 2150 m, KU 165905-09. 202516. Eleuthewdactylus ereniitus (16 specimens) Ecuador: Prov. Cotopa.xi: Las Pampas, KU 22 1 685- 86; Palo Quemado, MHNG 18739. Prov. Pichiiiclui: 2 km W Campamento Silante, 2 1 00 m, KU 1 40878; 8 km W Chiriboga, USNM 211209; 18 km NE La Palma, MCZ 90335, 25.7 km NE La Palma. 1820 m, MCZ 92103 (holotype). 92104; 3.5 km NE Mindo. 1540 m, KU 1 65884; Quebrada Zapadores. 5 km ESE Chiriboga, 2010 m, KU 179085-86, USNM 285924; 6.2 km E Tandapi. 1750 m, MCZ 92105, 94723; Tandayapa, USNM 2 1 1 208; 8.6 km SE Tandayapa, 2000 m, MCZ 98189. Eleutherodactylus eugeniae (15 specimens) Ecuador: Prov. Pichincha: Quebrada Zapadores, 5 km ESE Chiriboga, 2010 m. KU 165899 (holotype), 165900-04, 179382-83, MCZ 98031; Reserva Floristica-Ecologica Rio Guajalito, near Las Palnieras, 1 800 m, QCAZ 2629; 6.3 km E Tandapi. 1 700- 1 750 m. MCZ 94724, 98075, 98077, 98079-80. Eleutherodactylus fioridus (27 specimens) Ecuador: Prov. Cotopaxi: region of Sigchos, USNM 239672 (holotype) 239673. Prov. Imbahura: Lita, Rfo Mira, USNM 239674-75. Prov Pichincha: Bosque Protector Mindo-Nambillo. 1700 m. QCAZ 7296; re- gion of Gualea. USNM 239690; Milpe. 900 m. USNM 239691-92; immediate environs of Mindo. USNM 239686-89; road to Mindo. USNM 239680; 12 km Eleutherodactylus gentryi (69 specimens) Ecuador: Prov. Cotopaxi: 5 km (by road) E Pilalo, 3200 m, KU 131540 (holotype). 131541-65; 8 km (airline) E Pilalo, 2850 m, USNM 239741-62; 9 km (airline) E Pilalo, 3125-3200 m, USNM 239763-67, 239772-73; 10 km (airline) E Pilalo, 3200 m, USNM 239739^0; 24.6 km E Pilalo, 3190 m, QCAZ 7846- 5 1 ; 27.6 km E Pilalo. 3380 m. KU 2 1 8 1 09- 1 1 . QCAZ 7852-54. Eleutherodactylus gularis ( 1 1 specimens) Ecuador: Prov. Chimborazo: Huigra, 1235 m,ANSP 18113 (holotype of Hyloxalus hiiigrae). Prov. Esmeraldas: 1 kmSWCachabe,20m,BM 1947.2.15.82 (holotype). USNM 211977; Durango. QCAZ 4316- 17; Rfo San Miguel. MCZ 92909-10, 92919, 92926- 27; San Miguel de Cayapas, QCAZ 174. Eleutherodactylus hainiotae (5 specimens) Ecuador: Prov. Pichincha: 12 km W Nono, USNM 239843; 13.1 km NW Nono, 2140 m, MCZ 97486, 98024-25, 98027 (holotype). Eleutherodactylus hectus (3 specimens) Ecuador: Prov. Esmeraldas: El Cristal, 1200 m, ECO 88, 119-20. Eleutherodactylus helonotus (2 specimens) Ecuador: Prov. Pichincha: Mindo, USNM 195784; RfoPitsara. BM 1970.178. Eleutherodactylus illotus (24 specimens) Ecuador: Prov. Carchi: 5.9 km E Maldonado. 1780 m, USNM 286283. Prov Imbahura: above Cuellaje, 2560 m, ECO 1 35-36, 1 49, 1 7 1 . 1 75. Prov. Pichincha: 22 km NE Las Palmas, 1770 m. MCZ 90339-^0; Llambo(on Gualea road), USNM 239732-33; Miligali. USNM 239722; Mindo (road to). USNM 239727; Mindo (region). USNM 239730-3 1 ; 3.5 km NE Mindo, 1540 m, KU 165881 (holotype), 165882-83; Nanega- 202 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 lito. USNM 239724; Pachijal. USNM 239723; below Pacto, USNM 239725-26; Rfo Faisane.s, 1 5 km NE La Palma, 1 380 m; San Tadeo, near Mindo. USNM 239729. Questionable Locality: Prow Ficliinclui: Puerto de Ila, USNM 239728. Eleutherodactylus labiosus (109 specimens) Ecuador: Prov. Carchi: Rfo Baboso, near Lita, MECN-LAC 138. Prov. Esmeraldas: AltoTainbo, 830 m, QCAZ 1265; Hda. Equinox, 33 km NW Santo Domingo de los Colorados. USNM 2 1 2008 Rfo Cupa, USNM 239793. Prov. Imbabura: Lita, 570 m, QCAZ 1310. Prov. LosRios: Estacion BiologicaRfoPalenque, KU 165895-96, MCZ 90053-54, 91954-56, 92113, 94461, 94866, USNM 285348-49. Prov. Pichincha: Centinela, 14.1 km ESE Patricia Pilar, MCZ 97575- 9 1 , USNM 28559 1-6 1 3; 5 km NW La Florida, MECN- LAC 210, QCAZ 569-70; La Palma, KU 131612 (holotype); 5 km E La Palma, KU 165894; 16 km NE La Palma, MCZ 90006; Nanegal Grande, USNM 212005; Rfo Pitsara, USNM 239852; Santo Domingo de los Colorados, USNM 212006; 6 km E Santo Domingo de los Colorados, USNM 212007; 1 1 km E Santo Domingo de los Colorados, USNM 285981; Tinalandia, 800 m, MCZ 88392-93, 88891-93, 89974- 92, 89993-90004; 90049-52; 91953. Eleutherodactylus laticlavius (26 specimens) Ecuador: Prov. Carchi: 22 km E Maldonado, 2560 m, QCAZ 3740, 4186. Prov Esmeraldas: El Cristal, 1200 m. ECO 80-82, 84-86, 89-90, 92, 104-07, 124- 25, 128-31, 224-225, 240, QCAZ 6339. Prov. Pichincha: Rfo Faisanes, 14.3 NE La Palma, 1380 m, USNM 285876. Eleutherodactylus latidiscus (80 specimens) Ecuador: Prov. Carchi: Rfo Baboso, near Lita, MECN-LAC 33, 74-15. Prov Esmeraldas: AltoTambo. 830 m, QCAZ 1254, 1258, 1260-61, 1263-64, 1266, 1295, 1298; Bilsa, EPN-AA 2510, 2513, 2517; Cachabe, BM 1947.2.15.66-67 (syntypes); Lagarto, Reserva Mayronga, 100 m, QCAZ 4728; San Miguel, MCZ 92920. Prov. Los Rios: Estacion Biologica, Rfo Palenque, 200 m, KU 1 65468-70, MCZ 8839 1 , 900 1 4- 1 6, 92 1 1 1-1 2, 94455, 94457-58. Prov Manabi: 38 km NW El Carmen, 330 m, QCAZ 7855. Prov. Pichincha: Centinela, 14. 1 km SE Patricia Pilar, 550-600 m, MCZ 97555, 97557, 97574, USNM 285539-40, 28558 1-90; La Florida, MECN-LAC 205-06, 208; 5 km NW La Florida, KU 218016-17, QCAZ 5856, 5857 (eggs), 5858; Santo Domingo de los Colorados, KU 109059, 120222-26, 120232-33, 141957-61, 179397-98, 179400; Santo Domingo de los Colorados (Finca La Esperanza), USNM 239849; Tinalandia, MCZ 88422, 9001 7, 90330-33, 9 1 22 1 , 947 1 7. Eleutherodactylus leoni (43 specimens) Ecuador: Prov. Carchi: 2 km E La Gruel, 2590 m, KU 202410; 14 km SE Maldonado, 2500 m, KU 177320-40; 26.9 km W Maldonado, 2420 m, KU 218232-33; 51.3 km WTulcan, 3150 m, KU 218227- 31. Prov. Imbabura: above Cuellaje, 2560, QCAZ 8046-49; La Delicia, 2710 m, KU 132779; Nudo de Mojanda, N slope, 3400 m, KU 130870 (holotype), 130871-72. Prov Pichincha: 14 km W Chiriboga, 1960 m, KU 165897-98; Lloa, 2800 m, QCAZ 7969- 7 1 . Erroneous Locality: Pichincha: Rfo Blanco, near mouth of Rio Yambi, 700 m, USNM 212009. Eleutherodactylus longirostris (268 specimens) Ecuador: Prov. Esmeraldas: Bulum, 60 ft, AMNH 10698-700, MCZ 7600, UMMZ 83826 (5); Cachabe, BM 1947.2.15.56-60 (syntypes), USNM 233120; 1 km SW Cachabe. 20 m, USNM 23 1 10-14; 1 .5 km SW Cachabe. USNM 233115-17; Hda Equinox, 38 km NW Santo Domingo de los Colorados, USNM 233 1 08- 09; La Boca, MHNG 18503; Pambelar, BM 1901. 6.27. 14-1 5; ParroquiaCarondelet, Rfo Bogota, UIMNH 53558-59; Rfo Bogota, USNM 233118-19; Rfo Durango, 350 ft, AMNH 10706-07, MCZ 3891-92, UMMZ 51272; Rfo San Miguel, ca. 1 km upstream from Rfo Cayapas, MCZ 92929, 92934, 92953-71; Salidero, AMNH 10702, 10704; UMMZ 51265; San Javier, AMNH 10709-10. MCZ 7599; San Miguel, EPN 1 102, 1 104, 1 108-09, 1 1 1 5-1 6, 1 1 1 8, MCZ 85756, 85789, 92952-60; QCAZ 169-70; Urbina near San Lorenzo, EPN 1358. Prov. Guayas: Naranjito, Rfo Chimbo, USNM 233 107. Prov Prov. LosRios: Estacion Biologica Rfo Palenque, AMNH 89728-29, KU 147561-66, 152573, 165474-82. 165483-84 (C&S), 165485-95, 166277-78 (C&S). 180640, MCZ 8843 1 - 33,89919,89927,90055-107,91217-19,93476-511, 94838-41, 94888, 118034 (hatchlings), QCAZ 168, USNM 285350-58. Manabi: 38 km NW El Carmen, 330 m, KU 218018, QCAZ 7859. Prov Pichincha: Finca La Esperanza, NW of Santo Domingo de los Colorados, EPN 1 377. 1 382; Hda. Cerro Chico, 45 km N Quevedo, 1 70 m, USNM 285727-29; La Concordia, Bosque Protector La Perla, 190 m, QCAZ 7860; Pacto ELEUTHERODACTYLUS IN WESTERN ECUADOR 203 (road to Guayllabamba). USNM 233 1 24; Puerto tie Ila, USNM 233123; 8 km ESE Puerto Quito. 530 m, KU 1 65496-503 ; Rfo Baba, 5- 1 0 km SSW Santo Domingo de los Colorados, 500, AMNH K9727; Rio Baba, 24 km S Santo Domingo de los Colorados, UIMNH 93158; Rio Canoi, 4 km S Vicente Maldonado, 570 m, QCAZ 7861; lower Rio Toachi, USNM 233125; Santo Domingo de los Colorados, USNM 233 1 2 1 ; 1 km N, 2 km E Santo Domingo de los Colorados, 6290 m, KU 1 77804-06; 35 km E Santo Domingo de los Colorados, (Hda. Leiia), CAS-SU 10611; 2 km E, 1 km S Santo Domingo de los Colorados, 600 m, KU 1 77803; 1 9 km S, 5 km E Santo Domingo de los Colorados, UIMNH 936 1 0- 1 1 ; 5 km SE Santo Domingo de los Colorados, USNM 285794; 18 km W Santo Domingo de los Colorados (Ramsey Farm, Km 19 on road to Chone), USNM 233122. Prov. undetermined: Northwestern Ecuador, CAS 66296. Erroneous locality: Prov. Pastaza: Sarayacu, BM 80.12.5.229, 80.12.5.249. Eleutherodactylus loustes (20 specimens) Ecuador: Prov. Carchi: Maldonado, 1410 m, KU 179231-33. 179234 (holotype), 179235-50. Eleutherodactylus luteolateralis (81 specimens) Ecuador: Prov. Pichincha: 14 km W Chiriboga, 1960 m, KU 165160; 4 km NE Dos Rios, 1140 m, 165506 (&S), 165507-21, 166279 (C&S); 10.4 km E La Palma,MCZ 93430-31, 10.6 km ELa Palma, 1280 m, MCZ 9 1 957-58; 1 2.6 km E La Palma, MCZ 92848; 1 3 km E La Palma, MCZ 903 1 0; 1 4.4 km E La Palma, 1380 m, MCZ 91959-62, 92053; 16 km E La Palma, 1500 m, MCZ 90309; 3.5 km NE Mindo, 1540 m, KU 165522-38, 166280 (C&S); Rio Faisanes, 13.5 km E La Palma, 1 380 m, MCZ 93428-29, 94476-78. 94843. USNM 285884; Tandapi, 1460 m, AMNH 114490-95, KU 111378-84, 120151-54, 131672-73, 131674 (ho- lotype), 121675-77, 135448-51. Eleutherodactylus lymani (14 specimens) Ecuador: Prov. Azuay: 4 km SW Catavina, 1600 m, USNM212019;55.4kmEPasaje, 1000m, KU 152009, Rio Minas, 20 km W Santa Isabel, 1250 m, USNM 212020. Prov. El Oro: Cordillera de Chilla, Llanos de Guavos, AMNH 13738; El Chiral, AMNH 13961, 1 3964; Guainche, AMNH 1 6256; Pinas, AMNH 16257; Portovelo, 610 m, AMNH 16334, 16339, 16341-44. Prov. Loja: Loja, BM 1947.2.15.99 (holotype of Eleutherodactylus carrioni). Eleutherodactylus muricatus (16 specimens) EcuADOR:Prov. Los Rios: Estacion Biologica Rio Palenque, 220 m, MCZ 92110, 94456, 94460. Prov. Pichincha: RioFaisanes, 14.4 km ELa Palma, 1380m, MCZ 92091, 92095, 92100-01, 94469 (holotype), 94848, 97528-3 1 , USNM 2111 72-74; Santo Domingo de los Colorados. KU 179084. Eleutherodactylus necerus (58 specimens) Ecuador: Prov. Carchi: Maldonado, 1410 m, KU 179076-79; Prov. Cotopaxi: Las Pampas, 1500 m, MCZ 1 044 1 8, QCAZ 1 9, 227, 563-64. Prov. Pichincha: 13 km E Alluriquin, MCZ 88394; 4 km NE Dos Rios, 1140 m, KU 165541 (C&S). 165542-43, 166067; 14.4 km E La Palma, 1380 m, MCZ 91895-903; Mindo, USNM 195798 (holotype); 3.5 km NE Mindo, 1540 m, KU 1 65544-45; road from Pacto to Rio Guayllabamba, USNM 195851 ; Rio Blanco near mouth of Rio Yambi, USNM 195852; Rio Faisanes, 13.5 km E La Palma. 1380 m, MCZ 92882, 94844-45, USNM 285885-91; Rio Lelia, tributary of Rio Toachi, USNM 195799; Santo Domingo de los Colorados, MCZ 113574-75; Tandapi, 1460 m, KU 179080-82, MCZ 105457-59. 1 05467-68; 2. 1 km E Tandapi, 1 500 m, MCZ 956 1 3- 14, 98033; 2.1 km E Tandapi, Quebrada La Plata, 1550 m, USNM 28594 1 , 285975-77; 1 .6 km W Tandapi, KU 179083. Erroneous locality: Pastaza: Pastaza River, Canelos to Marafion, MCZ 19628. Eleutherodactylus nyctophylax (153 specimens) Ecuador: Prov. Cotopaxi: Galapagos, 1720 m, KU 221687; Palo Quemado, MHNG 18739; Reserva Otonga, 2100 m, QCAZ 4546; vicinity of San Fran- cisco de las Pampas, 1800 m. QCAZ 590. 592-94, 596-98. 2509. Prov. Pichincha: 4 km NE Dos Rios. 1140 m, KU 165546 (C&S). 165547; 10.4-10.6 km E La Palma, 1280-1300 m, MCZ 92087-89. 94725; 16 km E La Palma. MCZ 90108; 3.5 km NE Mindo. 1 540 m. KU 165548-49; Tandapi. 1460 m, AMNH 1 14506- 11. BM 1969.654-58; KU 110896-908. 110909 (holo- type), 110910-12, 110913 (C&S), 110914-22, 110923 (C&S), 110924-74, 111376-77. 117582-83, 130879- 86, 135428-46, 135459, 138789, 177812-13, MCZ 75158-63; 6.3 km E Tandapi, Rio Corazon, 1700 m, USNM 285938-39. Eleutherodactylus ocellatus (5 specimens) Ecuador: Prov. Carchi: 8 km W Maldonado, 1255 m, QCAZ 3741; 3 km up Rfo Blanco from Chical, 204 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 1450-1500 111. USNM 286293. Pnn: tiiihahura: above Cuellaje. 2560 m, ECO 168-170. Eleuthewdactyhts onnitissimiis (26 specimens) Ecuador: Prov. Canhi: Ri'o Baboso, near Lita, MECN-LAC 108. Prov. Cotopaxi: Galapagos, MHNG 18707; Palo Quemado. KU 221684; 20.3 km W Pilalo. 830 m. KU 141969 (C&S) 141970-71. Prov. Esmer- aldas: El Cristal. 1200 m, ECO 083, 095. Prov. Pichincha: Centinela, 14.1 km SE Patricia Pilar, 550- 600 m, MCZ 97593; Rio Baba, 4 km E, 10 km S Santo Domingo de los Colorados, 400 m, KU 141967-68; Rio Blanco, near mouth of Rio Yambf. 700 m. USNM 233205; Rio Pitsara, USNM 233206; Santo Domingo de los Colorados. KU 119744-48, 119749 (C&S), MCZ 1 06946; 6 km E Santo Domingo de los Colorados (Dyatt Farm), USNM 233201-02; 4 km W Santo Domingo de los Colorados (Finca La Esperanza), USNM 233203; 18 km W Santo Domingo de los Colorados, 650 m. USNM 233200; below Sigchos, USNM 233204; Tandapi, 1460 m, MCZ 105082. Eleutherodactylus parvillus (90 specimens) Ecuador; Prov. Aziuiy: Manta Real, Parroquia Molleturo, 650 m. EPN-AA 2550. Prov. Cotopa.xi: Reserva Otonga, 2000 m, QCAZ 6565. Prov. Esmer- aldas: Bilsa. 225 m. EPN-AA 2488; El Cristal, 1 200 m, ECO 219; Hda. Equinox, 38 km NW Santo Domingo de los Colorados, 300 m, USNM 239782. Prov. Los Rios: Estacion Biologica Rfo Palenque, MCZ 90115. Prov. Pichincha: Centinela, 14.1 km ESE Patricia Pilar, 550-600 m, USNM 285615-21, 285545; Mindo. 1 500 m. QCAZ 34 1 5- 1 7; Reserva Flori'stica-Ecologica Rfo Guajalito, near Las Palmeras, 1800 m. QCAZ 2628, 3263-64, 3448, 4254-55, 4821-22; Rfo Baba, 4 km E, 10 km S Santo Domingo de los Colorados. 400 m.KU 142146-50; Rfo Faisanes, 13.5 km E LaPalma. 1 380 m. MCZ 94479; Santo Domingo de los Colorados. 600 m. KU 120251. 179394-96, 179399. 179401-03; 3 km E Santo Domingo de los Colorados, MCZ 98202; 2 km N. I km E Santo Domingo de los Colorados. 570 m. USNM 285795; Tandapi. 1460 m. AMNH 1 14495- 99, KU 1 1 1345 (holotype), 1 1 1346-73, 1 1 1375 (C&S). 120242-45, 120246 (C&S). 135445, 177218-25; 2 km W Tandayapa, 1 840 m, QCAZ 4788; Tinalandia, MCZ 94711. Eleutherodactylus pho.xocephalus (221 specimens) Ecuador: Prov. Azuciy: 10 km SW Victoria del Portete, 2700 m, KU 131281-82. Prov. Bolivar: San Vicente. MECN 286, 288. Prov. Canar: 18.4 km NW El Tambo, 2960 m, KU 142118-31. Prov. Canar: 15 km NW El Tambo, 2840 m, KU 166059-65. Prov. Cotopa.xi: Las Pampas, MHNG 18512, 18531 (2), QCAZ 595; Pilalo, 2460 m, KU 131404-78, 131479 (young). 131480-88, 131698-716, KU 142075 (holo- type), 142076-103. 177826-36. 180296,202428-29, MHNG (7), MHNG 18737 (4), QCAZ 554-58. USNM 233240-52; 14.5 km E Pilalo. 3160 m. KU 142816. Prov. Imbabura: La Delicia, 2710 m, KU 179387. Pichincha: 1.1 km E Chiriboga (Finca Santa Lucia), 2 1 20 m, KU 1 42063-7 1 ; 4 km W Chiriboga. 2 1 20 m. KU 142072-74; Los Alpes, 2500 m, KU 140876-77; 9.5 km NW Nono, 2530 m. KU 165559-60; Quebrada Zapadoi-es, 5 km ESE Chiriboga, 20 1 0 m, KU 1 65556- 58, 180247; San Ignacio, 2030 m.KU 109 137; Tandapi, 1460 m, MHNG 18715. Eleutherodactylus pteridophilus (52 specimens) Ecuador: Prov. Imbabura: above Cuellaje, 2560 m, ECO 143-44, 150-52. 156-60, 172; La Delicia, 2710 m, KU 132629. 179106 (holotype), 179107-39. Pwu Pichincha: 14 km W Chiriboga, 1960 m. KU 165926; 13.1 km NW Nono. 2150 m. USNM 286043; San Tadeo. near Mindo. USNM 239848; 6.2 km E Tandapi. 1750 m. MCZ 92109, 94721-22. Eleutherodactylus pyrrhomerus (32 specimens) Ecuador: Prov. Bolivar: Bosque Protector Cashca Totoras, 3000 m. KU 218030-33. Prov. Cotopaxi: Pilalo. 2400-2580 m.KU 13 1606 (holotype). 177837- 38; 1 km E Pilalo, 2075 m. USNM 233253; 1.5 km E Pilalo, 2200 m. KU 187481, USNM 233254-63; 3 km E Pilalo, 2900 m, KU 131607-11; 4.6 km E Pilalo, 2600 m, KU 152038; 5 km E. Pilalo. 2400 m. USNM 233264-65; 6 km E Pilalo. 2670 m. KU 142167-70. Prov. Imbabura: La Delicia. 2710 m. KU 179387. Erroneous locality: Prov. Pichincha: Tandapi, QCAZ 584. Eleutherodactylus quinquagesimus (70 specimens) Ecuador: Prov. Carchi: Maldonado, 1410 m. KU 179391; 22 km E Maldonado. 2560 m. QCAZ 4 187. Prov. Cotopa.xi: Reserva Otonga. 2200 m. QCAZ 4550. Prov. Imbabura: above Cuellaje. 2560 m, ECO 162, 173-74, 176-78; La Delicia, 2710 m, AMNH 114517-21, KU 132755-56, 132770, 132777-78, 179365-68. Prov. Pichincha: 14 km W Chiriboga, 1960 m, KU 167875-80; Quebrada Zapadores, 5 km ESE Chiriboga, 20 10 m.KU 166292-93 (skeletons). 167852- ELEUTHERODACTYLUS IN WESTERN ECUADOR 205 58, 167859 (holotype), 167860-73. 179374-77; 1 km SW San Ignacio. 1920 m, KU 179378-81 ; Tandayapa, USNM 239846; 9 km SE Tandayapa, 2150 m, KU 167881-86. Eleuthewdactylus rosadoi (7 specimens) Ecuador: Prov. Carchi: Rio Baboso, near Lita, MECN-LAC 70. 116. Prov. Esmeraldas: Rio San Miguel, 1 km above Rio Cayapas, MCZ 92937 (holo- type), 92938. Prov. Imbabura: Lita, 520 m, KU 132780; Prov. Pichinclui: Rfo Canoi. 4 km S San Vicente Maldonado, 570 m, KU 218051, QCAZ 7862. Biologica Rfo Palenque, 220 m, MCZ 90 140-42, 94459. Prov. Maiiabf: Parque Nacional Machalilla, Cerro San Sebastian (Bola de Oro). 750 m, EPN-AA 2437-39. 2458. 2460-61. Prov. Pichiiwha: Centinela. 14.1 km ESE Patricia Pilar, 550-600 m, USNM 285622-26, 285538. 285541-45; 5 km NW La Florida, MECN- LAC 209; Rfo Baba, 10 km S, 4 km E Santo Domingo de los Colorados, 400 m, KU 146172; Santo Domingo de los Colorados, 580 m, KU 177839-41. USNM 233320-34; 2 km E, 1 km S Santo Domingo de los Colorados, 600 m, KU 177842-44, USNM 285796; 9 km W Santo Domingo de los Colorados (Hda. Espinosa), CAS-SU: 10508, 10513, 10516. Eleiitherodactylus niidus ( 1 6 specimens) Ecuador: Azuay: Molleturo. 2317 m, AMNH 17588-89, 17590 (holotype). 17591-603. Eleiitherodactylus scolodiscus (9 specimens) Ecuador: Prov. Carchi: Maldonado, 1410 m. KU 177651; Rfo Baboso. near Lita, MECN-LAC 31; Rfo Blanco, 3 km up from Chical, 1450-1500 m, USNM 286292. Prov Esmeraldas: El Cristal, 1200 m, ECO 78-79, 101-103, 110. Eleiitherodactylus simonbolivari (32 specimens) Ecuador: Prov. Bolivar: Bosque Protector Cashca Totoras, 10 km SE Santiago. 3200 m, KU 218252-56. QCAZ 936, 940^1, 943-44, 1458, 1459 (holotype), 1460-75, 1493-96 (C&S). Eleiitherodactylus surdus (91 specimens) Ecuador: Prov. Imbabura: La Delicia, 2710 m. KU 130888-90, 177845-59. Prov Pichinclui: 5 km W Aloag, west slope Cerro Corazon, 2945 m, KU 1 09073; 7 km W Aloag, west slope Cerro Corazon. 2810 m, KU 109074, 109077 (C&S); Chiriboga, QCAZ 585, 774; Chontapamba, USNM 239844; Los Alpes, 21 km W Aloag, 2600 m,KU 111387-94, 117584-617, 130891- 96; Otongoro, MECN (2), QCAZ 566-68, 586; Quebrada Silante Grande, Aloag-Santo Domingo de los Colorados road, 2300 m. MCZ 97482. 97483 (ho- lotype ), 97484-85; Salto Dos Novias, north slope Cerro Corazon, 2635. KU 109075-76; 4.7 km SW San Juan, 14 km W Chillogallo, 3 190 m, KU 141972-73; 2 km E Tandapi. Quebrada La Plata. 1550 m, MCZ 98034, USNM 285971; 9.4 km SW Tandayapa, 2390 m, KU 202516. Prov. undetermined: "West Ecuador," BM 1947.2.17.25 (syntype); "South America," BM 1947.2. 17.26 (syntype). Eleiitherodactylus sobetes (2 specimens) Ecuador: Prov. Pichincha: Quebrada Zapadores. 5 km ESE Chiriboga, 1920 m, KU 179389 (holotype), 179390. Eleiitherodactylus subsigillatus (65 specimens) Ecuador: Prov. Azuay: 1 1 .2 km W Luz Marfa, 930 m, KU 218147; 12.9 km W Luz Marfa, 740 m, KU 218248, QCAZ 7863-64. Prov. Bolivar: Balsapamba, 830 m, KU 131566-71. Prov. Canar: Manta Real, Parroquia Molleturo, 650 m, EPN-AA 2538, 2551. Prov. Esmeraldas: Alto Tambo, 830 m, QCAZ 1297; Bilsa, 225 m. EPN-AA 2516; Hda. Equinox. 38 km NW Santo Domingo de los Colorados, 300 m, USNM 233335; La Boca, MHNG 18816; Salidero, 350 ft., BM 1947.2.17.1 (holotype). Prov. Los Rios: Estacion Eleiitherodactylus tenebrionis (40 specimens) Ecuador: Prov. Esmeraldas: Alto Tambo, 830 m, QCAZ 1294. 1303-04. Prov. Los Rios: Estacion Biologica Rfo Palenque. 220 m. KU 146171. 152574. 1 6587V78. MCZ 94864-65, 94867. 98 1 64-66, USNM 21 1 175. 285359. Prov Pichincha: Centinela. 570-600 m, MCZ 97596-97, USNM 2 1 1 1 76, 285627-28; Santo Domingo de los Colorados, 580 m, KU 179224-27; 2 km E, 1 km S Santo Domingo de los Colorados. 600 m, KU 1 79228-30; Tinalandia. 800 m, MCZ 88890. 90325, 90326 (holotype). 90327-29. 92079-81. 94712. Eleutherodactylus thymalopsoides (27 specimens) Ecuador: Prov Cotopaxi: Pilalo, 2460 m. KU 1 3 1 533 (holotype). 131534-37, 177861-80,202517-18. 206 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Eleuthcrodactylus truchae (47 specimens) Eleuthewdactyhis w-nigrum (686 specimens) Ecuador: Frov. Bolivar: Bosque Protector Cashca Totoras. Quebrada Ramoshuiacao, 3000 m. KU 2181 42-45. QCAZ 2470, 2474-75, 2493, QCAZ 7865- 67; 14.1 km E Guaranda, 3030 m, KU 218139. Pmv. Caiiar: 20 km N Gun. 2909 m. USNM 239649-65; 4 km N Zhud, 3040 m, KU 202619, 202620 (holotype). Prov. Chimborazo: 1.1 km SW Compud, 2570 m, KU 141974; 35 km SW Desvio Panamericana, 2860 m. KU 218140; 15 km N Pallatanga. 2860 m. USNM 239648; 20 km N Pallatanga, 3000 m, USNM 239647. Prov. Cotopaxi: 5 km E Pilalo, 3200 m, KU 130887; 24.6 km E. Pilalo, 3190 m, KU 218103-08; road from Pilalo to Latacunga. 2870 m. KU 218113-15. QCAZ7868-69. Eleutherodactylus unistrigatiis (104 specimens) Ecuador: Prov. Cotopaxi: Pilalo. 2400 m, KU 177888-969, 143557, 202535-36. Prov. Imbabitra: 35 km E Apuela, 3220. KU 218057-60. Prov. Pichincha: 5 km W Aloag, N slope Cerro Corazon, 2945 m, KU 1 09066-67, 7 km W Aloag, KU 1 09068-69, 1 6 km W Aloag, 2810 m, KU 111132-35; 9.5 km NW Nono, 2530, KU 165591-92 4.5 km W Nono, 2600 m, KU 218088-89; San Juan, 3400 m, KU 177518-20. Eleutherodactylus verecundus (14 specimens) Ecuador: Prov. Cotopaxi: Las Pampas, QCAZ 582- 83. Prov. Carchi: Rio Baboso, near Lita, MECN-LAC 32; Rfo Blanco, 3 km upstream from Chical, 1450- 1500 m, USNM 286291, 286294. Prov. Esmeraldas: El Cristal, 1 200 m, ECO 9 1 , 93-94, 1 08-09, 121. Prov. Pichincha: Reserva Flori'stica-Ecologica Rfo Guajalito, near Las Palmeras, 1800 m, QCAZ 4820; 6469, 6562. Eleutherodactylus vertebralis ( 1 50 specimens) Ecuador: Prov. Cotopa.xi: Pilalo, 2400 m, KU 120103-10, 131290-300, 131538-39, 142173, 165871-72, 179140-63, 179165-215, 202546-55; 1 km W Pilalo, 2340 m, KU 131301; 4.6 km E Pilalo, 2600 m, KU 152032-33. Prov. Imbabura: Intac, BM 1947.2.17.5-6 (syntypes); La Delicia, 2710 m, KU 132737-48, 132750-53, 132757, 177970-84. Prov. Pichincha: Lloa, 2800 m, QCAZ 7966-67; Quebrada Zapadores. 5 km ESE Chiriboga, 20 1 0 m, KU 1 792 1 6; 3 km from Tandayapa, road to Quito, 1800 m, MCZ 90145. EcuADOR:Prav.y4cMav.Cuenca, CAS 63956; 10-15 km E Luz Maria, 2500 m, KU 218125-32, QCAZ 7870-77; 0.9 km W Luz Marfa, 1 770 m, QCAZ 7878; 1 1 .2 km W Luz Marfa, 930 m, KU 2 1 8 147; 12.9 km W Luz Marfa, KU 2 1 8 1 48-49; MoUeturo, 7600 ft, AMNH 17640-43; Zurucuchu, SMF 3804 (holotype) [only photograph examined]. Prov. Bolivar: Balsapamba, AMNH 33916-17; 10 km NE Bilovan, 2540 m, KU 141851-59; Bosque Protector Cashca Totoras, 3000 m, KU 218142-45; Guaranda, 2640 m, KU 131263- 7 1 ; 4.7 km E Guaranda, 2700 m, KU 2 1 8 1 2 1-22; 1 1 . 1 km E Guaranda, 2600 m, KU 202572; 29 km E Guaranda, 27 10 m, KU 2 1 8 1 36-38; 2.5 km S Guaranda, 2550 m, KU 141 293; 4.6 km S Guaranda, KU 202569- 70; 5.4 km S Guaranda, 2680 m, KU 179217-18; Hacienda Lizo, Rfo Tatahuazo, 2510 m, KU 218120; San Vicente, MECN 284-85. Prov. Carchi: 5 km W La Gruel, 2340 m, KU 202561, 202571, 202573; Maldonado, 1410 m, KU 177533-34; 14 km SE Maldonado, 2500 m, KU 177535; 27.3 km E Maldonado, 2420 m, KU 218133-35; Rfo Blanco, 3 km upstream from Chical, 1450-1500 m, USNM 286295. Prov. Chimborazo: Las Cochas, N of Huigra, 8100 ft, CAS-SU 9477; Paitanga, BM 80.12.5.273; 20 km NNE Pallatanga, 2550 m, KU 165840^3; 35 km SW junction Panamerican Hwy on road to Pallatanga, 2860 m, KU 218140. Prov. Cotopaxi: Las Pampas, QCAZ 607; 4.6 km E Pilalo, 2600 m, KU 152034-37; 10.5 km W Pilalo, 1520 m, QCAZ 7879. Prov. El Oro: El Chiral, AMNH 13960. Prov. Imbabura: 16.1 km E Apuela, 2400 m, MCZ 95539; 1 6.7 km E Apuela, 2470 m, MCZ 95537-38; 1 7.4 km E Apuela, 2500 m, MCZ 95544; Intac, BM 78.1.25.23-24; La Delicia, 2700 m, KU 130454-57, 177536-41; Quebrada San Miguel, 1 km N Otavalo, 2560 m. KU 1 1777 1 . Prov. Pichincha: Aloag-Santo Domingo road. 2900 m, MHNG 18724 ( 1 1 ); 7 km W Aloag. 2810. KU 1 09064; Bosque Protec- tor Mindo-Nambillo. 1700 m. QCAZ 7308; 9.3 km W Calacalf, 2380 m; 14.7 km W Calacalf, 2130 m, KU 218123-24, QCAZ 7880-81; 2 km W Campamento Silante, 2100 m, KU 130463-64; Chiriboga, BM 1940.2.20.1, EPN 1379, MECN 355, QCAZ 3227; 7.7 km E Chiriboga (Finca Santa Lucfa), 2120 m, KU 141836-43; 14.8 km EChiriboga, 2540 m, KU 141 833- 35 177564; 2.2 km W Chiriboga, 1 980 m, MCZ 98 182- 83 ; 4 km W Chiriboga, 2 1 20 m, KU 1 4 1 844-50; 1 4 km W Chiriboga. 1960 m, KU 165838-839, 166232 (young), 209553-55 (C&S); 14.4 km E La Palma, 1380 m, MCZ 92060-66; 16 km E La Palma, 1500 m, MCZ 90308; 1 8 km E La Palma. 1600 m, MCZ 90294- ELEUTHERODACTYLUS IN WESTERN ECUADOR 207 307; 20.6 km E La Palma. 1 600 m, MCZ 92056-58; 22 km E La Palma. 1 770 m. MCZ 90287-93; 23 km E La Palma. 1900 m, MCZ 90276-86; 25.7 km E La Palma. 1820 m, MCZ 92059. 94488. 94490-92. 94859-61, 949 1 9^4; Las Maquinas, AMNH 20 1 44; Las Palmeras. MECN 354. QCAZ 884; Loma Las Penas, 2 km due S Tandapi. 1 600 m. MCZ 9023 1 ; Los Alpes. 2600 m, KU 117772-73. 130458-62; Milligali. MCZ 3008; 3.5 km NE Mindo. 1540 m. KU 165733-39; above Mindo, 1500-1610 m, MCZ 113474. 113485; Mindo (Hda. San Vicente), 1465-1620 m, USNM 284356-57, 284360; road to Mindo, 7500 ft. UMMZ 55504; 10 km N Nono (road to Tandayapa). 2200 m. MCZ 90232-34; 15 km N Nono (road to Tandayapa). 2100 m, MCZ 90235-38; 9.5 km NW Nono, 2530 m, KU 1 65728-32; 10 km NW Nono (road to Nanegal), 2200 m. MCZ 90240-43; 12 km NW Nono (road to Nanegal). MCZ 90244-75; Quebrada La Plata. 3 km SE Tandapi. 1560 m. MCZ 90227-30; Quebrada Zapadores, 5 km ESE Chiriboga.2010m,KU 165740-82. 166287-88 (skel- etons), 1 77565-66; Rio Faisanes, 13.5 km E La Palma. 1380 m, MCZ 92864-7 1 , 92880-8 1 , 93447-48, 94480- 81, 94906-16, 95629; San Ignacio, 2030 m,KU 109065, 109070; 1 km W San Ignacio. 1900m.KU 177542-63; Tandapi, 1460 m, KU 110975-77 (C&S), 110978- 1084, 135339-42. 135447, 135469, 135471-82, 135485-86, 135489-90, 138779-87, MCZ 75164-69, QCAZ 606; 2 km E Tandapi, Quebrada La Plata, 1550 m. USNM 285967-69; 2.1 km E Tandapi. 1500 m, MCZ 92841-45. 94824-36, 95612, 95483-92; 6. 1 km E Tandapi. 1650 m. KU 177567; 6.2 km E Tandapi, 1 700- 1750 m, MCZ 92067-78, 93432-5 1 ; Tandayapa, 2300 m, CAS-SU 10351-52, MCZ 88417-18, USNM 239845; 7 km SE Tandayapa, 1900 m. MCZ 90239; 9 km SE Tandayapa, 2150 m, KU 165689-727; 1.4 km SW Tandayapa, 1820 m, KU 202591; 2.3 km SW Tandayapa, 1900 m, KU 202587-90; 9.4 km SW Tandayapa, 2390 m, KU 202574-86; Tinalandia, 800 m, MCZ 88419, 88423-25. Prov. undetermined: West Ecuador, BM 60.6.16.95, 60.6.16.98, 60.6.16.100, 60.6.16.102-03, 60.6.16.107. Erroneous locality: Prov. Pichincha: Santo Domingo de los Colorados, FMNH 172461. Eleutherodactylus walkeri (407 specimens) Ecuador: Prov. Azuay: 1 1.2 km W Luz Maria, 930 m, QCAZ 7882; 12.9 km W Luz Maria, 740 m, KU 218149, QCAZ 7883; 13.5 W Luz Maria, 740 m, KU 218119; Manta Real, Parroquia MoUeturo, 650 m, EPN-AA 2543-45, 2552-54, 2584. Prov. Bolivar: Balsapamba, 800 m, KU 131613-51; 6 km ESE Balsapamba, 1270 m, KU 142039-40. Prov. Cotopaxi: 0.7 km N Las Juntas, 320 m, USNM 286068; 1 1 km E Moraspungo, 840 m, USNM 286085; 1 8.6 km W Pilalo, 930 m. KU 1 42 172. 20.3 km W Pilalo. KU 142(J34-38, 142171. Prov. El Oro: 1 1.7 km W Pinas, 708-800 m, USNM 286253-56, 286258-62, 286269-76; 32.6 km SSE Portovelo, 990 m, KU 142041 . Prov. Esmeraldas: 3.9 km E Durango, QCAZ 8072; Hda. Equinox, 38 km NW Santo Domingo de los Colorados, 300 m. USNM 233423-24; vicinity of San Miguel, MCZ 92908, 92911-18,92921-25,92933,92935-36,92941,92950- 5 1 , QCAZ 171. Prov Guayas: Rio Frio, UMMZ 1 23900. Prov. Los Rios: Estacion Biologica Rio Palenque, 220 m, KU 146173, 147567-68, 165653-58, MCZ 90218, 92054, 94842, USNM 285360-62. Prov. Manabi: en- virons of Guale, Rio Ayampe, 500 m, QCAZ 4227; Parque Nacional Machalilla, Cerro San Sebastian ( Bola del Oro), 750 m, EPN-AA 2436. Prov. Pichincha: Centinela, 14. 1 km ESE Patricia Pilar, 550-600, USNM 285629-68; 4 km E Dos Rios, 1 1 20 m, KU 202557-59; 5 km NW La Florida, QCAZ 601-02; 5 km W La Florida, 860 m, KU 2 1 8 1 1 6- 1 7, QCAZ 406 1 . QCAZ 7884-85; La Palma, 920 m, CAS 134064-66, KU 131652 (holotype), 131653-63, 178013-14; 1-2 km E La Palma, MCZ 90206-08; 19.3 km E Patricia Pilar, 430 m. USNM 285687-90; Rio Baba. 5-10 km SSW Santo Domingo de los Colorados, 500 m, AMNH 89733-36; Rio Baba, 10 km S, 4 km E Santo Domingo de los Colorados, KU 142013-30; Rio Baba, 19kmS, 5 km E Santo Domingo de los Colorados, UIMNH 93560-6 1 ; Rio Canoi, 4 km S Vicente Maldonado, 570 m, KU 218118. QCAZ 7886; Rio Toachi. USNM 233439-40; San Miguel de los Colorados, USNM 233441; Santo Domingo de los Colorados, 600 m, FMNH 174092, 172464, 174334, KU 120215-21, 120227-31, 177985-8009; 2 km E, 1 km S Santo Domingo de los Colorados, KU 178010-11, 178012 (C&S); 6 km E Santo Domingo de los Colorados (Dyott Farm), USNM 233425-37; 8 km SE Santo Domingo de los Colorados (Hda. Delta), UMMZ 1 3 1 675 ( 10); 9 km W Santo Domingo de los Colorados (Hda. Espinosa), CAS-SU 10514. 10517; 18 km W Santo Domingo de los Colorados (Ramsey Farm). 650 m, USNM 233438; Tinalandia. MCZ 88395-411, 88421, 90146-204, 90209-17, 90219-26. 90334., 92055, 92851-63, 94710, 95482 USNM 217415; Toachi, 860 m, QCAZ 7887; 1 km E Vicente Maldonado. 570 m, QCAZ 7888. 208 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 APPENDIX II Gazetteer Following is a list of localities on the PaciHc lowlands and western slopes of the Andes in Ecuador where specimens of Eleutherodactylus were collected. After each place name the province is given in parentheses, followed by geographic coordinates, elevation and bioclimatic regime (see section on Western Ecuador). Spellings and coordinates given to seconds were taken from the Indice Toponfmico de la Repiiblica del Ecuador (no date); others were determined from the 1974 and 1991 editions of Mapa de Ecuador, Instituto Geografico Militar. Quito (1:1 ,000,000) and Salvador et al. (1995). Elevations were determined from altimeter readings at the sites or from listings in the Indice Toponfmico de la Repiiblica del Ecuador (1972). When known, specific sites and habitats are given, followed by names of collectors and months and years that they collected at the sites. Names of some of the collectors are abbreviated: JAP = James A. Peters. JDL = John D. Lynch, KM = Kenneth Miyata, LAC = Luis A. Coloma, RWM = Roy W. McDiarmid, SRE = Stephen R. Edwards, WED = William E. Duellman. The boldfaced number at the end of each entry refers to the number on the map (Fig. 87). Alhuiqiim (Pichincha)— 00°23' S, 78°58' W, 700 m; humid subtropical. Village on the lower slope of the Cordillera Occidental on the Aloag-Santo Domingo de los Colorados road. Collections made in disturbed humid lower montane forest at 1 2 km E, 940 m, by LAC in July 1989, and at 13 km E by KM. 69 Along (Pichincha)— 00°28' S. 78°35' W, 2890 m; humid temperate. Village on the highway from the junc- tion of the Panamerican Highway to Santo Domingo de los Colorados, which borders the north slope of CeiTO Corazon. Collections were made in remnants of humid upper montane forest at 5 km W Aloag (2945 m) and 7 km WofAloag (28 10 m) by WED in March 1967, and 16 km W Aloag (28 1 0 m) and 21 km W Aloag (2640 m) by JDL in July 1967.79 Alto Tambo (Esmeraldas)— 00°5r42" N. 78°30'54" W, 830 m; humid subtropical. A village on the Ibarra-San Lorenzo railroad and Rio Mira. Collection in disturbed lower humid montane forest by Stella de la Torre and others in December 1990. 15 /\/7(/('/a(Imbabura)— 00°I9"2I"N,78°30"36"W, 1670 m; humid subtropical. A village on the Pacific slope of the Cordillera Occidental. Collections were made in upper humid montane forest at various distances along the road from Apuela to Otavalo — 16. 1 km E (2400 m), 16.7 km E (2470 m). 17.4 km E (2500 m) by KM. 22 km E (2860 m) by LAC in June 1989, and in paramo 35 km E (3220 m) by LAC in June 1989. Collections also were made in the Cordillera de Intag. 2190 m. 23.2 km (by road) W of Apuela, by JDL in January 1978. 28 Ayampe (Guayas)— 01°40'31" S. 80°48'36" W. 12 m; dry tropical. A village and river 25 km N of Montafiita on the boundary of Provincia Manabf and Provincia Guayas. Collection from dry forest at edge of river 3.7 km E. 70 m, by LAC. 103 Balsapamha (Bolfvar)— 0r47' S, 79° 10' W. 800 m; dry subtropical. Village on the lower slopes of the Cor- dillera Occidental. Collections made in cutover forest in the immediate vicinity of the village by JDL in July 1 970 and 6 km ESE on the road to Chillanes. 1 270 m by WED in July 1 97 1 . Collection from •' 1 0 km E, 9500 ft" by JAP in June 1954. 109 Biblidn (Canar)— 02°42"42" S. 78°53'I2" W, 2630 m; humid temperate. Town in the northern part of the Hoya de Cuenca. Collections from disturbed subparamo at 0.5 km N, 2620 m by R. R. Montanucci in May 1 97 1 , and 10 km NE. 2540 m by WED in July 1971. 126 Bilovdn (Bolivar)— 0r48' S. 79°05' W; humid tem- perate. Village on western slopes of Cordillera Occiden- tal. Collection from 1 0 km NE. 2540 m. by WED in July 1971. 109B Bilsa (Esmeraldas) — See San Jose de Bilsa. Bold del Oro (Manabf) — See Parque Nacional Machalilla. Bosque Protector Cashca Totoras (Bolfvar) — 0r42' S. 78°53' W. 3000 m; humid temperate. A reserve in the uppermost drainage of the Rfo Chimbo on the Pacific slope of the Cordillera Occidental. Collections from humid upper montane forest by LAC in December 1986, November-December 1987, February 1989. and January 1990. 104 Bosque Protector Mindo-Nambillo (Pichin-cha) — See Mindo. Bucay (Guayas)— 02° 10' S. 79° 06' W. 275 m; dry tropical. Village on the railroad and Guayaquil-Riobamba road at the mouth of the Rfo Chimbo Valley at the base of the Cordillera Occidental. Collection by G. H. Tate in December 1921 and March 1922. 116 Buemi Fe (Los Rfos)— 00°53"40" S. 79°29'15" W, 200 m; humid tropical. Small village on the highway between Santo Domingo de los Colorados and Quevedo and situated in area predominately cultivated in bananas. ELEUTHERODACTYLUS IN WESTERN ECUADOR 209 Collections made I km N Buena Fe by KM in January 1979. 86 Buenavista (El Oro)— 03°2 r2 1" S. 79°50' 1 6" W, 27 m; dry tropical. Village on road 6 km SW Pasaje on Pacific lowlands; cutover semideciduous forest. Collec- tions made at 7 km SE, 30 m, by JAP in August 1 954. 134 Biiliun (Esmeraldas) — See San Francisco. Cac/zafee (Esmeraldas) — See San Javier de Cachabe. C«/«cY///'(Pichincha)— 00°00"I0" N. 78°30'40" W, 28 15 m; humid temperate. A small village in the Cordil- lera Occidental. Collections were made on the western slope of the cordillera at a point 14.7 km (by road) SW of Calacali on the road to Nono, 2 1 30 m by LAC in March 1990.43 Campamento Silante (Pichincha)— 00°26' S, 78°42' W, 2100 m; humid temperate. Road maintenance camp on highway between Aloag and Tandapi on the Pacific slope of Cordillera Occidental; disturbed humid upper montane forest. Collections in vicinity of camp and 2 km W 2100 m by JDL in August 1970. 73 Ca/7a/-(Canar)— 02°33'20" S, 78°56' 15" W, 3 140 m; humid temperate. Town in the Cordillera Occidental of the Andes. Collections made in disturbed low forest 5.6 km NNW. 2920 m by WED in March 1984 and 28.2 km N, 2915 m by R. R. Montanucci in May 1971. 122 Carondelel (Esmeraldas) — See Parroquia Caron- delet. Catov/w/(Azuay)— 03°I5"I2"S.79°15'42"W. 1580 m; dry subtropical. A hacienda in the valley of the Rio Jubones between Santa Isabel and Giron on the Pacific slope of the Cordillera Occidental. Collection from 4 km SW. 1600 m by JAP in February 1959. 130 Centinela, 14.1 km (by road) SE Patricia Pilar (Pichincha)— 00°35' S, 79°2r W, 550-600 m; humid subtropical. A site on a north-south ridge, Montaiias de Ila, on the Pacific coastal plain east of the Rio Palenque; humid lower montane forest, now cutover; collections by KM and RWM in January and October 1979. 81 Centra Cienti'fico Rio Palenque (Los Rfos) — 00°34' S, 79°20' W, 200-220 m; humid tropical. A biological station of 167 ha 56 km N Quevedo; humid tropical rainforest, oil palm plantation, and banana plantation. Collections made C. F. Walker and E. H. Taylor in August 1962, by WED in April 1972 and March 1975, by D. C. Dodson in January 1972, by J. E. Simmons in July 1972, by G. O. Vigle in December 1 977, by RWM in December 1978, and by KM and RWM in January and October 1979.82 Cerro San Sebastian (Manabi ) — See Parque Nacional Machalilla. Chinibo (Chimborazo)— 02°09' S, 79°05' W. 345 m; dry subtropical. Railroad bridge over Ri'o Chimho above Bucay at base of Cordillera Occidental. 115 Chirihoi>a (Pichincha)— 00° 13'42" S, 78°46'00" W, 1910 m; humid temperate. A small village on the "old road" between Quito and Santo Domingo de los Colorados. Collection from disturbed cloud forest in immediate vicinity by C. Fierro in May 1986 and May 1988. Collections from disturbed cloud forest at various distances along the road to the east and west of Chiriboga— 10 km E by JAP in March 1959; 14.8 km E (2410m)by WEDin July 1971 and by JDL in January 1978; 0.6 km W( 1880m) by RWM in March 1979.4 km W (2120 m) by WED in July 1971; 8 km W by JAP in May 1959; 14 km W( 1960 m) by WED in May 1975; and 25.8 km W ( 1 720 m; humid subtropical ) by WED in July 1971.50 Chontapamba (Pichincha) — Not located. Collection by Manuel Olalla in April 1955. awip//J(Chimborazo)— 02°20"28"S,78 56"03"W, 2570 m; humid temperate. A village on the Panamerican Highway in the Cordillera Occidental. Collection from disturbed humid upper montane forest 7.7 km SW, 2570 m by WED in July 1971. 119 Connina San Vicente de Asacoto (Chimborazo) — Dry temperate. A settlement at 2100 m in the Canton Pallatanga on the Pacific slope of the Cordillera Occiden- tal. Collection from small areas of secondary forest by A. Almendariz in September 1986. Not located. Cordillera de Chi I la. Llanos de Guavos (El Oro) — Mountain chain SW of Chilla (03°29' S, 79°36' W). Collection by H. E. Anthony in August 1920. 134 Cruz de Lizo (Bolivar)— Or43'30" S, 78°59-54" W. 2720 m; humid temperate. A village in the western Andean Chimbo Basin. 3.1 km S of Santiago. Collec- tions from disturbed areas by LAC in September 1983. 108 Cuellaje ( Imbabura)— 00°22' N, 78°32' W, - 1 500 m; humid subtropical. A settlement on the west slope of the Cordillera Occidental, below (west of Apuela). Collec- tions from "above Cuellaje, 2560 m" by A. Quiguango and others in January and September 1994. 26A Ciienca (Azuay)— 02°55' S, 79°00' W, 2543 m; hu- mid temperate. City in an inter-Andean basin. 128A De.viw (Manabi)—0 r03' S, 79°57' W dry tropical. A site on the Quevedo-Manta road, 16 km W of Pichincha in the Cordillera de Balzar. Collection from 2 km W, 250 m, by JAP in October 1965. 92 Desvio Panamericana (Chimborazo) — 0r42' S, 78°46' W, 3200 m; humid subtemperate. Junction of Panamerican Highway and road to Guayaquil. Collec- tion from disturbed upper montane forest 35 km SW on road to Pallatanga, 2860 m by LAC in April 1990. 105 Dos Rfos (Pichincha)— 00° 19' S, 78°5r W. 1270 m; humid subtropical. A small village (= 10 km NE Las Palmas) on the "old road" between Quito and Santo Domingo de los Colorados; disturbed lower montane 210 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 -CD •• ■•. ■ 1 I ELEUTHERODACTYLUS IN WESTERN ECUADOR 21 212 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 rainforest. Collections by T. H. Fritts in July 1970. Collections also made along the road — 3 km W ( 1 050 m ) by WnO in July 1971 and 4 km NE ( 1 140 m) in April 1975 and March 1984.65 Diircmgo (Esmeraldas)— 00°53' N. 78°48' W, < 300 m; humid tropical. A settlement on the Pacific coastal plain. Collection from lowland tropical rainforest by LAC and others in March 1994. 13 Echeamlia (Bolivar)— 01 '25"58" S. 79° 16" 15" W. 280 m; dry tropical. Small village on coastal plain east of Ventanas. Collection by R. Jimenez. 97 El Carmen (Manabi)— 00°16'08" S, 79°28' 20" W. 250 m; dry tropical. Town on Pacific lowlands. Collec- tion from 50 km WSW on road to Chone, 400 m; and 38 km NW on road to Pedernales, 330 m, by LAC, Decem- ber 1989.58 £/C/;/ra/(E10ro)— 03°40"36" S,79°28'48"W, 1630 m; dry subtropical. A settlement north of Zaruma on the south face of the Cordillera de Chilla. Collection by H. E. Anthony in July 1920. 136 El Corcovado (Cotopaxi)— 0r07'36" S, 79°07' 12", 2000 m; humid temperate. Settlement 3.5 km NW of El Corazon on western slope of Cordillera Oriental. Collec- tion by G. Onore in July 1991. 93 El distal (Esmeraldas)— 00°45' N. 78 30' W. 1200 m; humid subtropical. Settlement on the north slope of the Cordillera deLachas, SSWof Lita. Collection by M. L. Crump and others in December 1993 and by A. Quiguango and others in August 1994. 23 El Placer (Esmeraldas)— 00°5r 18" N, 78°33'42" W; dry tropical. Settlement on the Pacific coastal plain. Collection from 2-4 km W, 360-390 m by JAP in Novem- ber 1958. 16 El Tambo (Caiiar)— 02°31' S. 78°56' W, 2960 m; humid temperate. Village on Panamerican Highway in valley of upper Rfo Caiiar (Pacific drainage ). Collections from disturbed upper montane forest 1 5 km NW, 2840 m by WED in July 1 97 1 and from subparamo 1 8.4 km NW, 2960 m by WED in July 197 1 . 121 El Torueado (Bolivar)— 0r45'25" S, 79°08"08cT W. 2430 m; dry temperate. A site 6.4 km NE of Balsapamba. Collection by A. Almendariz in June 1987. 129A Estacion Biologica Rio Palenque (Los Rios) — See Centro Cientifico Rio Palenque. Estacion Forestal La Favorita (Pichincha) — Parroquia Chiriboga. 00°13"42" S, 78°45"54" W, 1900 m; humid temperate. A small field station located on the "old road" between Quito and Santo Domingo de los Colorados Pacific slopes of the Cordillera Occidental. The area of the station is mostly secondary forest amid remnants of upper humid montane forest; collections by A. Almendariz in July 1987. 51 Fifjca La Espenmza (Pichincha)— 00° 15' S. 79°09' W. 500 m; humid subtropical. A farm just NW of Santo Domingo de los Colorados. Collection by C. Duran. 55 Finca Santa L/k/'^/ (Pichincha)— 00 15' S. 78°42' W, 2120 m; humid temperate. A small farm 7.7 km E Chiriboga; disturbed humid upper montane forest. Col- lection by WED in July 1971. 54 Galapagos (Pichincha)— 00°24'2 1 " S. 78°58'00" W. 1 720 m; humid subtropical. A village in humid montane forest 2.5 km N of San Francisco de Las Pampas. Collec- tions by G. Onore. 71 Guainche (El Oro) — Collection by G. H. Tate in August 1921. Not located. Possibly this is Guaynche, Provicia Loja, 04°02'17" S, 79°59'13" W. 1 120 m. Guale (Manabi)— Or37"38" S. 80° 14" 15" W. 60 m; dry tropical. Collection from Finca Canta La Piedra on the Rfo Ayampe in the vicinity of Guale by Ana Agreda and Luis Vinueza in September 1993. 101 Gimlea (Pichincha)— 00°07'00" N. 78°44"48" W. 1300 m; humid subtropical. Settlement 3.9 km NW Nanegalito on Pacific slope of Cordillera Occidental. Specimens in BM. 33 Giialtaco{E\ Oro) — Not located; collection by Ramon Ollala in February 195 1 . Gm/n/(yV>(Bolfvar)— 01 °33"20"S. 79^00*24" W, 2900 m; humid subtemperate. Village in the upper Rfo Chimbo valley north of Guaranda. Collection in disturbed upper montane forest by LAC in December 1984. 98 G»flra7!Jrt (Bolfvar)— 0r35" 1 8" S, 78°59'58' W. 2670 m; humid temperate. Town in the upper Rfo Chimbo valley on the Pacific slope of the Cordillera Occidental. Collections from disturbed upper montane forest in im- mediate vicinity of town by JDL in July 1 970 and 4.7 km E (2700 m) by LAC in January 1990. Collections from subparamo along road to Riobamba — 1 1 . 1 km E (2600 m ) by WED in March 1 984, 1 4. 1 km E ( 3030 m ) by LAC in April 1990, and 29 km E (2710 m) by LAC in July 1989. Collections from disturbed upper montane forest on road to San Miguel and Chillanes — 2.5 km S ( 2550 m) by R. R. Montannuci in May 1 97 1 . 4.6 km S (24 1 0 m) by WED in March 1984. and 5.4 km S (2680 m) by JDL in January 1978. 99 Giiatea (Pichincha) — See Gualea. Guayas (Guayas)— 00°58'36" S, 79°23"45" W; hu- mid tropical. Village on Pacific coastal plain NW of Quevedo. Collection from 20 km W (90 m; dry tropical) USNM. 90 Gun (Cahar) — See Zhud. Hiiigra (Chimborazo)— 02°17'"06" S, 78°58"48" W. 1230 m; dry subtropical. Settlement on the railroad 6.9 km W of Chunchi in Rfo Chanchan Valley on the west slope of the Cordillera Occidental; subtropical, semiarid forest. Collection by S. N. Rhoades in 1911. 118 Intac (Imbabura) — See Intag. ELEUTHERODACTYLUS IN WESTERN ECUADOR 213 Intag (Imbabura)— 00°24' N, 78°36' W, 1200 m; humid subtropical. Settlement on north slope of Cordil- lera de Intag; lower humid montane forest. Collection by Buckley for BM. 26 Km 94(Giia\a(/iiil-Ciienc(i mad) — Not located. Col- lection by G. R Frymire and C. H. Dodson in January 1958. La5o(T/(Esmeraldas)— 0r08'N.78°46'W.<100m; humid tropical. Village at junction of Rio Bogota and Rio Tululbi. lA La Concordia (Pichincha)— 00°00" 1 5" N, 79°23' 12" W. 180 m; humid tropical. A village on the Pacific lowlands. Collections from a remnant of humid tropical rainforest, Bosque Protector La Perla. 190 m by LAC in April 1990. and from 5.3 km W. 190 m by LAC in April 1990.42 La Delicia (Imbabura)— = 00°22' N. 78°25' W. 2710 m; humid temperate. Settlement on road between Otavalo and Apuela near crest ofCordillera Occidental; disturbed upper humid montane forest. Collections by SRE in August 1 970 and by JDL in June 1977 and January 1978. 27 La Esperanza (Cotopaxi)— 00°54' S, 79°03' W. 1500 m; humid subtropical. Settlement on road and Quevedo and Pujili; cutover montane forest. Collection by JDL in July 1970. 87 La Florida (Pichincha)— 00° 16'48" S. 79°OrOO" W, 1 000 m; humid subtropical. A village on the lower slopes ofthe Cordillera Occidental, 1 5 km SE of Santo Domingo de los Colorados. Collections from 5 km NW, 860 m by LAC in September 1 984, December 1 989, and May 1 993 by LAC. 59 Lagarto, Resen'a Mayronga (Esmeraldas) — OTOO' N, 79° 1 3' W, 1 00 m; humid tropical. A village on the Rio Lagarto on the Pacific coastal plain. Collection from forest reserve by G. Onore in October and December 1993. 5 La Gruel (Carchi)— 00°53' N, 78°05' W, 2500 m; humid temperate. A village on the road between Tulcan and Maldonado. Collections from subparamo 2 km E, 2590 m, and from disturbed upper humid montane forest 5 km W, 2340 m by WED, February 1984. Name was provided by local inhabitants but does not appear on maps; possibly official name is Laurel, which does appear on recent maps. 12 Ui Palma (Pichincha)— 00°22'40" S, 78°55"49" W, 920 m; humid subtropical. A small village at the junction ofthe "old" and " new" roads between Quito and Santo Domingo de los Colorados; disturbed lower montane rainforest. Collections by JDL in August 1970 and July 1 977. Collections also from various distances and eleva- tions along the "old road" (via Chiriboga) — 3 km NE (1540 m) in April 1975 and 5 km E (900 m) in March 1 975 by WED; 1 . 1 km E, 1 0.4 km E, 10.6 km E, 1 2.6 km E. 13km E, 14.4km E, 16km E(1500m), 18km E, 20.6 kmE(1600m),22kmE(1770m),23kmE(19()0m),and 25.7 km E ( 1 820 m ) by KM in January and October 1 979. 68 La.vCoc/za5(Chimborazo)— 02°14'48"S,78°59'36" W, 2520 m; humid temperate. A site northwest of the village of Huigra in the upper Rio Chanchan valley. Collection by I. R. Wiggins in October 1944. 117 Las Juntas (Cotopaxi)— 01° 1 1 "06" S, 79° 1 3"48" W; humid subtropical. Village in foothills of the Andes. Collection from 0.7 km N (320 m) by RWM in October 1979.95 LasMdquinas{Pkh\ncha)—00°02'54" S, 78°36"45" W, 2800 m; humid temperate. A site 4.4 km NW of Nono on the Pacific slope ofthe Cordillera Occidental; humid montane forest. 45 Las Palmas (Pichincha) — See La Palma. L«,vP«//»m;.s (Pichincha)— 00° 12'00" S, 78°49\36" W, 1800 m; humid subtropical. A settlement 9 km W Chiriboga on the Pacific slope ofthe Cordillera Occiden- tal; disturbed humid montane forest. Collection by LAC in October 1984.49 Las Pampas (Cotopaxi) — See San Francisco de Las Pampas. Laurel (Carchi) — See La Gruel. L/Yo (Imbabura)— 00°50'24"N,78°27" 18" W, 570 m; humid subtropical. A village on the Ibarra-San Lorenzo railroad. Collections by Manuel Ollala in April 1 959 and from disturbed humid lower montane forest in the vicin- ity of the village by SRE in August 1970, LAC in September 1984. and Stella de la Torre and others in December 1990. 18 Lizo (Bolfvar) — See Cruz de Lizo. Llambo (Pichincha)— 00°01' N, 78°40' W; humid temperate. A settlement on the road between Nono and Gualea on the Pacific slope ofthe Cordillera Occidental; humid montane forest. Collection by C. Ollala in April 1929.40 Lloa (Pichincha)— 00 09' S. 78°35' W. 2800 m; hu- mid temperate. A village on the south slope of Volcan Pichincha. Collection by A. Quiguango and A. Rufz in March 1994. 58A Lonui Las Pemis (Pichincha)— 00°27' S, 78°46' W, 1600 m; humid subtropical. A hill 2 km due S Tandapi; humid montane forest. Collection by KM in June 1975. 78 Los Alpes (Pichincha)— 00°26' S, 78°39' W, 2500- 2600 m; humid temperate. A settlement 20 km W Aloag on the road to Santo Domingo de los Colorados; dis- turbed upper humid montane forest.. Collections by JDL in July 1967 and August 1970. 75 Luz Maria (Azuay)— 02°4ri8" S. 79°24'36" W, 214 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 1880 m; humid temperate. A village 7.5 km NW of Moileturo on the Pacific .slopes of the Cordillera Occi- dental. Collections were made in humid montane forest at 1-15 km E (2500 m) along the road from Luz Mari'a and in cloud forest (humid subtropical regime) 7.7 km W (1300m), 1 1.2 km W (930m), 1 2.9 km W (740m), 13.5 km W (740 m) by LAC in April 1990. 125 Mflc7!a/fl(E10ro)— ()3°15"24"S,79°57'20"W,35m; dry tropical. A town on the Pacific coastal plain; tropical dry forest. Collection from 7 km SE, 10 m, by JAP in March 1959. 131 MachaUlla (Manabi) — See Parque Nacional Machalilla. Macuchi (Cotopaxi)— 00°55'36" S, 79°02"48" W. 1600 m; humid subtropical. A settlement on the Rfo San Pablo 2 km SE of El Tinge on the road to Pilalo. Collection by JAP in January 1959. 88 Maldonado (Carchi)— ()0°54\39" N, 78°06'30" W, 1410 m; humid subtropical. Village in the valley of the Ri'o San Juan on the Pacific slopes of the Cordillera Occidental. Cut over cloud forest. Collection in vicinity of village and 14km(airline)SE(2500m)byJDL in May 1977. Collections from 22 km E along road to Tulcan (2560 m; humid temperate) by LAC in March and July 1993, from 26.9-27.3 km E along road to Tulcan (2420 m; humid temperate) by LAC in June 1989 and from 14 km W on road to Lita (1255 m) by LAC in March 1993. 10 Manglaralto (Guayas)— 0r50'40" S, 80°44'30" W, 5 m; dry tropical. Village on Pacific coast; tropical dry forest. Collections from along road N to Puerto Lopez — 23 km N (60 m) and 25 km N ( 120 m) by KM in August 1975. Ill Manta /?£'«/ (Caiiar)—02°34' S, 79°2 1 ' W, 250 m; dry tropical. A village at the base of the Andes. Collections were made above the village in cloud forest at an eleva- tion of 650 m by Ana Almendariz in July 1 99 1 . 123 Miligali (Pichincha)— 00°16' S, 78°46' W, 1625- 2000 m; humid subtropical. Settlement on western slope of Cordillera Occidental; in subtropical forest according to Paynter and Traylor (1977). Collections by Manuel Ollala in January 1955.60 Millegale (Pichincha) — See Miligali. Milligalli (Pichincha) — See Miligali. Milpe (Pichincha)— 00°02'42" N, 78°5 1 ' 16", 900 m; humid subtropical. A settlement on the western slope of the Cordillera Occidental. Collection by Manuel Olalla in February 1958.38 M/Wo(Pichincha)— 00°02'54"S,78°46'21"W, 1410 m; humid subtropical. A village on the Pacific slope of the Cordillera Occidental; disturbed humid upper mon- tane forest. Collections by L. Ponce in April 1953, M. Olalla in May 1959, M. S. Foster at Hacienda San Vicente in February 1985, LAC in September 1992, and from adjacent Bosque Protector Mindo-Nambillo by A. Lopez in March 1994. Collections from disturbed forest along roads— 18 km N (1500 m) by JAP in June 1966; 3.5 km NE (1540 m) by WED in April 1975; 1 km E ( 1400 m) by JAP in June 1966. 46 Mr;//f'/»w(Azuay)— 02°45'15"S,79°23'48"W,2560 m; humid temperate. Village on the Pacific slope of the Cordillera Occidental. Collection by LAC and others in October 1993. 128 Moraspungo (Cotopaxi)— OriO'48" S, 79°12'03" W; humid tropical. A village near the base of the Andes. Collections from 9. 1 km E (800 m) and 1 1 km E (840 m) road to El Corazon by Charles W. Myers in November 1979 and RWM in October 1979, respectively. 94 Nanegal Chico (Pichincha) — See Nanegalito. Nanegal Grande (Pichincha)— 00°08'36" N, 78°40' 36" W, 1 800 m; humid temperate. A village on the Pacific slope of the Cordillera Occidental; disturbed humid upper montane forest. Collection by Manuel Ollala in August 1950.33 Nanegalito (Pichincha)— 00°04'00" N, 78°40'50" W, 1600 m; humid subtropical. A village on the Pacific slope of the Cordillera Occidental; disturbed humid upper montane forest. Collection by Gonzalo Herrera in March 1 957. Collections in disturbed forest along roads — 4 km N by C. R Walker in July 1962; 1.5 km SW, 5 km NW, and 10 km NW by JAP in November 1958. 37 Naranjal (Guayas)— 02°40'30" S, 79°36'30" W, 24 m; dry tropical. A town on the Pacific coastal plain; cacao groves. Collection by G. H. Tate in May 1922. 124 Naranjito (Guayas)— 02°09' S, 79°27' W, < 200 m; dry tropical. A town on the Pacific coastal plain. Collec- tion by Manuel Ollala in May 1964. 114A Mwo (Pichincha)— 00°03' 50" S. 78°34'54" W, 2730 m; humid temperate. A village on the Pacific slope of the Cordillera Occidental; disturbed humid upper montane forest. Collections along road NE to Tandayapa — 4.5 km NW (2600 m) by LAC in December 1989; 9.5 km NW (2530 m) by WED in April 1975; 13.1 kmNW(2100m) by KM in March 1979. Collections along road NW to Nanegal— 10 km NW (2200 m) and 12 km NW by JAP in October 1958 and KM in March 1979. 48 Otavalo (Imbabura)— 00°13' N. 78° 15' W, 2554 m, humid temperate. City in an inter-Andean basin. 29B Otongow (Pichincha)— 00° 15" 18" S, 78°4r48" W; humid subtemperate. Asite on a tributary of the Rio Saloya on the upper slope of the Cordillera Occidental S W of Lloa. Collection from humid montane forest on Quito-Chiriboga road, 3300 m by LAC in October, 1984. 57 Pachijal (Pichincha)— 00°09'05" N, 78°57'36" W, = 1200 m; humid subtropical. Settlement on Pacific slope of Cordillera Occidental near Rio Pachijal, a tributary of ELEUTHERODACTYLUS IN WESTERN ECUADOR 215 the Rio Guayllabamba; lower humid montane forest. Collection by G. Olalla in September 1959. 30 PacroCPichincha)— 00°08'36"N,78°45'36"W, 1350 m; humid subtropical. A village on the Quito-Rfo Guayllabamba road on the Pacific slopes of the Cordil- lera Occidental. Collections in the vicinity by Pablo Mena in October 1954 and Manuel Olalla in December 1955.32 Paitanga Chimborazo) — See Pallatanga. Pallamnga (Chimborazo)— 0r59"48" S, 78°5V48" W, 1 520 m; dry subtropical. A village on the Cajabamba- Guayaquil road in the valley of the Rio Coco on the Pacific slope of the Cordillera Occidental; collections from 3 km N (2280 m), 5 km N (2450 m),and 15 km N (2860 m) by JAP in February 1959, and in cutover montane forest 20 km NNE. 2550 m by WED in May 1975. 113 Palma Real (Pichincha)— 00^05"54" N, 79°28'54" W, = 500-700 m; humid subtropical. Settlement on Rio Guayllabamba on lower Pacific slope of Cordillera Oc- cidental (fide Paynter and Traylor, 1977). Collection by Manuel Olalla in June 1955. 35 Palo Quemado (Cotopaxi)— 00°38' S, 78°58' W, 1 800 m; humid temperate. Settlement, 8.2 km NE of San Francisco de las Pampas on western slope of the Cordil- lera Occidental; disturbed humid upper montane forest. Collection by G. Onore in February 1988. 83 Pambeldr (Esmeraldas) — See Pambilar. Pambildr (Esmeraldas)— 0r03'36" N, 78°48'22" W; humid tropical. Settlement on Pacific coastal plain 3 km NE of Concepcion. Collection by W. F. H. Rosenberg for BM. 4 Paramba (Imbabura) — See Parambas. Parambas (Imbabura)— 00°48' 18" N, 78°2r03" W, 780 m; humid subtropical. A hacienda on the south bank of the Rio Mira on the Pacific slope of the Cordillera Occidental; lower montane forest. Collection by Rosenberg for BM in October 1897. 20 Parque Nacional Machalilla (Manabi) — A park of about 55,000 ha on the Pacific coast (dry tropical) and extending inland to encompass parts of the Cordillera de la Costa (dry subtropical). Collections made at Bola del Oro on Cerro San Sebastian (0 1 °36' S, 80°42' W, 750 m) by Ana Almendariz in January 1991. 100 Parwquia Carondelet (Esmeraldas) — Or07'52" N, 78°45'34" W, 200 m; humid tropical. Village on Rio Bogota in lowland humid tropical rainforest. Collection by Charles M. Fugler in August 1963. 1 Patricia Pz/ar (Pichincha)— 00°34'S, 79°22' W, 180 m. Town on Pacific lowlands in southern Pichincha. Collections from Centinela, 14.1 km SE of town. See Centinela. Pasaje (El Oro)— 03° 19' 12" S, 79°48'12" W, 28 m; dry tropical. Town on the Pacific coastal plain; mostly cultivated tropical dry forest. Collections from 3 km E (30 m) and 55.4 km E (1000 m, Prov. Azuay) in tropical dry forest in the valley of the Rfo Jubones by A. C. Echternacht in June 1971. 133 Pedro Vicente Maldonado (Pichincha) — 00''05'18" N, 79°05'24" W, 600 m; humid subtropical. A village 3.5 km NW of Alvaro Perez Intriago on the Pacific slopes of the Cordillera Occidental. Collections were made in disturbed areas 1 km E (670 m) and 4 km S (Rfo Caoni. 570 m) by LAC in March 1990. 36 Pilalo (Cotopaxi)— 00°56' 1 5" S, 78°59'40" W, 2400 m; humid temperate. Village on the Pacific slope of the Cordillera Occidental; disturbed humid upper montane forest with many arboreal bromeliads. Collections by JDL in June 1968, June 1977, and January 1978; by WED in July 1971 and March 1984; by LAC in August 1 986 and May 1 989. Also from 1 km W (2340 m) by JDL in July 1970 and from 2 km W (2220 m) and 3 km W (2160 m) by JAP in January 1959. Collections from eastward along road to Pujili — 1 km E (2475 m) by JAP in January 1959, 2 km E (2480 m) by JAP in January 1959, 3 km E (2900 m) by JDL in July 1970, 4.6 km E (2600 m) by A. C. Echternacht in May 1971, 5 km E (2900 m) by JAP in January 1959, 6 km E (2670 m) by WED in July 1971, 8 km E 2850 m) by JAP in January 1 959. Collections in paramo from eastward along road to Pujili— 24.6 km E (3 190 m) and 27.6 km E (3380 m) by LAC in April 1 989. Collections in disturbed lower mon- tane forest along road to west to Quevedo — 18.6 km W (930 m) and 20.3 km W (830 m) by WED in July 1971, 29.6 km W (600 m) by R. R. Montanucci in May 1971. 89 Pinas (El Oro)— 03°40'43" S, 79°40'48" W, 1 0 1 0 m; dry subtropical. Town in the upper valley of the Rio Puyango at the southern base of the Cordillera de Chilla; disturbed tropical dry forest. Collections from town by G. H. Tate in September 1921, from Villa Elvita by H. Vargas in December 1 994, and from riparian habitat 4.4 km NW ( 1 1 00 m) and in disturbed humid montane forest at 18 km W (780 m) by WED in July 1971 and March 1975, respectively. Also from 13-16.4 km W (585-800 m) by RWM in October 1979. 137 Placer (Esmeraldas) — See El Placer. Playa Rica (Pichincha)— 00° 10' N, 78°40' W, about 1 200 m. Asite north of Nanegal Grande. Collection by D. Lombeida in October 1987. 29C Portovelo (El Oro)— 03°42'48" S, 79°36"5 1" W, 6 10 m; dry subtropical. Village 4 km south of Zaruma in upper valley of Rfo Puyango; tropical dry forest. Collec- tion from village by G. H. Tate in September 1921 and from along stream 32.6 km SSE by WED in July 1971. 138 216 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Ponr/;//-(Bolivar)— or 1 6'24" S. 79"()7"48" W. 1 760 m; humid subtropical. A hacienda on the western slopes of the Cordillera Occidental. 1 1 km SW Facundo Vela. Collection by R O. Simons in March 1899. 96 Puerto de Ila (Pichincha)— 00 33' S. 79 22' 'W. 200 m; humid tropical. Settlement on Rfo Baba on Pacific coastal plain about 40 km SSW of Santo Domingo de los Colorados; remnants of humid tropical rainforest. Col- lection by Pablo Mena in December 1954. 80 Puerto Quito (Pichincha)— 00°08"42" N, 79M6' 10" W, 280 m; humid tropical. Town on Pacific coastal plain; cutover humid tropical rainforest. Collection from 8 km ESE (530 m) on road to Quito by WED in April 1975. 31 Puerto Velo. (El Oro) — See Portovelo. Puli'm (Esmeraldas) — See San Francisco. Pungald (Chimborazo)— 01 °48'48" S, 78°35'22" W, 2880 m; dry temperate. A village 2 km SE of Licto (S of Riobamba). Collections by LAC from 4 km W, 2900 m, in January 1 990. and from 7 km NW (road to Riobamba), 2880 m, in August 1989. 110 Quebrada La Plata (Pichincha)— 00°27' S, 78°47' W, 1560 m; humid subtropical. A small tributary of the Rfo Pilaton, 3 km SETandapi. Collections by KM and RWM inFebaiary 1979.77 Quebrada Silante Grande (Pichincha) — See Campamento Silante. Quebrada Zapadores, 5 km ENE Chiriboga (Pichincha)— 00° 15' S, 78°43' W, 20 1 0 m humid temper- ate. Tributary of Rio Saloya, 5 km ESE Chiriboga; disturbed upper humid montane forest. Collections by WED in April and May 1975. JDL in July 1977. and RWM in March 1979.53 Quevedo (Los Ri'os)— Oror30" S, 79°27"39" W. 53 m; humid tropical. Town on Pacific coastal plain; small remnants of humid tropical rainforest, but mostly planta- tions of cacao and bananas. Collections from Quevedo by Gonzalo Herrera in December 1955; I km N, 300 ft. by JAP in August 1 954; 1 .6 km N (Finca Playa Grande) by George Key in March 1965 and June 1966; 4 km N, 140 m; by JDL in July 1970; 46 km N (Hacienda Cerro Chivo) by H. Weed in July 1992; 3 km E (road to EI Corazon) by KM in July 1975. 91 e"/>;/m/e (Esmeraldas)— 00° 1 8'50" N, 79°27"40" W. 40 m (= Rosa Zarate); humid tropical. Town on the Rfo Blanco on the Pacific lowlands. Formerly humid tropical rainforest; now mostly cultivated in bananas. Collection from disturbed forest 10.5 km N, 10 m by WED in July 1971.29 Reserva Floristica-Ecologica Rio Guajalito (Pichincha)— 00° 14' S. 78°48' W. 1800-2200 m; humid temperate. Anatural reserve NE of Palmeras(Chiriboga- Santo Domingo de los Colorados road). Collections by LAC and others in November 1 99 1 , December 1 993 , and October and November 1994. 52 Re.serva Otonga (Cotopaxi)— 00°44' S, 78°59' W, 2000-2200 m; humid temperate. A natural reserve near San Francisco de las Pampas. Collections by LAC in August 1993, Molineros in March 1994, and I. Tapia in November 1994.85 Rfo Ayanipe (Guayas) — See Ayampe. Rfo Baba (Pichincha) — Humid tropical. Southward flowing river on Pacific coastal plain; riparian rainforest along river. Collections made at 10 km S, 4 km E Santo Domingo de los Colorados (08° 1 8' S, 79° 1 3' W, 400 m) by WED in July 1 97 1 and April 1 972, and at 1 9 and 24 km S Santo Domingo de los Colorados (00°25' S, 79° 17' W) by George Key in Febaiary-March 1965 and April- June 1966.62 Rio Baboso (Carchi)— 00°53' N, 78°27' W; humid subtropical. A tributary of the Rfo Mira N of Lita. Collection from headwaters of Rfo Baboso in primary forest about 4—5 km N Lita, 900 m by LAC in September 1984. 14 Riobamba (Chimborazo)— 01 °40' S, 78°38' W, 2750 m dry temperate. City in the Andes. Collection from 70 km W (road to Pallatanga), 2520 m (humid temperate) by LAC in August 1989. 102 Rfo Blanco (Carchi)— 00°54'35" N, 78°22'24" W, =700 m; humid subtropical. A village on the Rfo Blanco, a tributary of the Rfo San Juan, on the lower slopes of the Cordillera Occidental. Collections from along the Rfo Blanco, 2 km NE, 930 m, by JAP in November 1 958. and from Chical, 1450-1500 m, 3 km up the Rfo Blanco by Alwyn. H. Gentry in September 1979. 11 Rfo Blanco (Pichincha)— 00°I7' 12" S, 78°42'57"; humid subtropical. A tributary of the Rfo Saloya on the western slope of the Cordillera Occidental. Collections from along the river by and near mouth of the Rfo Yambi by Manuel Olalla in May 1953 and February 1959, respectively. 61 Rfo Caoni (Esmeraldas) — Not located. Collection from "region of, sector de Largartera" by Charles M. Fugler in August 1963. Rfo Chimbo, near junction of Rfo del Oro (Guayas ) — Not located. Collection by I. R. Wiggins in December 1944. Rfo Coco (Chimborazo) — Canton Pallatanga, 0I°59"48" S, 78°57'48" W, 1450 m; humid subtropical. River on Pacific slope of Cordillera Occidental; cutover montane rainforest. Collection along river near Pallatanga by A. Almendariz in September 1986. 114 Rfo Cuaque (Manabf)— 00°02' S. 79°57' W, 100 m; dry tropical. Small river on western slope of Cerros de Cuaque E of Pedemales on road to El Carmen; tropical dry forest. Collection by LAC in December 1989. 47 Rfo Cupa (Esmeraldas)— 00°26' N, 79°24' W, < 1 50 m ELEUTHERODACTYLUS IN WESTERN ECUADOR 217 (mouth); humid tropical. Small tributary of the Rfo Blancoon the Pacific lowlands. Humid tropical rainforest. Collection by A. Proaiio in January 1952. 25 Rio Ditrango (Esmeraldas)— OTOS' N, 78°4r W. <150 m; humid tropical. A tributary of the Rfo Bogota. Lowland tropical rainforest. Collection by W. P. H. Rosenberg for BM. 3 Rio Faisanes (Pichincha)— 00° 1 9' S. 78°49' W, 1 380 m; humid subtropical. Stream in humid lower montane forest, 13.5 km from La Palma on "old" road to Quito. Collections by G. O. Vigle in January 1978 and KM and RWM in January and October 1979. 66 Rio Frio (Quay as) — Not located. Collection by T. H. Hubbell in April 1963. Rio Giiichichi (Chimborazo)— Or57' S, 78°56' W, 1700 m; dry subtropical. In Canton Pallatanga. Collec- tions along banks of river in cutover forest by A. Almendariz in September 1986. 112 Rio Lelia (Pichincha)— 00° 1 8' S. 79°0r W, 1000 m; humid subtropical. A tributary of the RioToachi. Collec- tion by Manuel Olalla in April 1959. 64 Rio Lita (Esmeraldas) — Small river draining Pacific slope of Cordillera Occidental. 2 km S junction Rfo Lita and Rfo Mira. 00°56' N, 78°32' W, 520 m (humid sub- tropical). Collection by JAP in November 1958. 8 Rio Minas, 20 km W of Santa Isabel (Azuay) — 03° 18' 18" S, 79°22'24" W, 1250 m; dry subtropical. Collection by JAP in 1959. 132 Rio Parambas (Imbabura)— 00°01' S, 78° 14' W; hu- mid subtropical. Collection from edges of river where it crosses the Ibarra-Lita road by R. Barriga in August 1984. 44 Rio Pescado (Chimborazo) — Collection by G. H. Tate in May 1922. Not located. Rio Pitsard (Pichincha)— 00° I4'36" N, 79°06'48" W. Tributary of Rfo Guayllamba, SE of Golondrina. Collection by Manuel Olalla in July 1953. 29A Rio San Miguel (Esmeraldas) — River in Pacific low- lands. Collection from about 1 km above Rfo Cayapas, 00°5 r N, 78°57' W, < 1 00 m (humid tropical). Collection by KM in June 1977. 17 Rio Saloya (Pichincha) — Tributary of Rfo Blanco draining western slope of Volcan Pichincha. Elevation of 450' of specimens in BM must refer to Rfo Blanco (humid tropical). Rio Sapayo (Pichincha) — See Rfo Saloya. Rio Tatalmazo (Lizo) (Bolfvar)— 01 °43' S, 78°59' W, 25 10 m; dry temperate. A river in the Bosque Protector Cashca Totoras south of Santiago. Collection from near Cruz de Lizo by LAC in January 1990. 107 Rio Toachi (Pichincha)— 00°23' S, 78°56' W, 800 m; humid subtropical. River on Pacific slope of Cordillera Occidental; remnants of humid lowland rainforest. Col- lection from gallery forest along river 8 km E Alluriquin by WED in July 1971.70 Salidem (Esmeraldas)— 00 55' N. 78'34' W. 108 m; humid tropical. Village in Andean foothills. Collection by W. R H. Rosenberg in April-August 1919. 9 Salto dos Novias (Pichincha)— 00°26' S, 78°4r W, 2635 m; humid temperate. Waterfall on N slope of Cerro Corazon on road from Aloag to Santo Domingo de los Colorados; remnants of upper humid montane forest. Collection by WED in March 1967. 74 San Francisco (Esmeraldas)— Or05' N, 78°42' W, <150 m; humid tropical. Settlement on Rfo Bogota in disturbed humid tropical rainforest. Formerly known as Pulun. 2 San Francisco de Las Pampas (Cotopaxi ) — 00°40' S, 78°53' W, 1800 m; humid subtropical. A village on the Pacific slopes of the Cordillera Occidental. Collections from humid montane forest in the vicinity of the village at elevations of 1 600-2200 m by G. Onore and members of the Tapia family in November 1984, October 1985, March, May, and August 1986, December 1989, Febru- ary 1990, October 1991, and November 1992, and by LAC in August 1986 and August 1993. 84 San /g/;ar/V; (Pichincha)— = 00°27' S, 78°42' W, 2030 m; humid temperate. Settlement 10 km E of Tandapi on road to Aloag; upper humid montane forest. Collections from immediate vicinity by WED in March 1967 and from 1 km SW, 1920 m, by JDL in July 1977. 76 San Javier de Cachabe (Esmeraldas) — 00°58' N, 78°48' W, 20 m; humid tropical. Village on Rfo Cachabf and on Ibarra-San Lorenzo railroad; cut over humid tropical rainforest. Also spelled Cachabf and Cachavf. Collections by W. F. H. Rosenberg in April-August 1919, Charles M. Fugler in September 1 963, and from 1 km SW by L. Abuja and JAP in December 1958. 7 San Josede Bilsa (Esmeraldas )— 00°37' N. 79°5 1 ' W; humid tropical. A settlement on the western slope of the Montafias de Muise (northernmost part of the Cordillera de la Costa). Collections made in the vicinity of a camp in lowland tropical rainforest at an elevation of 225 m, 7 km E of the village by Ana Almendariz in January 1 99 1 . 24 San Juan (Pichincha)— 00° 17' S, 78°35' W, 3400 m; humid temperate. Settlement near crest of Quito- Chiriboga road south of Volcan Pichincha. Collection in disturbed high cloud forest by JDL in July 1977. 61 A Scm Miguel, 4 km del Campomento del MAG — See San Miguel de Cayapas. San Miguel de Cayapas (Esmeraldas) — 00°45' N. 78°54' W, <200 m; humid tropical. Village at the junction of the Rfo San Miguel and Rfo Cayapas. Disturbed humid tropical rainforest. Collections by KM in June 1977 and A. Almendariz in September 1984. 22 218 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 San Miguel de Congomd (Pichincha) — See San Miguel de los Colorados San Miguel de los Colorados {Pichincha) — 00°20' S, 79°15W, 500 m; humid subtropical. Settlement, also known as San Miguel de Congoma, south of Santo Domingo de los Colorados on road to Quevedo. Now mostly cultivated in bananas. Collections by Antonio Proatio in November 1953 and by Manuel Olalla in December 1956. 67 San Miguel-Telembf{Esmera\das)—00°45' N, 78°55' W, <200 m (Telembi); humid tropical. Rio Cayapas between San Miguel and Telembi; humid tropical rainforest. A. Almendariz; September 1984. 21 San Tadeo (Pichincha)— 00°0 1 ' N, 78°45' W, 1 500 m; humid subtropical. A site near Mindo. Collection by G. Olalla in August 1959.39 Santo Domingo de los Colorados ( Pichincha) — 00° 1 5' S. 79° 10' W, 600 m; humid subtropical. Town near base of the Andes on the Pacific coastal plain; remnants of lowland tropical rainforest but mostly cutover and now in banana plantations. Collections by C. Olalla in May 1929, by E, H, Taylor in 1962, by WED in March 1967 and July 1 97 1 ; by JDL in March, July, and August 1 968 and June 1977. Collections also from 1 km N, 2 km E, 620 m; JDL, December 1977; 2 km and 6 km E by P D. Spoecker in June 1962; 3.5 km E (Hacienda Lelia) by F. H. Funkhouser in September 1950; 5 km E by George Key in February and August 1965; 8 km SE by C. F. Walker in August 1962; 0.5 km S by JAP in May 1959; 5 km W by Cesar Duran in December 1965; 6 km W by E. S. Ross in February 1 955; 9 km W ( Hacienda Espinosa) by J. W. Funkhouser in September-November 1950; 18 km W on road to Chone by JAP in May 1959. 56 San Vicente (Bolivar)— 0r43' S, 79°01' W; humid temperate. Collection by LAC in December 1984. 106 Sigchos (Cotopaxi)— 00°42' S, 78°53' W, = 3000 m; humid subtemperate. Village in the Cordillera Occiden- tal; subparamo. Collection from "region of Sigchos" by Manuel Olalla in June 1957, and from "below Sigchos" (Prov. Pichincha) by Antonio Proano in October 1954. 84a Sitio Torneado (Bolivar) — See El Tomeado. Tandapi (Pichincha)— 00°24'52" S, 78°47'48" W, 1460 m; humid subtropical. Small village (also named Manuel Comejo Astorga) on south bank of Rio Pilaton on road from Aloag to Santo Domingo de los Colorados; disturbed upper humid montane forest. Collections by WED in March 1967; JDL in July and August 1967, March and August 1968, August 1970, and June and December 1977; and by Thomas H. Fritts in August 1970. Also collections along road to east and west — 2.1 km E ( 1500 m) and 6.2 km E ( 1750 m)by KM and RWM inFebruary 1979, and 1.6 km W( 1400 m) by JDL in June 1977.72 Tandayapa (Pichincha)— 00°01' N, 78°39' W, 1830 m; humid temperate. A village on the road from Quito to Puerto Quito; upper humid montane forest with many bromeliads. Collections in vicinity of town by Leonides Ponce in March 1952, JAP in March 1953, and Mauro Proafio in April 1956. Also collections from 2 km E by JAP in October and November 1 958, and 9 km SE (2 1 50 m) by WED in April 1975; also from various distances S W on road to Puerto Quito— 1 .4 km ( 1 820 m) by WED in April 1 984, 2 km ( 1 840 m) by LAC in September 1 993, 2.3 km (1900m) by WED in April 1984,4-5 km by LAC in September 1993, 7 km (1900 m) by KM and RWM in February 1979, and 9.4 km (2390 m) by WED in April 1975 and April 1984.41 Tinalandia (Pichincha)— 00° 18' S, 79°04' W, 700 m; humid subtropical. A hotel and park 15.5 km SE Santo Domingo de los Colorados; disturbed lowland tropical rainforest. Collections by KM in July- August 1972, July-August, 1975, and February 1979, E.E.Williams in August 1974, and by WED in March 1984. 63 Toacaso (Cotopaxi)— 00°45' S, 78°4r W, 2990 m; humid temperate. Small village SW of San Juan de Pastocalle. Collection by L. Lopez in October 1 989. 84B Tulcdn (Carchi )— 00°49' N, 77°44' W, 2960 m; humid subtemperate. City in the Andes in the upper valley of the Rio Carchi (a tributary of the Rfo San Juan). Collection from paramo 5 1 .3 km W on road to Maldonado, 3 1 50 m, by LAC in June 1989. 19 Urbina (Esmeraldas)— 00°58' N, 78°41' W, 300 m; humid tropical. Village on Rio Cachabi near base of Andes. 6 Vicente Maldonado (Vxchmcha) — See Pedro Vicente Maldonado. Victoria del Portete (Azuay)— 03°03' S, 79°03' W; humid temperate. Collection from 1 0 km S W, 2700 m by JDL in July 1970.129 Yerbabuena (Azuay)— 02 °45' S, 79°23' W, 2730 m; humid temperate. Collection by LAC and others in November 1992. 127 Zhud (Caiiar)— 02°28' S, 79°00' W, 2980 m; humid temperate. Small village on Panamerican Highway in upper valley of Rio Canar; disturbed subparamo. Collec- tion from 4 km N (3040 m) by WED in March 1984. 120 ELEUTHERODACTYLUS IN WESTERN ECUADOR 219 APPENDIX III Species of Eleutherodactylvs Names applied in the genus Eleiitherodactyhis with subgeneric assignments, species-group assignments, the state of the lengths of Toes III and V (description of conditions in text), and nomenclatural or taxonomic status (Comments). Series are employed (listed in parentheses) only for the subgenus Eleutherodactylus. (See text.) Species-group assignments are incomplete because not all taxa are regularly assigned to groups (e.g.. Hedges, 1989). In the case of West Indian taxa, we introduce a novel species group {E. abbotti group) to accomodate some taxa we believe wrongly assigned to Euhyas by Hedges (1989); those taxa are here assigned to the subgenus Eleutherodactylus. Additionally, we elect to recognize only a martinicensis group (of the auriculatus series) in the West Indies, combining much of what Hedges treated as the martinicensis and montanus groups, because Hedges did not assign many of the species of his auriculatus series to species groups. The subgeneric assignments used here include many novel ones and do not necessarily agree with the assignments made on the basis of the literature (e.g., Duellman, 1993). This is especially true for the subgenus Euhyas. Abbreviations for subgenera are: CR = Craugastor, EL = Eleutherodactylus, EU = Euyhas, PE = Pelorius, SY = Syrrhopus. Within the subgenus Eleutherodactylus, we employ several series abbreviated as follows: (a) abbotti, (b) binotatus, (c) conspicillatus, (m) martinicensis, (s) sulcatus, and (t) myersi. Toe conditions are: A = fifth toe shorter than third, B = fifth toe longer than third but not extending to distal subarticular tubercle of fourth toe, C = fifth toe much longer than third. Species Trivial name, author, date Subgenus group Toe condition Comments aaptus Lynch & Lescure 1980 abbotti Cochran 1932 acatallelus Lynch & Ruiz 1983 acerus Lynch & Duellman 1980 achatinus Bou\tr\gtr \%9% acmonis Schwartz 1960 actites Lynch 1979 acuminatus Shreve 1935 acutirostris Lynch 1 984 adamastiis Caxx\\)bt\\ 1994 affini sterner 1899 alalocophus Roa & Ruiz 1991 alberchi ¥\oxfi, 1988 albipes Barbour & Shreve 1937 albolabris Taylor 1940 albolabris Tay\or 1943 alcoae Schwartz 1971 a//r6'^/ Boulenger 1898 altae Dunn 1942 altamazonicus Barbour & Dunn 1921 alticola Lynn 1937 amadeus Hedges 1987 amelasma Schwartz 1958 amianthus Schwartz & Fowler 1973 amplinympha Kaiser, Green & Schmidt 1994 amulae Giinther 1900 anae Rivero 1986 anatipes Lynch & Myers 1 983 EL(m) unistrigatiis C EL(a) abbotti B EL(c) devillei B EL(m) unistrigatiis C EL(c) conspicillatus B EU ricordii A EL(c) conspicillatus B EL(m) unistrigatus C EL(c) - B CR milesi A EL(m) unistrigatus C EL(m) unistrigatus C - - - = surd us EU ricordii A - - - = pygmaeus - - - = dixoni EU - A CR alfredi B EL(m) unistrigatus C EL(m) unistrigatus C EU nubicola A EL(a) abbotti B EU ricordii A EU gossei A EL(m) martinicensis C - - - - nitidus - - - = curtipes CR anomalus A 220 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Appendix III. Continued. Species Trivial name, autlior. date Subgenus group Toe condition Comments anciano Savage et al 1 988 anderssoni hynch 1968 audi Savage 1 974 andicola Boettger 1 892 andrewsihynn 1937 angeliciis Savage 1975 angustidigitonim Taylor 1940 anolirex hynch. 1973 cDwmahis Boulenger 1898 anonynuis A. Lutz 1927 anotis Walkeri & Test 1955 cmtillensis Reinhardt & Liitken 1 863 anzuetoi Stuart 1 94 1 apantheatus Schwartz 1965 aphumis Campbell 1994 apicidatus Lynch & Burrowes 1990 aporostegiis Schwartz 1965 apostates Schwartz 1973 appendiculatus Werner 1 894 areolatus Boulenger 1898 armstrongi Noble & Hassler 1933 atkinsi Dunn 1 925 atratus Lynch 1979 audanti Cochxan 1934 augusti Duges 1 879 auriculatoides Nob\c 1923 auriciilatiis Cope 1862 aiihlegus Savage et al 1988 avocalis Taylor & Smith 1945 azuemensis Savage 1975 babax Lync\\ 1989 bacchus Lynch 1984 bakeri Coc\\xiix\. 1935 balionotus Lynch 1979 barbouri ^'leden 1923 barbiidensis Auffenberg 1958 barlagnei Lynch 1965 bartonsmithi Schwartz 1960 baryecuus Lynch 1979 batrachylus Taylor 1 940 bearsei DueWmdiXX 1992 beatae Boulenger 1903 beehei Coc\\vdX\ 1956 bellona Lynch 1992 berkenbuschii Peters 1 870 bemali Lynch 1 986 bicolor Rueda & Lynch 1983 bicumidus Peters 1 863 CR CR rugulosus fitzingeri ockendeni uncertain relationships EU nubicola A CR ntgulosus A SY nitidiis A EL(m) iiiiistrigatus C CR anomalus A EL - - = nomen nud EL(m) unistrigatus C EL(m) nuirtiniceiisis C - - - = lineatus EU I i cord a A CR biporcatiis A EL{m) unistrigatus C PE ruthae B EU ricordii A EL(c) devillei B - - - = chalceus EU ricordii A EU ricordii A EL(m) unistrigatus C EL(a) abbotti B CR augusti A EL(m) martinicensis C EL( m ) martinicensis C CR rugulosus A - - - = rugulosus CR rugulosus A EL(c) dolops B EL(m) unistrigatus C EL(m) C EL(m) unistrigatus C - - - - gundlachi - - - = johnstonei EL(m) martinicensis C EL(m) nuirtiniceiisis c EL(m) unistrigatus c EL(m) - c EL(m) unistrigatus c - - - = rhodopis - - - = inoptatus EL(m) unistrigatus c CR rugulosus A EL(m) unistrigatus c EL(m) unistrigatus c EL(m) - c ELEUTHERODACTYLUS IN WESTERN ECUADOR 221 Appendix III. Continued. Species Trivial name, author, date Toe Subgenus group condition Comments hilineatus Botcermann 1974 binghami Stejneger 1913 marsupiata binotatus Spix 1824 bipoiratus Peters 1 863 hisignatus Werner 1 899 blai h ^Mhoux 1928 bockermanni Donoso-Barros 1970 boconoensis Rivero & Mayorga 1973 bocourti 'Qrocc\\\ 1877 bogotensis Peters 1863 bolbodacnlus Lutz 1 925 ^o//v(7/7 Taylor 1942 bothroboans Schwartz 1965 boulengeri hynch 1981 brachypodius Rivero 1961 bransford a Cope 1886 brederi Dunn 1934 bresslerae Schwartz 1960 brevicrus Andersson 1945 brevifrons hynch 1981 brevipabnatiis Schmidt 1920 brevirostris Shxeve 1936 briceni Boulenger 1903 brittoni Schmidt 1920 brocchi Boulenger 1 882 bromeliaceiis Lynch 1979 buccinator Rodriguez 1 994 buckleyi Boulenger 1882 buergeri Werner 1 899 bufonifonnis Boulenger 1896 /7m/oaj/m5 Andersson 1945 bufonoides Lynch 1965 cabrerai Cochran & Coin 1970 cacau Lynch 1992 cactonim Taylor 1939 cadenai Lync\\ 1986 cajamarcensis Barbour & Noble 1920 calcaratus Boulenger 1908 ca/cY/ram.9 Andersson 1945 calcarulotus hynch 1976 calcitrans Gunther 1901 caliginosus Lynch 1996 campi Stejneger 1915 cantitans Myers & Donnelly 1996 caprifer hynch 1977 caribe Hedges & Thomas 1992 carmelitae Ruthven 1922 EL - - - = Gastrotheca EL(b) binotatus A CR bipoiratus A - - - = fene stratus - - - = podiciferus EL(m) unistrigatus C EL(c) - B CR alfredi B EL(m) unistrigatus C EL(b) lacteus B - - - = augusti PE ruthae B EL(m) unistrigatus C - - - = vilarsi CR rhodopis A - - - = achatinus EU ricordii A - - - = altamazonicus EL(m) unistrigatus C - - - = sierramaestrae EU ricordii A EL(c) - B EL(m) martinicensis C CR rugulosus A EL(m) unistrigatus C EL(c) conspicillatus B EL(c) curtipes B - - - = w-nigruni CR bufoniforniis A - - - = diadematus - - - = spatulatus EL(m) unistrigatus C EL(m) unistrigatus C - - - = augusti EL(s) sulcatus A EL(m) unistrigatus C EL(m) unistrigatus C - - - = anderssoni EL(m) unistrigatus c - - - - omiltemanus EL(c) conspicillatus B SY leprus A EL(m) unistrigatus C EL(c) conspicillatus B EU - A EL(c) conspicillatus B 222 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Appendix III. Continued. Species Trivial name, author, date carranguerorum Lynch 1994 carrioni Parker 1932 carvalhoi Lutz 1952 caryophyllaceus Barbour 1928 casparii Dunn 1926 cavernicola Lynn 1954 celator Lynch 1976 ceraemerus Schwartz 1968 cerasinus Cope 1 876 cerasoventris Rivero 1984 cerastes hynch 1975 ceuthospilus Duellman & Wild 1993 t//ar Savage 1987 chalceus Peters 1 873 cheiwplethus Lynch 1990 chersonesodes Schwartz 1965 chiastonotus Lynch & Hoomoed 1977 chica Noble 1918 chimboe Fowler 1913 chiquito Lynch 1965 chloronotus Lyv\c\\ 1970 chlowphenax Sc\\wanz 1976 chlorosoma Rivero 1984 cholorum Nt\\\ 1965 chrysozetetes McCranie et al 1989 cinereus Cope 1885 citriogaster DueWman 1992 cochranae Grant 1932 coerM/eM5 Andersson 1945 colodactylus Lynch 1979 colomai Lynch & Duellman 1996 colostichos LaMarca & Smith 1982 CO/7 Jor Lynch & Duellman 1980 conspicillatus Giinther 1 859 conspicuus Taylor & Smith 1945 cooki Gram 1932 cogw/ Thomas 1965 cornutus Jimenez de la Espada 1870 corona Hedges & Thomas 1992 cosnipatae Duellman 1978 costaricensis TayXox 1952 coiinouspeus Schwartz 1964 cramptoni Schmidt 1920 crassidigitus TayXov 1952 cremnobates Lynch & Duellman 1980 crenunguis \j)/x\c\\ 1976 crepitans ^oktrmann 1965 cristinae Lynch & Ruiz 1985 Subgenus group Toe condition Comments EL(c) conspicillatus B - - - = lymani EL(m) unistrigatus C EL(m) unistrigatus C EU ricordii A EU cundalli A EL(m) unistrigatus C EL(m) martinicensis C EL(c) cerasinus B — - - = vanadise EL(s) sulcatus A EL(m) unistrigatus C CR gollmeri A EL(m) diastema C CR anomalus A EU ricordii A EL(c) conspicillatus B - - - = diastema - - - = curtipes - - - = greggi EL(m) unistrigatus C PE inoptatus B EL(m) tubernasus C SY leprus A CR mile si A - - - - fene stratus EL(c) conspicillatus B EL(m) martinicensis C - - - = unistrigatus EL(m) unistrigatus C EL(m) unistrigatus c EL(m) unistrigatus c EL(c) conspicillatus B EL(c) conspicillatus B CR alfredi B EL(m) martinicensis C EL(m) martinicensis C EL(s) sulcatus A EL(a) abbotti B EL(c) - B - - - = bransfordii EU ricordii A — - - - locustus CR fitzingeri A EL(m) unistrigatus C EL(c) cerasinus B - - - - fene stratus EL(c) ELEUTHERODACTYLUS IN WESTERN ECUADOR 223 Appendix III. Continued. Species Toe Trivial name, author, date Subgenus group condition Comments croceoinguinis Lynch 1 968 EL(m) unistrigatus C crucifer Boulenger 1 899 EL(m) unistrigatus C cruentus Peters 1 863 EL(m) unistrigatus C crMra/z5 Boulenger 1902 EL(c) discoidalis A cruizi McCranie et al 1 989 CR milesi A cryophilius hynch 1979 EL(c) curtipes B cryptomelas Lynch 1 979 EL(m) unistrigatus C CMa^M^ra Savage 1980 CR fitzingeri A cubanus Barbour 1942 EU - A cunctator Dnnn 1928 - - - = orcutti cundalUDunn 1926 EU cundalli A cuneatus Cope 1 862 EU ricordii A curtipes Boulenger 1882 EL(c) curtipes B cystignathoides Cope 1 877 SY leprus A Ja«ae Duellman 1978 EL{c) - B darlingtoni Cochran 1935 EU ricordii A darlingtoni Cochran 1935 - - - = jugans Jarv/Ford& Savage 1984 CR gollmeri A decoratus Taylor 1942 CR alfredi B degener Lynch & Duellman 1996 EL(m) unistrigatus C delacruzi Estrada et al 1986 - - - = symingtoni delicatus Ruthven 1917 EL(c) galdi A delius Duellman & Mendelson 1995 EL(m) unistrigatus C dennisi Lynch 1970 SY longipes A Jev///^/ Boulenger 1880 EL(c) devillei B diadematus Jimenez de la Espada 875 EL(m) unistrigatus C diaphonus Lynch 1986 EL(m) unistrigatus C diastema Cope 1 876 EL(m) diastema C dilatus Davis & Dixon 1955 SY nitidus A dimidiatus Cope 1 862 EU ricordii A diplasius Schwartz 1973 EL(m) martinicensis C discoidalis Peracca 1 895 EL(c) discoidalis A dissimulatus Lynch & Duellman 1996 EL(m) unistrigatus C J/v/j^a Wandolleck 1907 - - - = guentherii dixoni Lynch 1991 SY nitidus A dolomedes Hedges & Thomas 1992 EU - A dolops Lynch & Duellman 1980 EL(c) dolops A dorsoconcolor Taylor 1 94 1 - - - = rhodopis dorsopictus Rivero & Serna 1987 EL(m) unistrigatus C douglasi Lynch 1 996 EL(c) galdi B dubitus Taylor 1952 - - - = cruentus duellmani Lynch 1980 EL(c) surdus B dunnii Barbour 1922 - - - = rhodopis eileenae Dunn 1926 EL(m) martinicensis C elassodiscus Lynch 1973 EL(c) nigrovittatus A elegans Peters 1 863 EL(m) unistrigatus C emcelae Lynch 1985 CR fitzingeri A emiliae Dunn 1926 EU ricordii A 224 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Appendix III. Continued. Species Trivial name, author, date emmetti Gorham 1966 emleni Dunn 1932 eneidae Rivero 1959 engytympanum Giinther 1900 epipedus Heyer 1984 eqiuitohalis Barbour 1908 eremitus Lsnch 1980 eremus Schwartz 1965 eriphus Lynch & Duellman 1980 ernesti Flores 1987 en'thromenis Heyer 1984 enthropleitra Boulenger 1 896 erythroproctus Schwartz 1960 escoces Savage 1975 estradai Lynch 1991 etheridgei Schwartz 1958 eugeniae Lynch & Duellman 1996 eimaster Schwarlz 1973 euphronides Schwartz 1969 eurydactylus Hedges & Schluter 1992 euryglossus Cope 1 894 fassilianus Werner 1916 femurlaevis Cochran 1935 fenestratus Steindachner 1 864 festae Peracca 1904 festae Peracca 1904 fitzingeh O. Schmidt 1857 flavescens Noble 1923 flavomaculatus Parker 1938 flavomaculatus fleischmanni Boettger 1892 floridus Lynch & Duellman 1996 florulentus Cope 1 893 footei Stejneger 1913 fowled Schwartz 1973 frater Werner 1 899 fraudator Lynch & McDiarmid 1987 fuhnnanni Peracca 1914 furcyensis Shreve & Williams 1963 fuscatus Davis & Dixon 1957 fuscofemora Zweifel 1956 fiiscus Lynn & Dent 1943 fusciis Davis & Dixon 1955 gaigeae Schmidt & Smith 1944 gaigei Dunn 1931 galdi Jimenez de la Espada 1 870 ganonotus Duellman & Lynch 1988 gehnnanni Schwartz 1958 Subgenus group Toe condition Comments _ — - = emleni - - - = rugulosus EL(m) martmicensis C - - - = fleischmanni EL(b) binotatus A - - - = unistrigatus EL(m) unistrigatiis C EU ricordii A EL(m) unistrigatiis C EL(c) devillei B EL(b) binotatus A EL(c) devillei B EU ricordii A CR rugulosus A EU ricordii A EU ricordii A EL(m) unistrigatus C EL(a) abbotti B EL(m) martinicensis C EL(m) unistrigatus C - - - = fleischmanni - - - = bogotensis _ — - = oxyrhyncus EL(c) conspicillatus B - - - - galdi — - - = trepidotus CR fitzingeri A EL(m) martinicensis C - - - = Phrynopus CR rugulosus A EL(t) myersi B — - - - biporcatus - - - - Gastrotheca marsupiata EL(m) martinicensis C EL{m) unistrigatus C EL - A — - - = elegans EU ricordii A - - - = mexicanus CR augusti A EU gossei A _ - - = maiiriis - - - = petrophilus EL(c) conspicillatus B EL(c) galdi B EL(c) - B = eileenae ELEUTHERODACTYLUS IN WESTERN ECUADOR 225 Appendix III. Continued. Species Trivial name, author, date Subgenus Toe group condition Comments g^/irr/ Miranda-Ribeiro 1926 gentryi Lynch & Duellman 1996 ginesi Rivero 1964 gladiator Lynch 1977 glandu lifer Cochran 1935 glanduliferoides Shreve 1936 glandulosus Boulenger 1880 glaphycompiis Schwartz 1973 glaucoreius Schwartz & Fowler 1973 glaucus hynch 1967 goini Schwartz 1960 goldmani Noble 1924 goUmeri Peters 1 863 gossei Dunn 1926 grabhami Dunn 1926 gracilis Lynch 1986 grahami Schwartz 1979 gratjdiceps hynch 1984 grandis Dixon 1957 grandoculis Van Lidth de Jeude 1904 granulosus Boulenger 1903 gravenhorstii Fitzinger 1 867 greggi Bumzahem 1955 greyi Dunn 1 926 griphus Cromb'\e 1986 ^m^a Hallowell 1861 gryllus Schmidt 1920 gualteri Lutz 1 974 guanahacahihes Estrada & Rodriguez guantanamero Hedges et al 1992 guayanensis Rivero 1968 guentherii Steindachner 1867 guentherii¥.Qitx%\.t\n 1868 guerreroensis Lynch 1 967 gw/om Boulenger 1898 gundlachi Schmidt 1920 guttilatus Cope 1 879 gutturalis Hoogmoed et al 1977 hahenatiis Cope 1 876 haitianus Barbour 1942 hamiotae Flores 1994 hectus Lynch & Burrowes 1990 hedricki R'wero 1963 helonotus Lynch 1975 heminota Shreve & Williams 1963 henselii Peters 1 870 hernandezi Lynch & Ruiz 1983 heterodactylus Miranda-Ribeiro 1937 1985 - - - = bolbodactylus EL(c) curtipes B EL - A EL(t) myersi B EU ricordii A EL(a) ahbotti B EL(m) unistrigatus C EU ricordii A EU cundalli A CR alfredi B EU ricordii A - - - = gollmeri CR gollmeri A EU gossei A EU luteolus A EL(m) unistrigatus C EU ricordii A EL(m) unistrigatus C SY nitidus A EL(m) unistrigatus C - - - = crural is - - - = nomen nudum CR gollmeri A EU ricordii A EU nubicola A - - - = fitzingeri EL(m) martinicensis C EL(b) binotatus B EU ricordii A EL(m) martinicensis C - - - = nomen nudum EL(b) binotatus A - - - = fitzingeri CR alfredi B EL(m) diastema C EU ricordii A SY marnockii A EL(c) conspicillatus B - - - = podiciferus EL(a) abbotti B EL(c) surd us B EL(t) myersi B EL(m) martinicensis C EL(s) sulcatus A EU - A - - - = guentherii EL(m) unistrigatus C EL(b) binotatus A 226 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Appendix III. Continued. Species Toe Trivial name, author, date Subgenus group condition Comments hidalgoensis Tay\ov 1942 — - = = decoratus hoharrsmithi Tdy\oT 1937 CR rhodopis A hoelmei Lulz 1958 EL(b) hinotatus B /loW Cochran 1948 EL(b) lacteus B huigrae Fowler 1913 - - - - gularis humeralis ¥ow\cT 19 1 6 - - - - gollmeri hyhotragus Lynch 1992 hylaeformis Cope 1 875 hylaeformis Melin 1941 hypostenor Sc\\v^SLX\z 1965 ignicolor Lynch & Duellman ileamazonicus Rivero 1961 illotus Lynch & Duellman 1996 imitatrix DueWman 1978 incanus Lynch & Duellman 1980 incomptus Lynch & Duellman 1980 ineptus Barbour 1928 ingeri Cochran & Coin 1961 inguinalis Parker 1940 inoptatus Barbour 1914 insertus Lutz 1927 insignitus R\i\h\itn 1917 intermedius Barbour & Shreve 1937 interme dius Cochxdin 1941 interorbitalis Langebartel & Shannon 1 956 inusitatus Lynch & Duellman 1980 ionthus Schwartz 1960 izecksohni Caramaschi & Kisteumacher 1988 jaimei Lynch 1 992 jamaicensis Barbour 1910 jasperi Drewy & Jones 1976 johannesdei Rivero & Serna 1987 johnstonei Barbour 1914 johnwrighti Lynch 1996 jorgevelosai Lynch 1994 jota Lynch 1980 jugans Cochran 1937 juipoca Sazaima & Cardoso 1978 junori Dunn 1926 karlschmidti Grant 1931 katoptroides Flores 1988 kirklandi Flores 1985 klinikowskii Schwartz 1959 koki UeWn 1941 labiosus Lynch et al. 1994 lacrimosus Jimenez de la Espada 1875 lactea Miranda-Ribeiro 1923 laevissimus Werner 1 896 EL(c) loustes B EL(m) diastema C PE ruthae B EL(m) unistrigatus C EL(c) conspicillatus B EL(m) unistrigatus C EL(m) unistrigatus C EL(m) unistrigatus C = melini = vilarsi _ - - = diastema EL(s) sulcatus A EL(m) unistrigatus C PE inoptatus B EL - - = nomen dudium EL(c) conspicillatus B EU ricordii A — - - = haitianus SY modestus A EL(m) unistrigatus C EL(m) martinicensis C EL(b) binotatus A EL(c) loustes B EU jamaicensis A EL(m) martinicensis C EL(c) conspicillatus B EL(m) martinicensis C EL(m) unistrigatus C EL(m) unistrigatus C CR rhodopis A EU ricordii A EL(b) binotatus A EU gossei A EU - A EL(m) unistrigatus C EL(m) unistrigatus C = incomptus EU ricordii A - - - = sulcatus EL(c) cerasinus B EL(m) unistrigatus C EL(b) lacteus B = rugulosus ELEUTHERODACTYLUS IN WESTERN ECUADOR 227 Appendix III. Continued. Species Toe Trivial name, author, date Subgenus group condition Comments lamprotes Scliwartz 1973 EL(m) martinicensis C lane iform is Cope 1877 - - - = gollmeri /ancm// Donoso-Barros 1965 EL(c) - B lanthanites Lynch 1975 EL(c) conspicillatus B lasalleorum Lynch 1995 EL(m) leptolophus C latensljynch 1989 EL(c) nigrovittatus A laticeps D\xm€n\ 1853 CR gollmeri A laticlavius Lynch & Burrowes 1990 EL(m) unistrigatus C latidiscus Boulenger 1898 EL(m) unistrigatus C latodactylus Taylor 1940 - - - = longipes latrans Cope 1880 CR augusti A lauraster Savage et al. CR rhodopis A leberi Sch-wsLXiz 1965 EL(m) martinicensis C lehmani Boettger 1892 - - - = unistrigatus lehmanvalenciae T\\orion 1965 - - - = moro lentiginosus Rivero 1984 EL(c) - B lentus Cope 1 862 EU ricordii A leoncei Shreve & Williams 1963 EU ricordii A leoni Lynch 1977 EL(t) myersi B leprus Cope 1 879 SY leprus A leptodacty hides Andersson 1 945 - - - = Leptodactylus leptolophus Lynch 1 980 EL(m) leptolophus C leucopus Lynch 1976 EL(m) unistrigatus C lewisi Lynn & Dent 1 942 - - - = luteolus librarius Flores & Vigle 1994 EL(m) unistrigatus c limbatus Cope 1 862 EU - A limbensis Lynn 1958 EU ricordii A lineatissimus Taylor 1941 - - - = mexicanus lindae Duellman 1978 EL(m) unistrigatus C //>j^(3rM5 Brocchi 1879 CR gollmeri A lirellus Dwyer 1995 EL(m) unistrigatus C lividus Lynch & Duellman 1980 EL(m) unistrigatus C locustus Schmidt 1920 EL(m) martinicensis c loki Shannon & Werler 1955 - - - = rhodopis longipes Baird 1859 SY longipes A longirostris Boulenger 1898 CR fitzingeri A loustes Lynch 1979 EL(c) loustes B /mc/o/ Schwartz 1980 EU ricordii A luscombei Duellman & Mendelson 1995 EL(m) unistrigatus C luteolateralis Lynch 1976 EL(m) unistrigatus c luteolus Gosse 1851 EU luteolus A lutitus Lynch 1984 EL(m) unistrigatus C lutosus Barbour & Dunn 1921 - - - = cruentus lymani Barbour & Noble 1920 EL(c) conspicillatus B lynchi Duellman & Simmons 1977 EL(m) unistrigatus C lynni Goin &. Cooper 1950 - - - = cundalli lythrodes Lynch & Lescure 1980 EL(m) unistrigatus C macdougalli Taylor 1942 - - - = lineatus 228 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Appendix III. Continued. Species Tri\ iai name, author, date macrocephalus Peracca 1904 macrotympamtm Taylor 1940 nuicitlata Daudin 1801 nuiciilosus Lynch 1991 ///(//A//;/ Lynch 1980 luantipus Boulenger 1908 iuary,antifi'r Boulenger 1912 mariposa Hedges et al 1992 mannorants Boulenger 1900 niarnockii Cope 1 878 marshal' Lynch 1964 martial' Lynch 1974 martinicensis Tschudi 1838 matudanay\or 1941 maiirus Hedges 1989 niaussi Boettger 1 893 medemi Lynch 1994 megacephalus Cope 1 875 megalops Ruthven 1917 megalotympaniim Shannon & Werler 1955 melanoproctus Rivero 1984 mi'kmostictus Cope 1 875 mehmotrigonwn Schwartz 1966 /«f'//'/!/ Bokermann 1958 memiax 'DxxtWm'dw 1978 mercedesae Lynch & McDiarmid 1978 mercudonensis Schmidt 1933 merostictus Lynch 1984 mexiccmus Brocchi 1877 mexicanus Boulenger 1 898 milesi Schmidt 1933 mimiis Taylor 1955 minimus Tay\or 1940 minutus Nohle 1923 /«/var<7/ Lynch 1984 modestus Tay\oT 1942 modipi'plus Lynch 1981 molinoi Barbour 1928 molybrignus Lynch 1986 mondolfii Rivero 1984 monensis Meerwarth 1 90 1 monnichoriim Dunn 1940 montanus Schm\d\. 1919 montanus l^ayXov 1942 mora Savage 1965 muricatus Lynch & Miyata 1980 museous Ibafiez et al. 1994 mversi Coin & Cochran 1962 Subgenus group Toe condition Comments EL(m) leptolophus C EL(c) conspicillatiis B EL(c) nigrovittatits A siilcatits vcrrucipes not applied = galdi EL(a) abbotti B EL(m) imistrigatus C SY marnockii A - - - = cruentus EL(m) imistrigatus C EL(m) martinicensis C CR milesi A SY nitidus A CR biporcatus A EL(c) conspicillatus B - - - = biporcatus EL(c) - A •v CR alfredi B EL(c) - B CR fitzingeri A EL(a) abbotti B - - - = ockendeni EL(m) unistrigatus C EL(c) - B CR rugulosus A EL(m) unistrigatus C CR rhodopis A — - - = occidentalis CR milesi A CR gollmeri A - - - = pygmaeus EU - A EL(m) unistrigatus C SY modestus A EL(m) unistrigatus C - - - = ridens EL(m) unistrigatus c EL(m) unistrigatus c EU ricordii A CR fitzingeri A EL(m) martinicensis C - - - = sartori EL(m) unistrigatus c EL(m) unistrigatus c EL(m) unistrigatus - EL(t) myersi A ELEUTHERODACTYLUS IN WESTERN ECUADOR 229 Appendix III. Continued. Species Trivial name, author, date Subgenus Toe group condition Comments mystaceus Barbour 1922 nanus Ahl 1933 napeus Jimenez de la Espada 1875 nasiitus Lutz 1 925 Aifl/rttor Taylor 1939 nebulosus TdLyXox 1943 nebulosus Henle 1992 necerus hync\\ 1975 neodreptus Schwartz 1965 nervicus Lynch 1 994 nicefori Cochran & Coin 1 970 nigriventris Lutz 1925 nigrogrisea Andersson 1 945 nigwvittatiis Andersson 1945 nitidiis Peters 1 869 nivocolimae Dixon & Webb 1 966 noblei Barbour & Dunn 1926 nortoni Schwartz 1976 notidodes Schwartz 1966 nubicola Dunn 1926 nubicola Dunn & Emien 1932 nubilis Giinther 1900 nyctophylax \jync\\ 1976 oaxacae l^ayXor 1940 obesiis Barbour 1928 obmutescens Lynch 1980 occidentalis Taylor 1941 ocellatus Lynch & Burrowes 1990 ockendeni Boulenger 1912 ocreatus Lynch 1 98 1 octavioi Bokermann 1965 oeus Heyer 1984 olibrus Schwartz 1960 oligaidax Schwartz & Fowler 1973 omdtemamis Gunihcx 1900 orarius Dixon 1957 orcesi Lynch 1 972 orcutti Dunn 1928 orestes Lynch 1979 orientalis Barbour & Shreve 1937 orientalis Barbour & Shreve 1937 ornatissimiis Dt?,^pax 1911 orocostalis Rivero 1961 orpacobates Lynch et al 1994 orphnolaimus Lynch 1970 oxyrhyncus Dumeril & Bibron 1841 pagmae Fowler pallidus Duellman 1958 - - - = rhodopis - - - = parvus - - - = cornutus EL(c) hinotatus B - - - = berkenbuschii SY pipihms A - - - = caliginosus CR bufonifonnis A EL(a) abbotti B EL(m) unistrigatus C EL(m) iinistrigatus C EL{b) Uicteiis B EL(m) unistrigatus C EL(c) nigrovittatus A SY nitidus A SY modestus A CR golbneri A PE inoptatus B EL(a) abbotti B EU nubicola A - - - = enileni - - - = fitzingeri EL(m) unistrigatus C - - - = mexicanus - - - = punctariolus EL(m) orcesi c CR rhodopis A EL(c) cerasinus B EL(m) unistrigatus C EL(t) myersi A EL(b) hinotatus A EL(b) hinotatus A EL(m) martinicensis C EU gossei A CR rhodopis A SY nitidus A EL(m) orcesi C EU nubicola A EL(c) orestes B EU - A - - - = estradai EL(m) unistrigatus C - - - = bicumulus EL(c) cerasinus B EL(m) unistrigatus C EU ricordii A - - - = achatinus SY modestus A 230 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Appendix III. Continued. Species Trivial name, author, date Subgenus group Toe condition Comments palmutus Boulenger 1882 palmeri Boulenger 1912 pantoni Dunn 1926 parahates Schwartz 1964 paralius Schwartz 1976 paramerus Rivero 1984 parapelates Hedges & Thomas 1987 pardalis Barbour 1928 panilliis Lynch 1976 parx'us Girard 1853 pan-US Barbour & Shreve 1937 pastazensis Andersson 1 945 patriciae Schwartz 1965 paulodutrai Boktrmsinn 1974 paulsoni Schwartz 1964 paululus Lynch 1 974 pecki Duellman & Lynch 1988 pelviculus Cope 1878 pentasyringos Schwartz & Fowler 1973 peraltae Barbour 1928 peraticus hynch 1980 p^rcM/rM5 Lynch 1979 permixtus Lynch et al 1994 persimilis Barbour 1926 peruvianusVieXm 1941 petersi 'DutWmdin 1954 petersi Lynch & Duellman 1980 petersorum Lynch 1991 petrobardus DuQWrmn 1991 petrophilus Firschein 1954 pezopetrus Schwartz 1960 pharangobates Duellman 1978 philipi Lynch & Duellman 1995 phoxocephahis Lynch 1979 phragmipleuwn Rivero & Serna 1987 phyzelus Schwartz 1958 pictissimus Cochran 1 935 pinarensis Dunn 1926 pinchoni Schwartz 1967 pipilans Taylor 1 940 pittieri Giinther 1900 pituinus Schwartz 1965 planirostris Cope 1 862 plat} dactyl us Boulenger 1903 platyrhynchus Giinther 1900 pleurostriatus Rivero 1984 /j//carM5 Giinther 1900 plic atus BdiXhoxxr 1914 = rugulosus EL(m) unistrigatus C EU gossei A EU - A EU ricordii A EL(c) - B PE ruthae B EL(m) unistrigatus C EL(m) unistrigatus C EL(b) parx'us A - - - = cubanus EL(m) unistrigatus C EL(m) martinicensis C EL(b) ramagii B EU ricordii A EL(m) unistrigatus C EL(m) unistrigatus C - - - = biporcatus EU gossei A - - - = cerasinus EL(m) leptolophus C EL(m) unistrigatus C EL(m) unistrigatus c - - - = bransfordii EL(c) conspicillatus B SY nitidus A - - - = johnwrighti - - - = johnwrighti EL(m) unistrigatus C SY marnockii A EU ricordii A EL(m) unistrigatus C EL(m) unistrigatus C EL(m) unistrigatus C EU ricordii A EU ricordii A EL(m) martinicensis C SY pipilans A EL(m) martinicensis C EU ricordii A EL(m) unistrigatus C = rhabdolaemus = varlevi = rugulosus = melanostictus EL(c) B : rhodopis -gundlachi ELEUTHERODACTYLUS IN WESTERN ECUADOR 231 Appendix III. Continued. Species Toe Trivial name, author, date Subgenus group conditior 1 Comments plicifer Boulenger 1 888 EL(b) binotatus A podiciferus Cope 1 875 CR rhodopis A polymniae Campbell et al 1989 CR alfredi B polyptychus Cope 1 886 - - - = bransfordii /7<3o/^/ Cochran 1938 EL(m) martinicensis C portoricensis Schmidt 1 927 EL(m) martinicensis C probolaeus Schwartz 1965 EU ricordii A prolatus Lynch & Duellman 1980 EL(m) unistrigatus C prolixodiscus \-,ync\\ 1978 EL(m) unistrigatus C proserpens Lynch 1 979 EL(m) unistrigatus c pruinatus Myers & Donnelly 1996 EL - B psephosypharus Campbell et al 1994 CR rugulosus A pseudoacuminatus Shreve 1935 EL(m) unistrigatus C pteridophilus Lynch & Duellman 1996 EL(m) unistrigatus C pugnax hynch 1973 EL(m) unistrigatus c pulchrigulus Cope 1862 - - - = fitzingeri pulidoi Rivero 1984 EL(m) tubernasus c pulvinatus Rivero 1968 EL(c) - B pumilio Boulenger 1920 - - - = parvus punctariola Peters 1863 CR rugulosus A purpuras Lynch 1967 CR alfredi B pusillus Bokermann 1967 EL(b) parvus B pycnodermis Lynch 1 979 EL(m) unistrigatus C pygmaeus Taylor 1937 CR rhodopis A pyrrhomerus hynch 1976 EL(t) myersi B quaquaversus Lynch 1974 EL(m) unistrigatus C quinquagesimus Lynch & Trueb 1980 EL(c) devillei B racemus Lynch 1980 EL(m) orcesi C racenisi Rivero 1961 - - - = bicumulus ramagii Boulenger 1888 EL(b) ramagii B ramosi Rivero 1959 - - - = cochranae randorum Heyer 1985 EL(b) - B raniformis Boulenger 1896 CR fitzingeri A ranoides Cope 1886 - - - = rugulosus rayo Savage & DeWeese 1979 CR fitzingeri A rearki Taylor 1952 - - - = bransfordii repens Lynch 1984 EL(t) myersi A reticulatus Walker & Test 1955 EL(m) - C rhabdolaemus Duellman 1978 EL(c) - B rhodesi Schwartz 1980 EU ricordii A rhodopis Cope 1 866 CR rhodopis A rhodoplichus Duellman & Wild 1993 EL(m) unistrigatus C richmondi Stejneger 1904 EU ricordii A ricordii Dumeril & Bibron 1841 EU ricordii A ridens Cope 1 866 EL(m) unistrigatus C riisei Reinhardt & Liitken 1 863 - - - = lentus riveroi Lynch & LaMarca 1 993 EL(m) unistrigatus C riveti Despax 1911 EL(m) unistrigatus c 232 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Appendix III. Continued. Species Trivial name, author, date rogersi Coin 1955 w««W/ Schwartz 1960 rosadoi ¥\oxts 1988 w seme linn s GoxUdLm 1966 roseiis Boulenger 1918 roseus Melin 1941 rostralis Werner 1 896 rozei Rivero 1961 nthkundus Jimenez de la Espada 1875 nibhmaculata Taylor & Smith 1945 nicillensis Cochran 1939 rufescens Duellman & Dixon 1959 nififemoralis Noble & Hassler 1933 riii^osus Peters 1873 riigidosa Cope 1 870 mgidosus Peters 1 870 nddits Lynch 1979 /■///.-/Lynch 1981 ntduie HobXt 1923 nithveni Lynch & Ruiz 1985 salaputium YiwtWm&n 1978 sallaei Gunther 1 868 .va/r«ro/- Taylor 1941 samaipatoe Kohler & Jungfer 1995 sanctaemartae Ruthven 1917 sandersoni Schmidt 1941 saivnartinensis Shannon & Werler 1955 sartori Lynch 1965 satagius Lynch 1 995 savagei Pyburn & Lynch 1981 saxatUis Webb 1962 schmidti Noble 1923 iY7;«/ff/ Duellman 1990 schwartzi Thomas 1966 sciagraphus Schwartz 1973 scitulus Duellman 1978 scolobelpharus Lynch 1 99 1 scolodiscus Lynch & Burrowes 1990 semipalmatus Shreve 1936 sernai Rivero 1985 i/zreve/ Schwartz 1969 sierramaestrae Schmidt 1920 sUvicola Lynch 1 967 simonbolivah Wiens & Coloma 1992 siinoterus Lynch 1 980 siopelus Lynch & Burrowes 1990 sisyphodemus Crombie 1 977 smithi Taylor 1940 sobetes Lynch 1980 Subgenus group Toe condition Comments EU ricordii A EL(m) martinicensis C EL(m) nnistrigatus C - - - = vilarsi ELtm) nnistrigatus C - - - = rosemelinus CR gollmeri A EL(m) nnistrigatus C EL(c) cerasinus B SY leprus A EU ricordii A SY nitidus A EU - A - - - = biporcatus CR rugulosus A - - - = binotatns EL(m) nnistrigatus C EL(s) snlcatus A PE ruthae B EL(c) - B EL(m) nnistrigatus C rhodopis CR rhodopis A EL(c) conspicillatns B EL(c) - B — - - = rugulosus - - - = rhodopis CR rhodopis A EL(c) cnrtipes B EL(c) conspicillatus B SY nitidus A EU ricordii A EL(m) nnistrigatus C EL(m) martinicensis C EU ricordii A EL(c) - B EL(m) leptolophus C EL(m) diastema C EU - A EL(s) snlcatus A EL(m) martinicensis C EU ricordii A CR a If red i B EL(c) orestes B EL(m) orcesi C EL(c) devillei B EU Inteolus A = guttilatns EL(c) surdus ELEUTHERODACTYLUS IN WESTERN ECUADOR 233 Appendix III. Continued. Species Trivial name, author, date Subgenus group Toe condition Comments sommeri Schwartz 1 977 sonans Dunn 1925 spanios Heyer 1985 spatulatus Smith 1939 spilogaster]jync\\ 1984 spinosus Lynch 1979 stadelmani Schmidt 1936 5?aAjto/H' Schmidt 1941 staurometopon Schwartz 1960 stejnegerianus Cope 1893 stenodiscits Walker & Test 1955 sternothylax Duellman & Wild 1993 stuarti Lynch 1967 subsigillatus Boulenger 1902 sulcatus Cope 1874 sulculus Lynch & Burrowes siipe mat is Lynch 1980 surdiis Boulenger 1882 5ymmgto«/ Schwartz 1957 syristes Hoy t 1 965 taeniatus Boulenger 1912 tcdamancae Dnnn 1931 tamsitti Cochran & Coin 1970 tarahumarensis Taylor 40 taurus Taylor 1958 taylo ri hynch 1966 tayrona Lynch & Ruiz 1985 tenebrionis Lynch & Miyata 1980 teretistes Duellman 1958 terraebolivaris Rivero 1961 thectopternus Lynch 1975 thomasi Schwartz 1959 thorectes Hedges 1988 thymalopsoides Lynch 1976 thymelensis Lynch 1972 tiptoni Lynch 1964 toa Estrada & Hedges 1991 ro/rflf Duellman 1978 trachyblepharis Boulenger 1918 trachydermus CampheW 1994 trepidotus Lynch 1968 trinidadensis Schwartz 1959 truebae Lynch & Duellman 1996 tubernasus Rivero 1984 turquinensis Barbour & Shreve 1937 tiirumiqiiirensis Rivero 1961 tychathnnis Schwartz 1965 underwoodi Boulenger 1 896 Mrt/co/or Tschudi 1838 EL(m) martinicensis C - - - = auriculatus EL(b) - B CR a If red i B EL(m) iinistrigatus C EL(m) unistrigatus C CR tnilesi A - - - = laticeps EL(m) martinicensis C CR rhodopis A EL(c) - B EL(m) unistrigatus C CR alfredi B EL(m) unistrigatus C EL(s) sulcatus A EL(c) devillei B EL(m) unistrigatus C EL(c) surdus B EU symingtoni A SY nitidus A EL(m) unistrigatus C CR fitzingeri A EL(m) unistrigatus C CR augusti A CR rugulosus A CR alfredi B EL(m) unistrigatus C EL(c) cerasinus B SY modestus A EL(c) conspicillatus B EL(c) conspicillatus B EU ricordii A EL(a) abbotti B EL(m) unistrigatus C EL(m) orcesi C - - - = cerasinus EU ricordii A EL(c) - B EL(m) unistrigatus C CR milesi A EL(t) myersi A EU ricordii A EL(c) devillei B EL(m) tubernasus C EU ricordii A EL(m) unistrigatus C PE ruthae B - - - = bransfordii - - - = Suppressed 234 UNIV. KANSAS NAT. HIST. MUS. SPEC. PUBL. NO. 23 Appendix III. Continued. Species Toe Trivial name, author, date Subgenus group condition A Comments unicolor Stejneger 1904 EU unistrigatus Giinther 1859 EL(m) unistrigatus ~ C imo Savage 1985 CR - A iiranobates Lynch 1991 EL(m) leptolophus C urichi Boettger 1 894 EL(m) martinicensis C vanadise LaMarca 1984 EL(c) - B variabilis Lynch 1968 EL(m) unistrigatus C varians Gundlach & Peters 1 864 EL(m) martinicensis C varleyi Dunn 1 925 EU - A venancioi Lutz 1958 EL(b) - A ventrilineatus Shreve 1936 EU - A ventrimarmoratus Boulenger 1912 EL(m) unistrigatus C ventrivittatus Andersson 1945 - - - = ventrimarmoratus venustus Giinther 1900 - - - = rhodopis verecundus Lynch & Burrowes 1990 EL(m) unistrigatus C verrucipes Co^t 1885 SY marnockii A verruculatus Peters 1 870 SY - - = nomen dubium vrr^/co/or Lynch 1979 EL(m) unistrigatus C vertebralis Boulenger 1886 EL(c) devillei B vicarius Lynch & Ruiz 1983 EL(m) unistrigatus C vidua Lynch 1979 EL(c) orestes B vilarsi Melin 1941 EL(c) conspicillatus B vm/jfl/ Bokermann 1974 EL(b) - B viridicans Lynch 1977 EL(c) conspicillatus B voca//5 Taylor 1940 CR rugulosus A vocator layXor 1955 EL(m) diastema C vulcani Shannon & Werler 1955 - - - = berkenbuschii w-nigrum Boettger 1892 EL(c) conspicillatus B walked Lynch 1974 EL(m) unistrigatus C warreni Schwartz 1976 EU ricordii A weinlandi Barbour 1914 EU ricordii A werleri Lynch & Fritts 1965 - - - = laticeps wetmorei Cochran 1932 EL(m) martinicensis C whymperi Boulenger 1882 - - - = curtipes wiensi Duellman & Wild 1993 EL(m) unistrigatus c wightmanae Sc\im\d\ 1920 EL(a) - B wilUamsi Rivero 1961 - - - = bicumulus williamsi Schwartz 1968 - - - = diplasius xestus Lynch 1995 EL(c) curtipes B xucanebi Stuart 1941 CR alfredi B yaviensis Myers & Donnelly 1996 EL(m) unistrigatus C yucatanensis Lynch 1965 CR alfredi B zayasi Schwartz 1 960 EU ricordii A zeuctotylus Lynch & Hoogmoed 1 977 EL(c) conspicillatus B zeus Schwartz 1958 EU - B zimmermanae Heyer & Hardy 1991 EL(m) unistrigatus C zugi Schwartz 1958 EU ricordii A zygodactylus Lynch & Myers 1983 CR anomalus A ELEUTHERODACTYLUS IN WESTERN ECUADOR 235 1 ^ (73 r-» 3 ^ ^ o ex C -^ C/3 o 'o o > "B. >< X (U ■ s ^ s^ o > C/3 f. ^' a ^ 3 c/5 "c J3 Si II X) > II CI. o kH o -o t4-H o , s '2 e 3 c h ^ ^ o 1^ ^ II II < 3 X kH ^ II e 'C c c >. ^ w c/3 < • — - o e II H 'So -3 ^ b >. . . * X ^ t to -2 .2 '^' c/3 'q. 13 c •3 3 i 2 X) c tij 4J 2 C/5 ■^ t1 II V3 II s 13 rt 3 13 g CJ X S o Q § " j^ o "o t .2 3 C (U (U i3 '=3 c« C/2 ^ ^ 1^ kj II II N c« CI, II s "o a C 3 O C/5 (U B X s 2 00 00 r-r-~ONOON>/~ir\io 00 -^ VO -^ r<-i T^ \D rr-, \0 -^ — ' — '^ (^ OJ in rNOvD^O'Or^-^OOONr«-iONOOO TfinO — ■^— <^t~~— ir, 0Or<-, Tj- r4 00 — >' •^' On cn — ' — ' — ' m' — <" oooo^'^^'^^'^'^'^'^'^^Z^'-'^Zt/^o^^c/jon&o Tf IT) '^ n in in in Tj- o o ^ o in '^ -^ ~ m >n OrnO'O^S-OO—^'^rNl — '^OO ii 1 o (N O (~N| m m o o o o o o O r-J O 00 O O t^ — 00 r-- 00 r- I (N «N --I -H CN O I I I I o o o o o ^ in ^ in -^ r^ '^ in ^ oooooooooooo O^f^'^OO — — O — OOO r^iinoovDoocNt^i^ — OOc^ — — — '(N(N(N(N(Nr<-) 1 I I I I I I I I I o ooooooooo (N oominin-^ooin oocNiooinininr^t^oo I I O O in o r^ m I I I I I I I I -^ O I (Ti I I I I I m I I I I (N 00 in ONllvDIIIII— ll'OIIIIIIIIII 00 I I I I I I I I I I I -^ I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I < ° .^ .^ c .- - OOCCCIUOCCOC3 C/5 C/^ C^ C/: C/^ GO "^ CA C/l ,2 'c u 'S 'S 'S 3 "S 'c C3 C3 ^ s S -i c: 5 g ^ a c c Ci, ex t3 a C3 a C3 -Q S a Cj Cj cx-S C3 ^ Cj ;j s N ?« h s s 1^ -C Ci C !^ Cj Cj tj o o "<3 -Q "t3 a 3 a ^ -^ cu ^ 5 s ;2 ■-: s 's; C S o w kjkji-tjijjijjkikikiujkikiiiikiijjkikiijQkikiuj 236 HIST. MUS. SPEC. PUBL. NO. 23 093 353 142 CNOOOOOOO — — ■^ — O0s00r<"i CN — ^ O Ov C7> I^r<-i--fvD0N'^O'nrjr<-jvOI^'^r-l00I~~-O^OOO^00O — '^ON^^vO oooo>r"jO>nooO — ^ooOO^ON-^t^ — in — — Orooo — '— r^l — — ^^-Drl oi ri — m oi — — — r<-j (N — ' — ^ C?> OO (» Cn| vo -^ -^ r~- ON viu^'^viUiQnuicAv^i/i'Z^^^^'^'^Z'^^^^^'^'Z.'^'Z,^^^^^'^'^'^'^^^^ •^ ON — o o o o o o o o O O O O O O (N I 1 I I I I z z z z z z — „ m lO o I I 00 or) ^ O^ — "/-lOooX- o b r) ^ — " o ^ o o o o o — O (N O O I I I z z z in — < no O '^ — r) O 'I- mj o ri — ^ rj ^ — r) -t oi 00 in O -+ oooooor^ooooooooooooo OO — O — t^ — OOO — r<-)ONrnoOONOOOOt^ oo — i^Or^inr<-)r^r<-iOOs^oo^^Ttoooorin ~~ rr^ n r^i C^\ in — ^ m r) \ r^i I (N r*"! f; — rl r<-, — IIIIIO II OIOIIIIIII ooomoin oo oonooooooo "OOr>— oo — inrivor-OOOOO no in -^ 00 -^ ON 00 00 — — r<-, — n n n — I I I I I llllllll— I— llll — I NO — I I I I r) I I — 1 -H in I I I I I I n — n r) n -t r- ON I — — ^ — r-iTf-^oo I'Tf — o inNOt^r^roi^noi \ —' m Tj-i I I iml^ioi-^ <^l|||-H||||^l— llllllni I I I I I I I I I r- I I I I llllllllll— 'lllllll'^NO I I 1 I I I —^ I n I I I I I I I I I I I I I I — I I I I I I I I — N':= ^ S S S c o _ c^ C/5 v^ QJ c 'c =^ i:i == § 3 e « ° c/3 c3 c/} ^ p O 3 p 3 3 ED c c .S 3 3 -O i-3C3ajC(UCCiL>OC Oc«3(/3U3T333'Ou3 i 2 to -^ .5 5 -2 2 tq kj "3 ."^ '^ 5u ^ -2 "r- '^ ^ :^ ? "3 -r O S 5u ttj kj kj "3 .c; ~ ~ '^ 2; to >. J" Sr^cT:=:a-^-2s^^-33 kj kj ci. Ci. o, a, cr c 2 O S ^ "5 to ~5 5 ^ ^ •2? to "g 2 S ii o s s ^) -r k ^ :^ -o nr 5ij -^ Ji ^ i kj kJ kq kJ kj kjkjkjkjkjkjkjkjklkjkjkjkjkjklkqkqkjkjkj SPECIAL PUBLICATIONS OF THE UNIVERSITY OF KANSAS NATURAL HISTORY MUSEUM (Numbers cuirently available) 3. Maintenance of Rattlesnakes in Captivity. By James B. Muiphy and Barry L. Amistrong. Pp. 1^0. 29 December 1978. ISBN 0-89338-006-7. 5. The Natural History of Mexican Rattlesnakes. By Barry L. Armstrong and James B. Murphy. Pp. 1-88. 14 December 1979. ISBN 0-89338-010-5. 7. A Diapsid Reptile from the Pennsylvanian of Kansas. By Robert R. Reisz. Pp. 1- 74. 18 February 1981. ISBN 0-89338-01 1-3. 9. The Ecological Iinpact of Man on the South Florida Herpetofauna. By Larry DavidWilson and Louis Porras. Pp. 1-89. 8 August 1983. ISBN 0-89338-018-0. 10. Vertebrate Ecology and Systematics: A Tribute to Henry S. Fitch. Edited by Richard A. Seigel, Lawrence E. Hunt, James L. Knight, Luis Malaret, and Nancy Zuschlag. Pp. 1-277. 21 June 1984. ISBN 0-89338-019-0. 13. Geographic Variation Among Brown and Grizzly Bears ( Ursus arctos) in North America. By E. Raymond Hall. Pp. 1-16. 10 August 1984. 15. Spring Geese and Other Poems. By Denise Low. Pp. 1-84. September 1984. ISBN 0-89338-024-5. 18. A Checklist of the Vertebrate Animals of Kansas. By George D. Potts and Joseph T. Collins. Pp. 1^2. September 1991. ISBN 0-89338-038-5. 1 9. The Compleat Cladist. A Primer of Phylogenetic Procedures. By E. O. Wiley, D. Siegel-Causey, D. R. Brooks, and V. A. Funk. Pp. 1 - 1 58. October 1 99 1 . ISBN 0- 89338-035-0. 21. Amphibian Species of the World: Additions and Corrections. By William E. Duellman. Pp. 1-372. July 1993. ISBN 0-89338-045-8. 22. Guide to the Frogs of the Iquitos Region, Amazonian Peru. By Lily O. Rodriguez and William E. Duellman. Pp. 1-80 + 12 color plates. December 1994. ISBN 0- 89338-047-4. pPTODACTYLiD FROGS of the genus Eleutberodactylus are the most diverse and abundant terrestrial vertebrates in the American tropics. They form a conspicuous ele- ment of the frog fauna on the Pacific lowlands and slopes of the Andes in western Ecuador. John D. Lynch and Wil- liam E. Duellman have studied these frogs in western Ec- uador for the past 30 years. In this book, they recognize 61 species (with 9 species described as new) and pro- vide diagnoses of each species and accumulated infor- mation on their ecology, life history, and distribution. Keys in English and Spanish facilitate identification. Dis- tribution maps are provided for each species, and 57 spe- cies are illustrated in 64 color photographs. Nine com- munities oiEleutherodactylus are analyzed ecologically, and the entire eleutherodactyline fauna is treated bio- geographically. This volume also contains definitions and illustrations of taxonomic characters of Eleutherodacty' lus, definitions and subgenera and series, and a list of all species (including synonyms) in the genus with alloca- tions to subgenera and species groups. Thus, this is the definitive work on the largest genus of vertebrates and a thorough account of the genus in western Ecuador.