UNX HARVARD UNIVERSITY Library of the Museum of Comparative Zoology <. io^- XU^t^^^-^^^ UNIVERSITY OF KANSAS MUSEUM OF NATURAL HISTORY MUS. COM P. ZOOU. LIBRARY SEP 5 1978 MISCELLANEOUS PUBLICATION NO. 65 HARVARD UNIVERSITY The Biology of an Equatorial Herpetof auna in Amazonian Ecuador By William E. Duellman UNIVERSITY OF KANSAS LAWRENCE 1978 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers fonnerly published in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publications, Museum of Natural History, began with number 1 in 1946. Longer research papers are pub- lished in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subjected to critical review by intra- and extramural specialists; final acceptance is at the discretion of the publications committee. Institutional libraries interested in exchanging publications may obtain the Occa- sional Papers and Miscellaneous Publications by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas 66045. Individuals may purchase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Law- rence, Kansas 66045. The University of Kansas Museum of Natural History Miscellaneous Publication No. 65 August 30. 197S The Biology o£ an Equatorial Herpetofauna in Amazonian Ecuador By William E. Duellman Curator Division of Herpetologij Museum of Natural History Professor Department of Systematics and Ecology The University of Kansas Lawrence, Kansas 66045 U.S.A. Publication Supported by a Grant from the National Science Foundation (DEB 77-28380) The Unwersity of Kansas Lawrence 1978 University of Kansas Publications, Museum of Natural History Editor: Linda Trueb Miscellaneous Publication No. 65 pp. 1-352; 198 figures; 34 tables Published August 30, 1978 Museum of Natural History The University of Kansas Lawrence, Kansas 66045 U.S.A. Printed by Univehsity or Kansas Prixtixg Ser\ice Lawrence, Kansas CONTENTS INTRODUCTION 9 Acknowledgments 10 Scope of study 11 Methods 13 Reliability and Limitations of Data 13 Description of the Area 14 Physiography and Hydrography 17 Climate 19 Vegetation 22 Habitats 27 IDENTIFICATION OF SPECIES 29 Key to the Orders and Suborders of Amphibians and Reptiles 30 Key to the Caecilians 31 Key to the Salamanders 31 Key to the Frogs 31 Key to the Turtles 48 Key to the Crocodilians 49 Key to the Lizards and Amphisbaenians 49 Key to the Snakes 58 Key to the Known Tadpoles 70 SPECIES ACCOUNTS 78 Caecilians 79 Caeciliidae 79 Caecilia disossea Taylor 79 Caecilia tentaculata (Linnaeus) 79 Microcaecilia albiceps (Boulenger) 79 Oscaecilia hassleri (Dunn) 80 Siphonops annulatus (Mikan) 80 Salamanders 80 Plethodontidae 80 Bolitoglossa ecuadoriana Brame and Wake 80 Bolitoglossa peruviana (Boulenger) 80 Frogs 82 Pipidae 82 Pipa pipa (Linnaeus) 82 Leptodactylidae 83 Adenomera andreae (Miiller) 83 Ceratophrys cormita (Linnaeus) 85 Edalorhina perezi Jimenez de la Espada 86 Eleiitherodactijlus acuminatus Shreve 86 Eleutherodactyhis altamazonicus Barbour and Dunn 87 Eleutherodactijlus conspicUlatus (Giinther) 89 Eleutherodactyhis croceoinguinis Lynch 90 Eleutherodactyhis diadematus (Jimenez de la Espada) 91 Eleutlierodactylus lacrimosus (Jimenez de la Espada) 92 Eleutherodactyhis lanthanites Lynch 93 Eleutlierodactylus martiae Lynch 94 Eleutlierodactylus nigrovittatus Andersson 96 Eleutlierodactylus ockendeni (Boulenger) 97 Eleiitherodacfyhis orphnolaimus Lynch 98 Eleutherodactyhis pauhdus Lynch 98 EJeutherodactyJus pseudoacuminotiis Shreve 99 Eleutherodactyhis quaqtiaversus Lynch 100 Eleutherodactyhis sulcatus (Cope) 102 Eleutherodactyhis variahdis Lynch 103 Ischnocncma quixensis (Jimenez de la Espada) 104 Leptodactyhis discodactylus Boulenger 105 Leptodactylus mystaceus (Spix) 107 Leptodactyhis pentadactylus (Laurenti) 108 Leptodactylus rhodomystax Boulenger 111 Leptodactylus stenodema Jimenez de la Espada 112 Leptodactylus ivagneri (Peters) 112 Lithodytes lincatus (Schneider) 114 Physalaemus petersi (Jimenez de la Espada) 115 Bufonidae 116 Bufo glaberritiius (Giinther) 116 Bufo inarinus (Linnaeus) 117 Bufo typhonhis (Linnaeus) 118 Dendrophryniscus muiutus (Melin) 120 Dendrobatidae 121 Colostethus marchesianus (Melin) 121 Colostethus saidi Edwards 123 Dendrobates parvulus Boulenger 124 Dendrobates pictus pictus Dumeril and Bibron 125 Dendrobates quinquevitfatiis Steindachner 125 Phyllobates femoralis (Boulenger) 126 Hylidae 127 Hemiphractus proboscideus (Jimenez de la Espada) 127 Hyla alboguttata Boulenger 128 Hyla bifurca Andersson 130 Hyla boons (Linnaeus) 131 Hyla bokermanni Coin 133 Hyla brevifrons Duellman and Crump 135 Hyla calcarata Troschel . 137 Hyla cruentonima Duellman 139 Hyla fasciata Giinther 140 Hyla favosa Cope 142 Hyla funerea ( Cope ) 143 Hyla garbei (Miranda-Ribeiro) 144 Hyla geographica Spix 146 Hyla granosa Boulenger 148 Hyla lanciformis (Cope) 150 Hyla leucophyllata (Bereis) 152 Hyla marmorata (Laurenti) 153 Hyla minuta Peters 155 Hyla parviceps Boulenger 156 Hyla punctata (Schneider) 158 Hyla rhodopepla Giinther 159 Hyla riveroi Cochran and Goin 161 Htjla rossalleni Goin 162 Hijla rubra Laurenti 163 Hyla sarayacuensis Shreve 165 Hyla triangulum Giinther 167 Nyctimantis rugiceps Boulenger . 168 Osteocephalus huckleyi (Boulenger) 170 Osteocephahis leprieurii (Dumeril and Bibron) 171 Osteocephalus taurinus Steindachner 172 Phyrnohyas coriacea ( Peters ) 173 Phrynohyas venulosa (Laurenti) 174 Phyllomedusa paUiata Peters 176 PhyUomcdusa tarsius (Cope) 177 Phyllomedusa tomopterna (Cope) 179 Phyllomedusa vaillanti Boulenger 180 Sphaenorhynchus carneus (Cope) 181 Sphaenorhynchus eurhostus Rivero 182 Centrolenidae 183 Centrolenella midas Lynch and Duellman 183 Centrolenella munozorum Lvnch and Duellman 184 Centrolenella resplendens Lynch and Duellman 185 Ranidae 186 Rana palmipes Spix 186 Microhylidae 187 Chiasmocleis anatipes Walker and Duellman 187 Chiasmocleis hassleri Dunn 188 Chiasmocleis ventrimacidata (Andersson) 189 Ctenophryne geayi Mocquard 190 Hamptophryne holiviana (Parker) 190 Syncope antenori Walker 192 Turtles 192 Chehidae 192 Chelus fimbriatus (Schneider) 192 Mesoclemnujs gibba (Schweigger) 192 Phrynops geoffroanus tuberosus (Peters) 193 Platemys platycephala (Schneider) 193 Kinosternidae 193 Kinosternon scorpioides (Linnaeus) 193 Testudinidae 194 Geochelone denticulata (Linnaeus) 194 Crocodilians 194 Crocodylidae 194 Caiman crocodilus crocodilus ( Linnaeus ) 194 Paleosuchus trigonatus (Schneider) 195 Lizards 195 Gekkonidae 195 Gonatodes concinnatus (O'Shaughnessy) 195 Pseudogonatodes guianensis Parker 196 Thecadactylus rapicauda (Houttuyn) 197 Iguanidae 198 Anolis chrysolepis scypheus Cope 198 Anolis fuscoauratus fiiscoouratus D'Orbigny 199 Anolis ortonii Cope . 200 Anolis punctatiis boulengeri O'Shaughnessy 201 Anolis trachyderma Cope 201 Anolis transversalis Dumeril 202 Enyalioides cofanonim Duellman 203 Enijalioides laticeps fesfae Peracca 204 Plica umbra ocJirocoIlaris (Spix) 206 Polychrus marmoratus (Linnaeus) 207 Uracentron flaviceps (Guichenot) 208 Scincidae 208 Mabitya mabouya (Lacepede) 208 Teiidae " 209 Alopoglossus atriventris Duellman 209 Alopoglossus copii Boulenger 210 Ameiva ameiva petersii Cope 210 Arthrosatira reticulata (O'Shaughnessy) 213 Bachia trinasale trinasale (Cope) 213 Dracaena guianensis Daudin 214 Iphisa elegans Gray 215 Kentropyx pelviceps Cope 215 Leposoma parietale ( Cope ) 217 Neusticurus ecpleopus Cope 218 Prionodactylus argulus (Peters) 220 Prionodacfylus manicatus (O'Shaughnessy) 221 PtycJioglossus brevifrontalis Boulenger 222 Tupinambis teguixin (Linnaeus) 223 Amphisbaenians 224 Amphisbaenidae 224 Amphisbaena fuliginosa Linnaeus 224 Snakes 224 Aniliidae 224 Anilius scytale (Linnaeus) 224 Boidae 225 Boa constrictor constrictor Linnaeus 225 Corallus caninus (Linnaeus) 225 Corallus enydris enydris (Linnaeus) 226 Epicrates cenchria cenchria (Linnaeus) 226 Eunectes murinus murinus (Linnaeus) 227 Colubridae 227 Atractus claps (Giinther) 227 Atractus major Boulenger 229 Attractus occipitoalbus (Jan) 230 Chironius carinatus (Linnaeus) 231 Chironius fiiscus (Linnaeus) 232 Chironius midtiventris Schmidt and Walker 233 Chironius scurrulus (Wagler) 234 Clelia clelia clelia ( Daudin ) 235 Dendrophidion dcndrophis (Schlegel) 236 Dipsas catesbyi (Sentzen) - 237 Dipsas indica ecuadorensis Peters 238 Dipsas pavonina Schlegel 239 Drepanoides anomalus (Jan) _.._ 240 Dnjmohius rhomhifer (Gunther) 240 Drymohiber dichrous (Peters) 241 Enjthrolamprus aescidapu aescidapii (Linnaeus) 242 Helicops angulatiis (Linnaeus) 242 Helicops petersi Rossman 243 Imantodes cenchoa cenchoa (Linnaeus) 244 Imantodes lentiferus (Cope) 245 Leimadopliis reginae (Linnaeus) 246 Leimadophis sp. 247 Leptodeira aimidata annuJata (Linnaeus) 247 Leptophis aluietuUa nigroinarginatus (Giinther) 249 Liophis cohella (Linnaeus) 250 ISlinia hudsoni Parker 250 Oxybelis argenteus (Daudin) 251 Oxyrhopus formosus (Wied) 253 Oxyrhopits meJanogenys (Tschudi) 253 Oxyrhopus petola digitalis (Reuss) 254 Pseudoboa coronata Schneider 255 Pseustes sidphureiis sidphureus (Wagler) 255 Rhadinaea brevirostris ( Peters ) 255 Siphlophis cervimis (Laurenti) 256 Tantilla melanocephala melanocephala (Linnaeus) 256 Tripanurgos compressus (Daudin) 258 Xenodon severus (Linnaeus) 258 Xenopholis scalaris (Wucherer) 260 Elapidae 260 Leptomicrurus nardiiccii (Jan) 260 Micrurus langsdorffi langsdorffi Wagler 260 Micrurus lemniscatus helleri Schmidt and Schmidt 261 Micrurus spixii obscurus (Jan) 261 Micrurus surinamensis surinamensis (Cuvier) 262 Crotahdae 262 Bothrops atrox (Linnaeus) 262 Bothrops biUneatus smaragdinus Hoge 264 Bothrops castehiaudi Dumeril, Bibron and Dumeril 264 Lachesis muta muta (Linnaeus) 265 REPRODUCTIVE BIOLOGY 265 Annual Reproductive Patterns 266 Anurans 266 Lizards 270 Snakes 271 Fecundity 271 Anurans ^ 272 Lizards 273 Snakes 276 Reproductive Strategies 280 Anurans 280 Lizards 285 Snakes 287 Discussion 288 Reproduction in Other Groups 288 Reproductive Potential 288 General Conclusions 289 Seasonal Versus Aseasonal Environments 289 COMMUNITY STRUCTURE 291 Habitat Analysis 294 Primary Forest 298 Secondary Forest 310 Clearings 311 Diel Activity 312 Diet and Feeding 313 Size Ratios 314 Tadpoles 315 Anuran Mating Calls 318 Niche Dimensions 319 Species Diversity 323 Discussion — 327 Seasonal Versus Aseasonal Environments 330 BIOGEOGRAPHY 331 Distribution of the Herpctofauna 331 Origin of the Herpctofauna 335 Tertiary and Quaternary History of the Amazonian Biota 336 Evolution of the Herpetofaunal Community 338 EPILOGUE 339 RESUMEN 341 LITERATURE CITED 345 PLATES FOLLOW PAGE 192 EQUATORIAL HERPETOFAUNA 9 INTRODUCTION Civilized man has walked on the moon and is carrying out exploratory probes of outer space; yet there remain on earth two nearly unknown ecosystems — the depths of the oceans and the low- land tropical rainforest. Technological problems limit the exploration of oceans, but the tropical lowlands? Is the "fore- boding jungle," to which the exaggerated writings of travelers and adventurers all- too-often refer, beyond the limits of mod- ern technology? In the Nineteenth Centuiy a few nat- uralists ventured into the Amazon Basin in South America. The vast collections and provocative writings of Henry W. Bates, Alexander von Humboldt, Richard Spruce, and Alfred R. Wallace should have provided the impetus for continued and intensive exploration and study. Af- rica, with its relatively small Congo Basin, was the "dark continent" to be explored, colonized, and exploited, whereas South America, which so richly deserves the appelation given to Africa, languished. Although the short-lived "rubber boom" late in the last century left an opera house in Manaus, the Ama- zon Basin remained a biological tierra incognita. Now the Amazon Basin is being ex- ploited— before it has been adequately explored and studied. Human popula- tion pressure coupled with food and en- ergy shortages seem to make it "practi- cal" to settle the vast region, constiTict roads, clear the forests, cultivate the soil, and drill for oil. The lateritic soil is leached and eroded after two or three crops; the rivers are polluted; and we are losing forever a part of the world's biota. For the most part biologists are still ignorant of the fauna and flora of the Amazon Basin and have little accu- rate conception of the ecology of the rainforest. Ironically, the Biome Study Committee of the International Biologi- cal Program never succeeded in initiating a study of the tropical rainforest. Today many biologists are concerned primarily with theoretical aspects of pop- ulation biology, ecology, and evolution. Models of species diversity, niche breadth and overlap, resource partition- ing, competition, predator-prey, and r and K selection abound in the literature. But how do these models apply to the tropics? Where and how can they be tested? Although inferences frequently are made in the literature, base line data are wanting in most cases. The present report is an attempt to present base line data and preliminary interpretations. Hopefully, they will pro- vide future investigators with the in- formation and impetus for more inten- sive and varied ecological and popula- tional studies, which in trnn will furnish the bases for testing models. This report concerns the amphibians and reptiles of Santa Cecilia, a small area in the upper Amazon Basin. It deals with the amphib- ians and reptiles for two reasons. First, I am a herpetologist and am most knowl- edgeable about those groups. Second, amphibians and reptiles are ideal organ- isms for faunal study in the tropics. They are closely tied to the environment, usu- ally are independent of given species of plants, and are sufficiently numerous and well enough known to be studied feasi- bly. The choice of the study area was completely fortuitous. During the past ten years numerous publications dealing with the herpeto- fauna of the Santa Cecilia region have appeared. Many of these have been descriptions of new species: Eleuthero- dactyhis croceoinguinis and variahilis (Lynch, 1968); E. orphnolaimtis (Lynch, 1970); E. martiae, pcmhdus, and qiiaqtia- versus (Lynch, 1974); E. lanthanites (Lynch, 1975); CoJostethus satili (Ed- wards, 1974); Hyla ])revifrons (Duell- man and Crump, 1974); }Iijlo cruentom- ma (Duellman, 1972a); Centrolenella midas, munozorum, and resplendens (Lynch and Duellman, 1973); Chiasmo- 10 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY cleis anatipes (Walker and Duellman, 1974); Syncope antenori (Walker, 1973); Emjaliokles cofonorum and Alopo^Iossiis atriventris (Duellman, 1973c); and HeJi- cops petersi (Rossman, 1976). Other papers deal with the taxonomic status of various taxa: Phrynohyas coriacea (Duellman, 1968); Hyla funerea (Duell- man, 1971b); Hyla fl.(irhci (Duellman, 1970a); Hyla rhodopepla (Duellman, 1972c); Hyla punctata, rossalleni, leuco- phyllata group and Sphaenorhynchus carneus (Duellman, 1974a); and Ama- zonian PhyUomcdusa (Duellman, 1974b). Material from Santa Cecilia has been used in the following systematic reviews: Hyla geographica group (Duellman, 1973b); Hyla parviceps group (Duell- man and Crump, 1974); Hyla rostrata group (Duellman, 1972b); Osteocepha- lus (Trueb and Duellman, 1971); Hemi- phractus (Trueb, 1974); Nyctimantis (Duellman and Trueb, 1976); and Ecua- dorian caecilians (Taylor and Peters, 1974). Fitch (196S) reported on body temperatures of several species of lizards. Crump ( 1974) provided a thorough anal- ysis of reproductive strategies in the anurans, and Simmons (1975) analyzed the female reproductive cycle in Ameiva ameiva. ACKNOWLEDGMENTS Field studies were initiated at Santa Cecilia primarily because of the existence of a base camp, "Muiiozlandia." Our work there was made possible by Ing. Ildefonso Murioz B., who provided hous- ing and a laboratory. He and his family made it possible for us to devote full time to biological studies while enjoying rustic, but comfortable, accommodations in the jungle. For their gracious hospi- tality, which included numerous fiestas and despedidas, I extend my sincere miichissimas gracias to the Muiioz fam- ily. I am grateful to the personnel of Gulf Oil Ecuador and Texaco Petroleum Com- pany, Ecuador, for providing air trans- portation between Quito and the Oriente and also for many other courtesies. Ec- uadorian anny personnel stationed at Santa Cecilia aided us in many ways; I am especially indebted to Colonel Ra- mirez. Our work in Quito was greatly facilitated by personnel at the Universi- dad Catolica del Ecuador; I especially thank Father Gustavo Maldonado, Dra. Olga Hcrrera de MacBryde, and the late Francisco Leon. Muiiozlandia was "discovered" by Charles M. Fugler, fomierly of Auburn University. I will be forever grateful to Fugler for his telephone call to me in October 1966 in which he suggested Santa Cecilia as a study site. The data and specimens upon which this report is based were accumulated mostly by field parties from The Univer- sity of Kansas. I thank each person for their contribution and list them in de- scending order of the amount of time spent at Santa Cecilia: Martha L. CiTunp, John E. Simmons, Linda Trueb, Stephen R. Edwards, James W. Wad- dick, Thomas H. Fritts, Charles F. Walker, John D. Lynch, Henry S. Fitch, Abraham Goldgewicht K., Werner C. A. Bokermann, Joseph T. Collins, Arthur C. Echternacht, and Robert W. Henderson. In addition to these persons, all of whom were primarily concerned with studies on the herpetofauna, I am indebted to Philip S. Humphrey, Stephen R. Humph- rey, Frank B. Cross, Gerald R. Smith, and especially William G. Saul for their con- tributions made incidental to their stud- ies on the birds and fishes. Charles M. Fugler, George Key, and Terry D. Schwaner collected amphibians and rep- tiles at Santa Cecilia; their specimens were deposited at Auburn LTniversity Museum and the Museum of Compara- tive Zoology at Harvard University. I am indebted to Schwaner for field notes and photographs and to Robert H. Mount and Ernest E. Williams for the loan of specimens. Charles F. Walker kindly made available material that he collected and deposited in the University of Mich- EQUATORIAL HERPETOFAUNA 11 igan Museum of Zoology. Ronn Altig, Mississippi State University, collected at Santa Cecilia in 1973-1975; I am grateful to him for making material available to me. Some of the limited information pro- vided herein on the geology and vegeta- tion was derived in part from infonna- tion provided by Richard Govett of the Texaco Petroleum Company, John Dwyer of the Missouri Botanical Garden, and Bruce MacBryde, fonnerly of the Universidad Catolica del Ecuador. I was fortunate in being able to spend time in the field with each of them. The identification of the material col- lected at Santa Cecilia has been a long and arduous task that required visits to many museums; I am grateful to curators of these collections in the United States, Europe, and South America for their hos- pitality and provision of working space. Specimens of some groups have been studied by others, who have provided me with identifications and insights into the systematics of these groups. Thus, I am grateful to Edward H. Taylor (cae- cilians), Federico Medem (pleurodire turtles), Charles W. Myers (Leima- (lophis and Rhadinaea) , Douglas A. Rossman (Helicops), W. Ronald Heyer (LeptodacfyJits), and Charles F. Walker (microhylids), and especially John D. Lynch for his efforts on the difficult genus Eleutherodoctyhis. I am extremely appreciative of the aid provided by the late James A. Peters, who on many oc- casions shared with me his extensive knowledge of the Ecuadorian herpeto- fauna. The field studies were variously sup- ported by the Herpetological Research Fund, F. William Saul Fund, and the Watkins Museum of Natural History Grants, all of The University of Kansas. Crump's field work in 1971-72 was sup- ported by the National Science Founda- tion (GB-29557). Visits to European museums were made possible by a grant from the Penrose Fund of the American Philosophical Society (No. 5063), and part of the laboratory work was sup- ported by a grant from the General Re- search Fund, The University of Kansas. A grant from the National Science Foun- dation (GB-35483) made possible the completion of the research. During the preparation of this report I have been assisted by Jan Caldwell, Albert Fisher, Julian C. Lee, John E. Simmons, and especially Martha L. Crump. I sincerely thank them for their careful and extensive work. I am in- debted to James R. Dixon and John D. Lynch for critically reviewing the taxo- nomic accounts and to Martha L. Crump for her provocative discussions and criti- cal review of the manuscript. I am grate- ful to Juan R. Leon and Jaime E. Pefaur for translating the keys and summary into Spanish. Many of the photographs in this report were taken by Crump and Simmons; I thank them for permission to reproduce the photographs, and I am grateful to Simmons for his painstaking work in the darkroom. Indeed I have been fortunate in hav- ing the opportunities to undertake this research and in having the cooperation of so many associates. But my most cher- ished fortune is my wife, Linda Tmeb. She accompanied me on four trips to Santa Cecilia, assiduously worked in the field, and collaborated in several prelim- inary reports. She has provided many ideas for the organization, analysis, and interpretation of the data, and she has critically read the entire manuscript. But her major contributions to this report are her illustrations, the usefulness of which shall, I fear, long outlive my less colorful prose. Piers has been a labor of love, which I can never repay in kind. SCOPE OF STUDY The material presented herein is the result of 48 man months of field work at Santa Cecilia on the Rio Aguarico, Provincia Napo, Ecuador, plus smaller collections of specimens and data from four nearby localities along the Rio 12 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Table 1. — Taxonomic Summary of the Herpetofauna of the Upper Rio Aguarico, Ecuador. ( The numbers given first are for the entire area; numbers in parentheses are for Santa CeciHa only ) Order: Family Genera Species Specimens Gymnophiona Caeciliidae 4 (2) 5 (3) 65 (22) Caudata Plethodontidae 1 (1) 2 (2) 122 (22) Anura Pipidae 1(1) 1(1) 8(8) Leptodactylidae - 8 (8) 28 (25) 2109 (1602) Bufonidae 2 (2) 4 (4) 473 (456) Dendrobatidae 3 (3) 6 (5) 579 (526) Hylidae 7 (7) 38 (37) 3060 (2859) Centrolenidae 1 (1) 3 (3) 27 (20) Ranidae 1 (1) 1 (1) 98 (97) Microhylidae 4 (3) 6 (5) 111 (97) Testudines Cheliidae 4 (4) 4 (4) 18 (17) Kinostemidae 1(1) 1(1) 9(9) Testudinidae 1 (1) 1 (1) 7 (3) Crocodilia Crocodylidae 2 (2) 2 (2) 16 (13) Sauria Gekkonidae 3 (3) 3 (3) 62 (47) Iguanidae --- 5 (5) 11 (10) 849 (749) Scincidae 1 (1) 1 (1) 44 (35) Teiidae 12 (12) 14 (13) 823 (724) Amphisbaenia Amphisbaenidae 1(1) 1(1) 6(3) Serpentes Aniliidae 1(1) 1(1) 7(3) Boidae ^^,- 4 (4) 5 (5) 21 (19) Colubridae 26 (24) 38 (36) 459 (384) Elapidae 2 (2) 5 (5) 18 (15) Crotalidae 2 (2) 4 (4) 41 (35) Totals 97 (92) 185 (173) 9035 (7765) Aguarico. The area that was studied at Santa Ceciha consisted of about 3 square kilometers. During the course of the field work, efforts were made to collect series of each species from every month of the year and to obtain data on the micro- habitat, activity, and life history of each species. As a result of the field work, 9035 specimens (+292 lots of tadpoles) representing 185 species were obtained; 7765 specimens ( +2n.AAVO^ Meters Fig. 5. — Map of tlie study area at Santa Cecilia. All work was done between tlie Rio Aguarico and the Rio Conejo and from the small lake at the extreme left to die eastern end of die airstrip. (1975), the velocity frequently exceeds 2.0 meters per second, and the surface temperature varies between 18° and 20° C. Water level fluctuations of about 5 meters were common and mostly asso- ciated with heavy rains on the Andean slopes. At times of high water, the size of the island at Santa Cecilia was markedly reduced, and local residents claimed to have seen the entire island covered with water. In contrast, at times of low water a broad, boulder and silt bar connects the western end of the island with the north bank of the river (Fig. 6), resulting in an extensive backwater la- goon on the north side of the island. The Rio Conejo is a shallow (up to 2 meters), meandering stream about 6 meters in width. In undisturbed areas the river is concealed by the forest. The velocity is 0.3-0.6 meters per second, and the surface temperatures are 22-23° C. (Saul, 1975). Other streams in the area are small (< 3 meters wide) and shal- low. They vary in gradient from some spring-fed rivulets that descend over rock and gravel from the ridge to the Rio Aguarico to silt-bottomed streams drain- ing lakes and swamps. Two pennanent lakes are present in the area. The southern lake is the larg- est (± 150 X 300 m), whereas the northern is about 100 m in diameter. Both lakes are deep; actual depths arc unknown but exceed 10 m. Originally these eutrophic lakes were bordered by EQUATORIAL HERPETOFAUNA 19 M^ K: 1 1 Ml Ifi ^1 .^ M B 1 g ' y^^ if^ r tam -«■•>— ^ P*^MWiM H 1 '..:,.. •;<»'!* p /S^ ^&'' ^■jfck • .j^ • i • •J' 1 ^ '. ,. r ■ ^* -i.-*. "?^v ^1^ :f-^T,^^^-7^^ * flj -'^*i S^-^'^Lr- ♦<»^.- - .., 'V -^^ .iLiJJh?-^' ''■^!^ K^ -^^'^ • V ■ ■', * ^^^CJ^'*' _f .i 'M n. -^i':r%J Fig. 6. — The Rio Aguarico, looking downstream from Muiiozlandia. In the foreground is the bar extending to tlie island at a time of low water. primary forest (Fig. 7). Surface tem- peratures of the southern lake fluctuate daily from 23.8° to 27.8° C (Saul, 1975). A smaller, shallow lake ( ± 25 X 75 m) exists in the western part of the study area. Infrequently it is reduced to a small, muddy pool. There are numerous swamps in de- pressions, and after heavy rains these and other low-lying areas offer extensive aquatic habitats that are drained by the numerous small streams. Climate The climate at Santa Cecilia is typi- cally Af in the Koppen classification, the climate type characteristic of the upper Amazon Basin (Eidt, 1969). Here we have essentially an aseasonal climate. This is not to say that fluctuations do not occur, but there are no distinct seasons. The only complete year of climatic data is for the period 29 June 1971 through 14 July 1972 (Fig. 8; Table 2). The fol- lowing discussion also includes data from June-July, 1967, June-August, 1968, April-May, 1969, and April-May, 1973. Winds are uncommon at Santa Ce- cilia; as often as not, a heavy rain may be preceded by a gentle breeze. Day length is nearly constant, and effective daylight is controlled more by cloud cover than by the seasonal position of the sun. The daily temperature fluctuation is usually about 10° C. The greatest daily fluctuation recorded was 22-37° C on 9 August 1968; this also was the highest temperature recorded. The least fluctua- tion was 21.5-23.0 C on 22 April and 18 June 1972. The lowest temperature (17.0° C) was on the night of 17 June 1968. The lowest daily maximum tem- peratures and the lowest daily fluctua- tions are during periods of several days of continuous heavy cloud cover. Con- versely, the highest daily maximum tem- 20 xMISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 7. — The west shore of tlie lower lake at Santa Cecilia. 2 3 X < Q Z < Z) z >- _l X (- z o z < LlI 2 cr 3 < q: Ui Q. 2 35° 30' 25"- 20'- TEMPERATURE RAINFALL ■■600 -■500 JAN FEB MAR APR MAY — 1 — JUN — I 1 — JUL AUG -+- ■400 --300 3 O o < z o 2 < q: 200 SEP OCT NOV DEC Fig. 8. — Monthly mean minunum and maxinnnn ambient air temperatures and monthly rainfall accumulation at Santa Cecilia for tlie period July 1971-June 1972. EQUATORIAL HERPETOFAUNA 21 Table 2. — Summary of Climatic Data from Santa Cecilia. (July 1971- June 1972) Rainfall (mm) Cloud Cover ( Days ) Temperature ( ° C ) Montli Amount Jan 372.5 Feb 243.0 Mar 619.0 Apr 340.0 May 306.5 Jun 250.5 Jul 367.0 Aug 220.5 Sep 251.5 Oct 422.0 Nov 530.5 Dec 366.5 TOTAL 4289.5 Mean/Day Days Heavy Clear Maximum Minimum 12.0 22 8 8.4 17 10 20.0 25 12 11.3 21 21 10.0 25 14 8.4 21 9 11.8 23 7 7.1 20 24 8.4 21 19 13.6 25 13 17.7 21 16 11.8 17 3 215 156 7 4 9 3 3 7 3 4 4 4 6 15 69 24.5-30.0 24.0-31.5 23.0-35.0 23.0-32.0 23.0-31.0 23.0-31.0 24.0-32.5 22.0-31.0 23.0-33.0 23.5-32.0 23.0-32.0 24.0-32.5 22.0-35.0 19.0-23.0 18.5-23.0 19.0-23.0 18.0-23.0 20.0-23.0 18.5-23.0 20.0-23.0 18.0-22.0 18.5-22.0 18.0-22.0 20.0-23.0 19.0-22.0 18.0-23.0 peratiires and the greatest daily fluctua- tions are during periods of clear weather. Rainfall is abundant, but erratic; 4390 mm of rain were recorded in one year (Table 2). Local residents claim that normally August-September and Decem- ber-February are drier than other times and that April-May and October-Novem- ber have the heaviest rains. These obser- vations are at least partially substan- tiated by data from 1971-1972, although in 1972, March with 619 mm of rain was the wettest month. In the period 3-23 March 1972 there were 393 mm of rain; in the same period in 1967 there were only 150 mm. Although in April-May 1972 there were only 646.5 mm of rain, the contention of the local residents was borne out in 1973, when 1335.5 mm fell in 30 days (Table 3). December 1971 was a relatively dry month with only 234 lum of rain in the first 29 days, but 114 mm fell on December 30. The only other records of rainfall in excess of 100 mm in a 24 hr period were in 1973: 26 April (390 mm), 1 May (105 mm), and 8 May ( 165 mm ) . Among the rainfall records for 20 months at Santa Cecilia, there are six instances of rainfall on more than ten consecutive days: 15-27 June 1967, 130 mm (1-34/day); 10-20 July 196S, 171 mm (2-46/day); 6-19 October 1971, 209 mm (1-54/day); 29 December 1971-9 January 1972, 2S1 mm (1-114/ day); 8-21 May 1972, 166 mm (1-54/ day); 29 April-9 May 1973, 614 mm (2-165/day). The number of consecutive days with- out rain varied from one in May to five in December, although there were more days that were predominately clear. The greatest number of consecutive clear days was nine in December. The rainfall pattern usually consists of three or four rainy days with a maximum of 50-70 mm for the period and a maximum of 30-50 mm on any given day, followed by one or two days without rain. The general pattern does fluctuate, however, and sometimes there are 8-10 consecutive days of moderate to heavy cloud cover with only 1-5 mm of rain each day. Con- versely, it is possible to have several consecutive days of heavy rainfall, such as 29 April-5 May 1973,' in which the daily rainfall was 70, 19, 105, 29, 90, 42, Table 3. — Comparative Rainfall Data (mm) for Three Years. 10 June-9 July Year l967 l968 1972 Total 322.0 304.0 330.5 No. Days 23 22 21 Daily Mean 10.7 10.1 11.0 Daily Range 0-71.0 0-50.0 0-22.5 19 April-18 May Year 1969 1972 1973 Total 378.0 316.0 1335.5 No. Days .- 20 24 22 Daily Mean 12.6 10.5 46.6 Daily Range 0-45.0 0-53.5 0-390.0 22 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 9. — Aerial view of Santa Cecilia looking nortlieast. The Rio Aguarico is visible in the lower right corner. and 17 mm. This was followed by 2, 71, 165, and 4 mm, 3 days with no rain, and then 86, 38, and 46 mm. Generally, rain falls more frequently by day than at night. Rain was recorded on 215 days in the period July 1971-June 1972; of these, rain fell during the day and night on 47 days and only at night on 40 oc- casions. Sometimes rainfall is extremely localized during brief but heavy showers, whereas at times of extensive heavy cloud cover the entire region receives rain. Vegetation Although plant collections have been made for the Shaw Herbarium of the Missouri Botanical Garden in St. Louis and the herbarium at the Universidad Catolica in Quito, no published summary of the flora is available. Because the physiognomy determines the numbers and kinds of habitats available for ani- mals, the following commentary is con- cerned more with the structure of the vegetation than with the floristics. Santa Cecilia is near the western lim- its of the largest expanse of tropical rain- forest in the world. This biotope called the Hijlea by most South American biol- ogists is referred to locally as the selva. The region of Santa Cecilia is at a suffi- ciently high elevation to preclude the seasonal flooding of great expanses of forest. Thus, the hajiaJes of the lower elevations of the Amazon Basin, char- acteristic as far inland as Iquitos, Peru, are absent. Prior to 1965 we can assume that, with the exception of a few clearings along the river and scattered swamps, the entire region was covered with primary rain- forest (Fig. 9-10). This forest is best developed on level, well-drained ground. In such places the largest trees form a canopy 30-35 meters above the ground. Some stilt palms protrude through the canopy. The largest trees in the primary EQUATORIAL HERPETOFAUNA 23 Fig. 10. — Aerial view of Munozlandia. The Rio Aguarico is to the left. The large square building in the upper center is the laboratory. forest are scattered 30-50 meters apart. Most such trees have extensive but- tresses, that may extend 3 meters up the trunks, which may be up to 2 meters in diameter (Figs. 11-12). In some areas definite stratification can be found. A secondary stratum, when present, is 15- 20 meters above the ground. This usually consists of broad-leafed trees, with or without buttresses, and stilt palms. A third layer at 8-12 meters consists of smaller, broad-leafed trees and spiny palms. The ground cover consists of a great variety of broad-leafed herbs 10-25 cm high and small ferns. In the primary forest there is a deep (5-10 cm) and continuous mulch layer. At midday usu- ally no more than 10 percent of the ground cover receives sunlight. Lianas are numerous, and heavy growths of epi- phytes are present on the horizontal limbs of the large trees, although brome- liads are uncommon. The large, but- tressed trees forming the canopy have shallow, extensive root systems. Growth seems to be limited by the ability of the shallow soil to hold the towering trees. These forest giants frequently fall over, with the uprooting of the extensive root system creating broad shallow craters and the interwoven system of lianas tear- ing a large swath in the forest. Thus, Fig. 11. — Buttressed base of large tree in pri- mary forest. 24 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 12. — A trail through primary forest. Note the abundance of liana.s. EQUATORIAL HERPETOFAUNA 25 Fig. 13. — Secondary forest. The large-leafed trees are Cecropia. 26 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 14. — A forest-edge situation showing dense shrubby growth; in the foreground is a marsh. there are numerous natural clearings in the primary forest. Clearings, either natural or man- made, soon are overgrown by bushes, saplings, and the successional Cecropia. These trees attain heights of 10-12 me- ters. The secondary forest is character- ized by dense, bushy undergrowth, few lianas, epiphytes, and herbs, and little mulch (Fig. 13). At the edge of either primary or secondary forest, especially along the borders of man-made clearings, there is a dense growth of various suc- cessional plants, especially leafy vines, which form an impenetrable wall of veg- etation (Fig. 14). Stands of bamboo with individual stalks reaching heights of 15-18 meters are scattered through the forest, al- though bamboo is uncommon in climax primary forest (Fig. 15). Within both primary and secondary forest there are depressions that contain water at least intermittently. These swamps vary from small, shallow depres- FiG. 15. — Bamboo grove in secondary forest. EQUATORIAL HERPETOFAUNA 27 Fig. 16. — A forest swamp that contained water throughout most of the year. sions that are flooded only after heavy rains to large semi-permanent swamps (Fig. 16). The swamps in primary forest characteristically have numerous spiny palms, whereas the swamps in secondary forest usually support dense growths of Heliconia, the immense leaves of which attain lengths of 2 meters, exclusive of stems (Fig. 17). Man-made clearings are either culti- vated or maintained as clearings by man or grazing animals. Some maiz and pine- apple are grown, but pldtanos and ijtica are the most common crops. As a result of topographic irregularities, there are low-lying areas in clearings that contain water. The marshes in these open de- pressions contatn sawgrass, small Heli- conia, and various aroids (Fig. 18). Habitats Throughout the accounts of the species and in the ecological analysis, the data are organized by major types of habitat. These are defined as follows: Primary Forest. — Mature forest char- acterized by nearly continuous canopy, stratification of vegetation, and deep mulch layer. Secondary Forest. — Successional stages and partially lumbered primary forest. The cutting of the large trees re- sults in a physiognomy resembling that of inteiTnediate successional stages. Forest Edge. — The ecotone, usually resulting from human disturbance, be- tween forest and clearings. Bamboo. — Any of the stands of bam- boo in forest or cleared areas. Clearing. — Uncultivated clearings, usually man-made and characteristically supporting a variety of grasses. Ctdtivated Fields. — Any cleared areas bearing crops. Swamp. — Forest depressions contain- ing water either ephemerally or perma- nently. 28 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 17. — Dense growth of Heliconia in a shallow swamp. Fig. 18. — A forest-edge marsh with a thick growth of sawgrass. EQUATORIAL HERPETOFAUNA Water-filled depressions in 29 Marsh clearings. Lake. — This category includes only the two pennanent lakes in the area. River. — Only the two major water- ways (Rio Aguarico and Rio Conejo) are included in this category. Stream. — All flowing water not in- cluded in the fonner category. IDENTIFICATION OF THE SPECIES One of the most difficult problems that faces biologists working with the large Amazonian biota is the identifica- tion of the species. The following keys are offered as an aid to the identification of the species of amphibians and reptiles from the upper Rio Aguarico. They should be useful for the herpetofauna of the entire upper Amazon Basin in Ecua- dor, southern Colombia, and northern Peru. Identifications made by use of the keys should be checked against the de- scriptions given in the accounts of the species. The keys have been designed for use in the field with living or freshly pre- served specimens. Unless stated other- wise, colors pertain to living animals. All characters used in the keys are visible with no more magnification than that provided by a 10 X hand lens. The keys are satisfactory for specimens of all ages, except that some juvenile frogs will pre- sent difficulties. All character states, except the most obvious ones, are illustrated. Internal structures, such as dentition and hemi- penes, have been avoided. The most con- fusing scutellation character in snakes is the loreal, a scale on the side of the head between the nasal and the preocular (Fig. 41a). Some snakes lack either the loreal scale or the preocular scale. If the scale that is present is higher than long, it is the preocular (Fig. 41c), whereas, if it is longer than wide," it is the loreal (Fig. 43b). Uno de los problemas mas dfficiles que han enfrentado los biologos a traba- jar con la abundante biota amazonica ha sido la identificacion de las especies. Las siguiente claves se entregan como una ayuda para la identificacion de las especies de anfibios y reptiles de la re- gion del alto Rio Aguarico. Ellas de- berian ser igualmente utiles para la her- petofauna de toda la Hoya Amazonica tanto en Ecuador, el sur de Colombia, como en norte de Peru. Las identifica- ciones hechas a traves de estas claves deberian ser confrontadas con las de- scripciones dadas en los registros de es- pecies. Estas claves han sido disefiadas para su uso en el campo, trabajando con especimenes vivos o recientemente pre- servados. Si no menciona otra cosa, las colores corresponden a animales vivos. Todos los caracteres usados en las claves son visibles con no mas aumento que el dado por una lupa manual de lOX. Las claves con satisfactorias para especi- menes de toda edad, excepto para al- gunos sapos juveniles que presentaran un poco mas de dificultad. Todos los caracteres senalados, ex- cepto los mas obvios, son ilustrados. Las estructuras internas, tales como denti- cion y hemipenes, han sido evitados. De los caracteres de escutelacion el que mas confunde en las culcbras es la escama loreal, ubicada lateralmente en la cabeza entre las escamas nasal y la preocular (Fig. 41a). En algunas culcbras la loreal o la preocular estan ausentes; en tal caso, si la escama presente es mas alto que larga, es la preocular (Fig. 41c); si fuese mas larga que ancha, correponde a la loreal (Fig. 43b). 30 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Key to the Orders axd Suborders OF Amphibians and Reptiles 1. Skin lacking external scales; limbs, if present, lacking claws on digits _. 2 Skin covered with scales; limbs, if present, having claws on digits 4 2. Limbs absent Order Gymnophiona (p. 79) Limbs present 3 3. Tail present; hind limbs and fore- limbs equal in size Order Caudata (p. 80) Tail absent; hind limbs much longer than forelimbs . Order Anura (p. 82) 4. Body encased in a bony shell Order Testudines (p. 192) Body not encased in a bony shell ^ 5. Limbs present 6 Limbs absent 7 6. Nostrils dorsal; hind feet webbed; anal opening longitudinal Order Crocodilia (p. 194) Nostrils lateral; hind feet not webbed; anal opening transverse .___ Suborder Sauria (p. 195) 7. Eyelids present; body cylindrical with small rectangular scales around body; anal opening nearly teniiinal .— Suborder Amphisbaenia (p. 224) Eyelids absent; body usually not cy- lindrical; scales not rectangular and not equal in size around the body; tail present posterior to the anal opening Suborder Serpentcs (p. 224) Clave de Ordenes y Subordenes de Anfibios y Reptiles 1. Piel sin escamas externas; extremi- dades, cuando presentes, no poseen uiias en los dedos 2 Piel cubiertas de escamas; extremi- dades, cuando presentes, tienen unas en los dedos 4 2. Extremidades ausentes Orden Gymnophiona (p. 79) Extremidades presentes 3 3. Cola presente; extremidades poste- riores y anteriores de igual tamano Orden Caudata (p. 80) Cola ausente; extremidades poste- riores mas largas que las anteriores Orden Anura (p. 82) 4. Cuerpo encerrado en una concha osea Orden Testudines (p. 192) Cuerpo no encerrado en una concha osea 5 5. Extremidades presentes 6 Extremidades ausentes 7 6. Nostrilos dorsales; pies posteriores con membranas; ano con abertura longitudinal Orden CrocodiHa (p. 194) Nostrilos laterales; pies posteriores sin membranas; ano con abertura transversal Suborden Sauria (p. 195) 7. Parpados presentes; cuerpo cilin- drico con pequenas escamas rec- tangulares alrededor del cuerpo; ano con abertura casi tenninal .... Suborden Amphisbaenia (p. 224) Parpados ausentes; cuerpo usual- mente no cilindrico; escamas no rec- tangulares e iguales en tamano al- rededor del cuerpo; cola presente posterior a la abertura anal — - - Suborden Sei-pentes (p. 224) EQUATORIAL HERPETOFAUNA 31 Key to the Caecilians 1. Small species, less than 200 mm in length; head pink or white, distinct- ly different from dark gray body _ Microcaecilia alhiceps (p. 79) Larger species; head same color as body 2 2. Body slender, length more than 80 times diameter 3 Body thick, length less than 50 times diameter 4 Clave de los Cecilios 1. Especie pequefia, menores de 200 mm de largo; cabeza rosada o blan- ca, notoriamente diferente del cuer- po gris oscuro Microcaeclia alhiceps (p. 79) Especie mayor; cabeza no diferente en color al cuerpo 2 2. Cuerpo delgado, largo mayor que 80 veces su diametro 3 Cuerpo grueso, largo menor que 50 veces su diametro 4 3. Eye visible beneath thin layer of bone^ Oscaecilia hassleri (p. 80) Eye in bony socket, not covered by bone Caecilia disossea (p. 79) 4. Snout tapered; body unifonii dull bluish gray Caecilia tentaculata (p. 79) Snout bluntly rounded; body bluish black with narrow white rings Siphonops annulatus (p. 80) 3. Ojo visible bajo una delgada capa osea^ Oscaecilia hassleri (p. 80) Ojo en zoquete oseo, no cubierto por hueso Caecilia disossea (p. 79) 4. Hocico puntiagudo; cuerpo de color gris-azulado apagado uniforme- mente __ Caecilia tentaculata (p. 79) Hocico romo, redondeado; cuerpo negro-azulado con anillos delgados blancos Siphonops annulatus (p. 80) Key to the Salamanders 1. Belly dark with large, irregular pale blotches; snout- vent length to 60 mm -_._ Bolitoglossa equatoriana (p. 80) Belly dull gray to brown, unifonn or with minute pale flecks; snout-vent length to 40 mm Bolitoglossa peruviana (p. 80) Clave de las Salamandras 1. Vientre oscuro con manchas palidas grandes e irregulares; distancia hocico-ano de 60 mm Bolitoglossa equatoriana (p. 80) Vientre gris apagado a marron, uni- forme o con pequeiias pecas palidas; distancia hocico-ano de 40 mm Bolitoglossa peruviana (p. 80) Key to the Frogs 1. Fingers lacking expanded discs (Fig. 19c) 2 Clave de los Sapos 1. Dedos sin discos tenninales expan- didos (Fig. 19c) 2 1 In order to determine the nature of tlie eye, it is necessary to remove a flap of skin over the eye, which may be visible as a dark spot beneath the skin. 1 Para determinar la naturaleza del ojo es necesario remover un pedacito de piel sobre el; el ojo puede verse como una mancha debajo de la piel. 32 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 19. — Digits of fingers: A. Pipa pipa, B. Hyla p,co^rup]iicia, C. Lcptodactylus wugncri, D. Eleuthcrodactyhis diadcmatus, E. Dendwhates pictus; A-D ventral, E dorsal. Fingers with expanded discs or other terminal modifications (Fig. 19a-b, d-e) 26 2. Skin on belly smooth 3 Skin on belly granular 21 3. Tympanmii visible (Fig. 20b) 4 Tympanum concealed ( Fig. 20a ) 17 Dedos con discos expandidos u otra modificacion terminal ( Fig. 19a-b, d-e) 26 2. Piel del vientre suave 3 Piel del vientre granulosa 21 3. Timpano visible (Fig. 20b) 4 Timpano oculto (Fig. 20a) 17 ■rrpcieB f/ Fig. 20. — Lateral views of heads: A. Eleuthewdactylus acuminatus with tympanum concealed, B. Eleuthcrodactylus lacrimosus with tympanum (stippled) visible. 4. First finger shorter than second 5 First finger longer tlian or equal to second 7 5. Four toes on foot (Fig. 21a) Syncope antenori (p. 192) Five toes on foot ( I"ig. 21b) 6 4. Primer dedo anterior mas corto que el segundo 5 Primer dedo anterior miis largo que el segundo 7 5. Cuatro dedos en los pies ( Fig. 21a ) ____ Syncope antenori (p. 192) Cinco dedos en los pies (Fig. 21b) 6 EQUATORIAL HERPETOFAUNA 33 Fig. 21. — Ventral views of feet: A. Syncope antenori with four toes, B. Hamptophrync boliviana with five toes. 6. Dark brown anal patch; no dark tri- angular mark in occipital region; skin on dorsum smooth; snout broad- ly rounded Eleutherodactylus nigrovittatus (p. 96) No dark brown anal patch; dark tri- angular mark present in occipital region; skin on dorsum pustular; snout pointed Adenomera andreae (p. 83) 7. Outer toes webbed nearly to tips __ Rana palmipes (p. 186) Outer toes not or only basally webbed 8 8. Tubercles present on heels and eye- lids Edalorhina perezi (p. 86) Tubercles absent on heels and eye- lids 9 9. Parotoid glands present (Fig. 22) Bufo glaherritnus (p. 116) Parotoid glands absent 10 10. Toes with distinct lateral fringes (Fig. 19c) 11 Toes lacking lateral fringes 12 6. Mancha anal marron oscura; sin marcas triangulares oscuras en la region occipital; piel del dorso suave; hocico ampliamente redon- deado -_ Eleutherodactylus nigrovittatus ^ (p. 96) Sin mancha anal marron oscura; marcas triangulares presentes en la region occipital; piel del dorso pus- tulosa; hocico puntiagudo Adenomera andreae (p. 83) 7. Dedos posteriores externos con membranas hasta cerca de su ex- tremo Rana palmipes (p. 186) Dedos posteriores externos sin o solamente con membranas en su base 8 8. Tuberculos presentes en los talones y parpados Edalorhina perezi (p. 86) Tuberculos ausentes en los talones y parpados 9 9. Glandulas parotidas presentes (Fig. 22) Bufo glaherrimus (p. 116) Glandulas parotidas ausentes 10 10. Dedos posteriores con flecos late- rales (Fig. 19c) 11 Dedos posteriores sin flecos laterales 12 34 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 22. — Biifo glaberrimiis showing paratoid glands (stippled). 11. First finger longer than second; ven- 11. ter mottled .— Leptodactylus wagneri (p. 112) First and second fingers equal in length; throat and chest dark; rest of venter usually creamy white -._ Leptodactylus discodactijlus (p. 105) 12. Dorsum black with distinct cream or 12. tan dorsolateral stripes; large red spots on thighs Lithodytes lineatus (p. 114) Not as described 13 Primer dedo anterior mas largo que el segundo; vientre moteado __.. Leptodactylus ivagneri (p. 112) Primer y segundo dedos anteriores de igual longitud; garganta y pecho oscuros; el resto del vientre usual- mente bianco cremoso Leptodactylus discodactylus (p. 105) Dorso negro, con bandas dorsolat- erales distintas de color crema o marron palida; grandes manchas rojas en los muslos Lithodytes lineatus (p. 114) Diferente a lo descrito 13 13. Dorsum tuberculate; orange spot with black mottling in groin; venter mottled pale gray and black with median white stripe Physalaemus petersi (p. 115) Dorsum smooth; coloration not as described 14 13. Dorso tuberculado; mancha anaran- jada con pintas negras en la ingle; vientre moteado gris palido con linea medial blanca Physalaemus petersi (p. 115) Dorso suave; coloracion diferente a la descrita 14 14. Distinct cream or tan labial stripe 15 14. Franja labial resultante de color crema o marron claro 15 Lips barred 16 Labios con barras 16 EQUATORIAL HERPETOFAUNA 35 15. Venter uniform cream _ Leptodacfyhis imjstaceus (p. 107) Venter mottled cream and black .... __- Leptodactyhis rhodomi/stax : (p. Ill) 16. Venter uniform grayish brown _- Lepfodactylus stenoclema (p. 112) Venter mottled cream and black __._ — Leptodactyhis pentadactylus .... (p. 108) 17. Outer toe at least one-third webbed 18 Outer toe not, or only basally, webbed 19 18. Dorsum and flanks dark; venter pale gray with dark spots Chiasmocleis anatipes (p. 187) Dorsum pale, sharply delineated from darker flanks; venter dark with pale flecks Ctenophryne geayi (p. 190) 19. Pale dorsal color meeting dark ven- tral color on a distinct line _ Hamptophryne boliviana (p. 190) Pale dorsal color blending into ven- tral color 20 20. Throat, chest, belly, and ventral sur- faces of thighs white with black spots and mottling - Chiasmocleis ventrimaculata (p.l89) Throat and chest brown; belly and ventral surfaces of thighs white with black spots Chiasmocleis hassleri (p. 188) 21. Parotoid glands present (Fig. 22); outer toe more than one-third webbed 22 Parotoid glands absent; outer toe not, or only basally webbed 23 15. Vientre unifomiemente crema - Leptodactyhis mystacetis (p. 107) Vientre moteado de crema y negro Leptodactylus rhodomystax (p. Ill) 16. Vientre unifomiemente cafe-grisa- ceo -. Leptodactylus stenodema (p. 112) Vientre monteada de crema y negro Leptodactylus pentadactylus : (p. 108) 17. Dedo externo posterior al menos con membrana hasta un tercio de su lon- gitud 18 Dedo externo posterior sin o solo basalmente con membrana 19 18. Dorso y flancos oscuros; vientre gris palido con manchas oscuras Chiasmocleis anatipes (p. 187) Dorso palido, netamente delineado de los flancos oscuros; vientre oscuro con manchitas palidas Ctenophryne geayi (p. 190) 19. Dorso de color palido reuniendose con colores ventrales oscuros en una linea distinta Hamptophryne boliviana (p. 190) Dorso de color palido mezelandose con colores ventrales 20 20. Garganta, pecho, vientre, y caras ventrales de los muslos blancos con manchas y pintas negras Chiasmocleis ventrimaculata (p. 189) Garganta y pecho marrones; vientre y caras ventrales de los muslos blan- cos con manchas negras Chiasmocleis hassleri (p. 188) 21. Glandulas parotidas presentes (Fig. 22); dedo externo posterior con mas de un tercio con membrana 22 Glandulas parotidas ausentes; dedo externo posterior sin o solo basal- mente con membrana 23 36 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 22. First finger shorter than second; snout pointed; spinous lateral fold of skin Biifo typhonius (p. 118) First finger longer than second; snout rounded; no spinous lateral fold of skin .. Biifo marinus (p. 117) 23. First finger shorter than second ^ 24 First finger longer than second ... 25 24. Tympanum concealed (Fig. 20a); snout pointed; skin on dorsum spin- ous; venter reddish lavender ____ Dendroplin/niscus minutus : (p. 120) Tympanum visible ( Fig. 20b ) ; snout rounded; skin or dorsum with elongate ridges; venter grayish cream __- Eleutherodactylus sulcatus (p. 102) 25. Dermal "horn" on upper eyelid; head broad ( > 50% snout-vent length) Ceratophrys cornuta (p. 85) No dermal "horn" on upper eyelid; head narrow ( < 40% snout- vent length ) Ischnocnema quixensis (p. 104) 26. Fingers temiinating in star-shaped structures (Fig. 19a); body greatly depressed Pipa pipci (p. 82) Fingers terminating in round or truncate discs (Fig. 19b, d, e); body not greatly depressed 27 27. Discs truncate; pair of dermal scutes on dorsal surface of eacli disc (Fig. 19e) ..._ 28 Discs truncate or round; no dermal scutes on dorsal surface of each disc (Fig. 19b, d) - 33 22. Primer dedo anterior mas corto que el segundo; hocico puntiagudo; pliegue lateral de piel espinoso Bufo typhonius (p. 118) Primer dedo anterior mas largo que el segundo; hocico redondeado; sin pliegue lateral de piel espinoso Bufo marinus (p. 117) 23. Primer dedo anterior mas corto que el segundo 24 Primer dedo anterior mas largo que el segundo 25 24. Timpano oculto ( Fig. 20a ) ; hocico puntiagudo; piel del dorso espinosa; vientre lavanda-rojizo Dendrophryniscus minutus (p. 120) Timpano visible ( Fig. 20b ) ; hocico redondeado; piel del dorso con lo- mos alargados; vientre crema-grisa- ceo Eleutherodactylus sulcatus (p. 102) 25. "Cuerno" dermico en el parpado su- perior; cabeza ancha ( > 50% de la longitud hocico-ano) Ceratophrys cornuta (p. 85) Sin "cuerno" dermico en el parpado superior; cabeza mas estrecha ( < 40% de la longitud hocico-ano) Ischnocnema quixensis (p. 104) 26. Dedos terminados en estructuras con forma de estrellas ( Fig. 19a ) ; cuerpo fuertemente deprimido Pipa pipa (p. 82) Dedos terminados en discos redon- dos o truncados (Fig. 19b, d, e); cuerpo no fuertemente deprimido - 27 27. Discos truncados; un par de escudos dermales en la cara dorsal de cada disco (Fig. 19e) 28 Discos truncados o redondeados; sin escudos dermales en la cara dorsal de cada disco (Fig. 19e) 33 EQUATORIAL HERPETOFAUNA 37 2S. First finger shorter than second Colostethiis sauli (p. 123) First finger equal to or longer than second 29 28. Primer dedo anterior mas corto que el segundo Colostcthu.s sauli (p. 123) Primer dedo anterior mas largo que el segundo 29 29. Venter uniform white or pale yellow __ Colostethiis marchesianus (p. 121) Venter mottled black and blue or gray 30 29. Vientre uniformemente bianco o amarillo palido -_ Colostethiis marchesianus (p. 121) Vientre monteado negro y azul o gris 30 30. Dorsimi l)rown with or without black flecks 31 Dorsum red, or black with yellow lines 32 30. Dorso marron con o sin pintas ne- gras 31 Dorso rojo, o negro con lineas ama- rillas 32 31. Pale ventrolateral stripe present; bright yellow spot on dorsal surface of upper arm and another in groin .___ P]ujIIo])ates femoralis (p. 126) Pale ventrolateral stripe absent; bright yellow spot in groin and another on ventral surface of shank __- Dendrohates pictus pictus (p. 125) 31. Banda ventrolateral palida presente; mancha amarilla brillante en la cara dorsal del brazo superior y otra en la ingle PhyUobates femoralis (p. 126) Banda ventrolateral palida ausente; mancha amarilla brillante en la ingle y otra en la cara ventral de la canilla _ Dendrohates pictus pictus (p. 125) 32. Dorsum red Dendroljates parvuhis (p. 124) Dorsum l)lack with yellow lines — _ Dendroljates quinquevittatus (p. 125) 32. Dorso rojo Dendrohates parvuhis (p. 124) Dorso negro con lineas amarillas ____ __-_ Dendrohates quinquevittatus _.._ (p. 125) 33. Skin on belly smooth 34 Skin on belly granular 35 34. Throat dark gray with median white stripe; no broad black mark posterior to eye Eletitherodacti/his lanthanites : :-^--(p.93) Throat and belly uniform creamy white; broad black mark present posterior to eye Eleutlierodacti/kis conspicillatus : (p. 89) 33. Piel del vientre suave 34 Piel del vientre granulosa 35 34. Garganta gris oscura con banda blanca; sin ancha marca negra pos- terior al ojo .___ Eleutherodacti/his lanthanites ._ _ : (p. 93) Garganta y vientre uniformemente blanco-cremoso; amplia marca ne- gra posterior al ojo presente -___ Eleutherodactyhis conspicillatus (p. 89) 38 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 35. First finger Ioniser tlian second -.- 36 First finger shorter tlian second 39 35. Primer dedo anterior mas largo que el segundo 36 Primer dedo anterior mas corto que el segundo 39 36. Head triangular with flesh)- probos- cis - Hcnxiphrucius prohoscideus (p. 127) Head normal without fleshy probos- cis '. 37 36. Cabeza triangular con proboscis car- nosa Ilcinipliractus prohoscideus (p. 127) Cabeza nonnal sin proboscis carnosa 37 37. Snout long, sloping; dermal fringes on outer edges of foreann and foot; white warts on dorsum ., CenfroleneUa rcsplendens (p. 185) Snout short, rounded; no dermal fringes on limbs; pale flecks, if pres- ent, not elevated 38 37. Hocico largo, en declixe; flecos der- micos en el canto externo del ante- brazo y pie; \'errugas blancas en el dorso . Ccntrolenello resplendens (p. 185) Hocico corto, redondeado; sin flecos dermicos en las piernas; pintas palidas cuando presentes, no eleva- das 38 38. Dorsum dark green with small gold- en flecks; heart not \isible through ventral skin; bones green Centrolenella midas (p. 183) Dorsum pale green with diffuse greenish yellow spots; heart \isible through \entral skin; bones white .. .... Centrolenella niunozorum (p. 184) 38. Dorso \erde oscuro con pequeiias manchas de color oro; corazon no \'isible a traves de la pared ventral; huesos verdes Centrolenella midas (p. 183) Dorso verde palido con manchas di- fusas amarilla-verdosas; corazon \isible a traves de la pared ventral; huesos blancos Centrolenella niunozorum (p. 184) 39. First toe longer than second 40 First toe shorter than second 43 39. Primer dedo posterior mas largo que el segundo 40 Primer dedo posterior mas corto que el segundo 43 40. Flanks tan or orange with dark markings 41 Flanks green with or without cream, blue, or orange spots 42 41. Calcars absent (Fig. 23a); ventral side of head, flanks and \enter cream or tan with brown or black flecks ..., Phyllomedusa palliata (p. 176) 40. Flancos marron claros o anaranja- dos con manchas oscuras 41 Flancos verdes con o sin manchas cremas, azules, o anaranjadas 42 41. Calcares ausentes (Fig. 23a); cara ventral de la cabeza, flancos v vien- tre crema o marron claro con pecas marrones o negras Tlujllomedusa palliata (p. 176) EQUATORIAL HERPETOFAUNA 39 Fig. 23. — Dorsal views of heels showing presence and shape of calcars and tubercles: A. Hijla punctata, B. Hyla calcarata, C. Hijla garhei, D. Hijla jasciata. Calcars present (Fig. 23a); side of head green; flanks orange with verti- cal black bars ___- Phyllomedusa tomopterna - (P- 179) 42. Parotoid region round; no dorsolat- eral row of white tubercles; iris deep bronze with black reticulations Phyllomedusa tarsius (p. 177) Parotoid region angular with dorso- lateral row of white tubercles; iris pale green Phyllomedusa vaillanti (p. 180) Calcares presentes (Fig. 23b); lados de la cabeza verdes; flancos anaran- jados con barras verticales negras .. - Phyllomedusa tomopterna (p. 179) 42. Region parotidal redondeada sin fila dorsolateral de tuberculos blancos; iris bronce-oscuro con reticulaciones negras Phyllomedusa tarsius (p. 177) Region parotidal angulosa con fila dorsolateral de tuberculos blancos; iris gris palido Phyllomedusa vaillanti (p. 180) 43. Tympanum completely concealed (Fig. 20a) 44 At least ventral part of tympanum visible 46 43. Timpano completamente oculto (Fig. 20a) 44 Timpano visible al menos en su parte ventral 46 44. Dorsum green with black stripe from snout to eye Eleutherodactylus acuininatus (p. 86) Not as described 45 44. Dorso verde con banda negra desde el hocico al o|o .... Eleutherodactylus acuminatus .... (p. 86) Diferente a lo descrito 45 45. Small tubercle on heel (Fig. 23d); dorsum smooth Eleutherodactylus quaquaversus .... (p. 100) No tubercle on heel; dorsum rugose . Eleutherodactylus martiae (p. 94) 45. Pequeiio tuberculo sobre el talon (Fig. 23d); dorso suave _ . Eleutherodactylus quaquaversus (p. 100) Sin tuberculo sobre el talon; dorso rugoso _. Eleutlierodactylus martiae (p. 94) I 40 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 46. Outer toe not, or onl\' basally, webbed 47 Outer toe more than one-third webbed 55 47. One elongate tubercle on heel (Fig. 23c); and two tubercles on margin of upper eyelid Eleutherodactijhis orphnolaimiis : (p. 98) No tubercles on heel and eyelid 48 48. Bright yellow or orange spot in 49 grom No vellow or orange spot in groin ...J 50 49. Spot in groin deep yellow or orange; spots widely separated ventrally; dorsum rugose. Eleutherodacfi/his croceoing^uinis _— --(p-90) Spot in groin yellow, narrowly bor- dered by black; spots confluent, or nearly so, ventrally; dorsum smooth _— EleiitJicrochictijIus variabilis (p. 103) 50. Dorsum pale green, with or without cream dorsolateral stripe; discs on fingers round EleiitlierodactyJiis pseudoacumina- tus (p. 99) Dorsum not pale green; discs on fingers truncate . 51 46. Dedo externo posterior sin o con membrana solo en parte basal .... 47 Dedo externo posterior con mem- brana por mas que un tcrcio 55 47. Un tuberculo alargado sobre el talon (Fig. 23c); y dos tuberculos sobre el margen del parpado superior _.-. Eleutherodactijhis orphnolaimiis (p. 98) Sin tuberculos sobre el talon y el parpado 48 48. Mancha amarilla brillante o anaran- jada en la ingle 49 Sin mancha amarilla o anaranjada 1 50 i en la ingle 49. Mancha inguinal amarilla oscura o anaranjada; manchas ampliamente separadas ventralmente; dorso ru- goso Eleutherodactijlus croceoinguinis . .. (p. 90) Mancha inguinal amarilla, bordeada de negro; manchas juntandose o muy cercanas ventralmente; dorso suave Eleutherodactijlus variabilis (p. 103) 50. Dorso verde palido, con o sin banda dorsolateral crema; discos de los de- dos anteriores redondeados Eleutherodactijlus pseud oacuminatus (p. 99) Dorso no verde palido ; discos de los dedos anteriores truncados 51 51. Venter pale green with white flecks Eleutherodactijlus paululus ._._ ; (p. 98) Not as described 52 51. Vientre verde palido con pecas blan- cas Eleutherodactijlus paululus (p. 98) Diferente a lo descrito 52 52. Upper lips with dark vertical or diagonal bars; venter not pale yel- low 53 52. Labios superiores con barras verti- cales o diagonales oscuras; vientre no amarillo palido 53 EQUATORIAL HERPETOFAUNA 41 Upper lips not barred; venter im- maculate pale yellow Eleutherodactylus hicrimosus .. (p. 92) 53. Groin marked with bold vertical or diagonal dark marks separated by gray, blue, orange, or red areas _ _ 54 Groin unifomi brown Eleiitherodacfyliis ockendeni (P- 97) 54. Venter cream with brown reticula- tions Eleutherodactylus diadematus __.. (p-91) Venter brown or gray with pale flecks Eleutherodactylus altamazonicus (P- 87) 55. Outer finger not webbed 56 Outer finger at least one-fourth webbed 64 56. Tubercle or calcar present on heel (Fig. 23b-d) 57 No tubercles or calcars on heel ( Fig. 23a) 59 57. Tubercle on heel (Fig. 23c); snout pointed, with tubercle on tip. Hyla garbei (p. 144) Calcar on heel (Fig. 23b, d); snout rounded, without tubercle on tip .— 58 58. Venter uniform white; flanks and thighs white or pale blue with verti- cal black bars Hyla calcarata (p. 137) Venter white with small brown flecks; flanks and thighs white or pale blue with irregular black spots Hyla fasciata (p. 140) Labios superiores sin barras; vientre amarillo palido -__- Eleutherodactylus lacrbnosus : (p. 92) 53. Ingle marcada con notorias marcas verticales o diagonales separadas por areas grises, azules, rojas, o ana- ranjadas 54 Ingle uniformemente marron Eleutherodactylus ockendeni (p. 97) 54. Vientre crema con reticulaciones marrones ____ Eleutherodactylus diadematus .... (p. 91) Vientre marron o gris con pecas pa- lidas Eleutherodactylus altamazonicus . . ; (p. 87) 55. Dedo anterior externo sin membrana 56 Dedo anterior externo al menos con membrana hasta un cuarto de su longitud 64 56. Tuberculo o calcar presente sobre el talon (Fig. 23b-d) 57 Sin tuberculos o calcares sobre el talon (Fig. 23a) 59 57. Tuberculo sobre el talon (Fig. 23c); hocico puntiagudo, con proboscis sobre la punta Hyla garhei (p. 144) Calcar sobre el talon (Fig. 23b, d); hocico redondeado, sin proboscis sobre la punta 58 58. Vientre bianco uniforme; flancos y muslos blancos o azul-palidas con barras verticales negras Hyla calcarata (p. 137) Vientre bianco con pequenas pecas marrones; flancos y muslos blancos o azul-palidas con manchas irregu- lares negras _ Hyla fasciata (p. 140) 42 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 59. Skin co-ossified with roofing bones of skull; dorsum tan; other surfaces dark brown with large pale yellow spots on flanks and thighs Nyctiinantis rugiceps (p. 168) Not as described 60 59. Piel co-osificada con los heusos de la boveda craneal; dorso marron claro; otras superficies marron oscuras con grandes manchas amarillo-palidas sobre los flancos y muslos Nyctimantis rugiceps (p. 168) Difercnte a lo descrito 60 60. Throat and chest grayish brown with cream spots on chest Hijla lanciformis (p. 150) Throat and chest immaculate cream or yellow 61 60. Garganta y pecho marron-grisaceo con manchas cremas sobre el pecho Hyla lanciformis (p. 150) Garganta y pecho de color crema o amarillo immaculados 61 61. Groin and anterior and posterior sur- faces of thighs with black mottling enclosing yellow spots Hyhi rubra (p. 163) Groin and thighs not black with yel- low spots 62 61. Ingle y caras anteriores y posteriores de los muslos moteados de negro encerrando manchas amarillas Hyla rubra (p. 163) Ingle y muslos no negros con man- chas amarillas 62 62, Dorsum pale brown with small black flecks; iris metallic green with black reticulations Hyla alboguttata (p. 128) Not as described 63 62. Dorso marron palido con pequeiias pecas negras; iris verde metalico con reticulaciones negras Hyla alboguttata (p. 128) Difercnte a lo descrito 63 63. Dorsum tan or pale olive-green with longitudinal brown marks; iris silver with median horizontal red streak. Hyla cruentomma (p. 139) Dorsum pale green or yello\\'ish tan with dark flecks, interorbital bar, and pairs of longitudinal or diag- onal marks in scapular and sacral areas; iris bronze Hylafunerea (p. 143) 63. Dorso marron claro o verde-oliva palido con marcas longitudinales marrones; iris plateado con banda media horizontal roja Hyla cruentomma (p. 139) Dorso verde palido o marron ama- rillento con pecas oscuras; barras in- terorbitales, y pares de marcas longi- tudinales od diagonales en la esca- pula y areas sacrales; iris bronceado Hyla funerea (p. 143) 64. Calcar present on heel (Fig. 2.3b); pigmented reticulation present in lower eyelid (Fig. 24b) 65 No calcar on heel (Fig. 23a); no reticulation in lower eyelid (Fig. 24a) 66 64. Calcar prcsente sobre el talon ( Fig. 23b); parpado inferior con reticula- cion pigmentada ( Fig. 24b ) 65 Sin calcar sobre el talon (Fig. 23a); parpado inferior sin reticulacion (Fig. 24a) 66 EQUATORIAL HERPETOFAUNA 43 TKjed'fV Fig. 24. — Lateral views of heads: A. Hijla calcarata with unpigmented lower eyelid, B. Hijla geograpliica with reticulated lower eyelid. 65. Outer finger webbed to base of disc; projecting prepollex in males (Fig. 25a) Hyla boans (p. 131) Outer finger about three-fourths webbed; no projecting prepollex in males (Fig. 26b) Hyla geographica (p. 146) 65. Dedo anterior externo con mem- brana hasta la base de los discos; prepolex saliente en los machos (Fig. 25a) Hyla boans (p. 131) Dedo anterior externo con mem- brana hasta tres cuartos de su longi- tud; sin prepolex saliente en los machos (Fig. 25b) Hyla geographica (p. 146) Fig. 25. — Palmar views of hands: A.' Hyla hoans witli prepollical spine (stippled), B. Hijla geographica without prepollical spine. 66. Dorsum predominantly green, with or without red flecks or dorsolateral stripes 67 Not as described 70 66. Dorso predominantemente verde, con o sin pecas o bandas dorsolat- erales rojas 67 Diferente a lo descrito 70 44 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 67. Snout pointed and angular in profile, projecting beyond margin of lip; vo- cal sac huge, pleated, pectoral ( Fig. 26a) _......^ 68 Snout round, not projecting beyond margin of lip; vocal sac smooth, sub- gular (Fig. 26b) 69 67. Hocico puntiagudo y anguloso en i perfil, projectandose mas alia del margen labial; immenso saco vocal, rugoso, pectoral (Fig. 26a) 68 Hocico redondeado, no projectan- dose mas alia del margen labial; soco vocal suave, subgular (Fig. 26b) 69 Fig. 26. — Ventral and lateral views: A. Sphaenorhynchus eurhostus, B. Hyla punctata. 68. Dorsum uniform green; outer finger webbed to base of disc; snout-vent length to 40 mm in males .__. Sphaenorhijnchus eurhostus (P- 182) Dorsum green with narrow red dor- solateral lines; outer finger about one-half webbed; snout-vent length to 20 mm in males __.- Sphaenorhijnchus carneus (p. 181) 68. Dorso verde unifomie; dedo anterior externo con membrana hasta la base del disco; longitud hocico-ano de hasta 40 mm en machos Splwcnorhi/nchus eurhostus : (p. 182) Dorso verde con lineas delgadas dorsolaterales rojas; dedo anterior externo con membrana hasta su mitad; longitud hocico-ano de hasta 20 mm en machos _ .. Sphaenorhynchus careus (p. 181) 69. Dorsum uniform green or with trans- verse red marks; prepollical spine projecting in males Hyla granosa (p. 148) Dorsum green with red flecks and/ or dorsolateral stripes; no projecting prepollical spine Hyla punctata (p. 158) 69. Dorso verde uniforme o con marcas rojas transversales; espina prepolical saliente en machos Hyhi granosa (p. 148) Dorso verde con pecas y/o bandas dorsolaterales rojas; sin espina pre- polical saliente Hyla punctata (p. 158) 70. Outer finger at least two-thirds webbed 71 70. Dedo anterior externo con mem- brana hasta al menos dos tercios de su longitud 71 EQUATORIAL HERPETOFAUNA 45 Outer finger no more than one-half webbed 74 71. Belly orange or yellow with bold black mottling; webbing orange; demial fringes present on outer edges of forearm and foot Hyla mormorata (p. 153) Not as described 72, 72. Broad, vertical cream bar below eye, expanded into labial stripe in some females; skin on dorsum of males spinous, of females, smooth Osteocephahis taurinus (p. 172) No suborbital bar or labial stripe; skin on dorsum thick and glandular 73 73. Large black post-tympanic spot present; webbing red; large dark brown quadrangular dorsal blotch narrowly outlined with cream Phrynohyos coricea (p. 173) No black post-tympanic spot; web- bing tan; dorsal pattern variable, but usually not consisting of a large quadrangular blotch Phrynohyas venulosa (p. 174) 74. Dorsum tan or olive usually with narrow transverse dark markings; cream labial stripe usually expanded posteroventral to eye; dorsum in males bearing many small spinous tubercles Osteocephalus leprieurii (p. 171) Not as described '— 75 75. Dorsum mottled dark brown, green, and tan; lips at least faintly barred with dark brown or dark green; dor- sum in males bearing mixture of large and small tubercles -___ Osteocephalus buckleyi (p. 170) Dedo anterior externo con mem- brana hasta no mas alia de su mitad 74 71. Vientre anaranjado o amarillo con motas negras oscuras; membranas anaranjadas; cantos denriicos pre- sentes en los costados externos de los antebrazos y pies Hyla marmorata (p. 153) Diferente a lo descrito 72 72. Amplia barra vertical crema bajo eo ojo, expandiendose hacia una banda labial en algunas hcmbras; piel de los machos espinosa, de las hembras suave Osteocephalus taurinus (p. 172) Sin barra suborbital o banda labial; piel del dorso gruesa y glandular ._. 73 73. Gran mancha negra postimpanica presente; membranas rojas; gran mancha dorsal cuadrangular marron limitada por fino borde crema Phrynohyas coriacea (p. 173) Sin mancha negra postimpanica; membranas marrones claras; colores dorsales variables, usualmente sin una gran mancha cuadrangular Phrynohyas venulosa (p. 174) 74. Dorso marron claro u oliva, corrien- temente con marcas transversales oscuras; banda labial crema casi siempre expandida posteroventral- mente al ojo; dorso de los machos con pequefios tuberculos espinosos _-.. Osteocephalus leprieurii (p. 171) Diferente a lo descrito 75 75. Dorso moteado marron oscuro, verde, y marron claro; labios al menos suavemente pintados con bar- ras marrones oscuros o verdes os- curos; dorso de los machos con una mezcla de tuberculos grandes y pe- quefios -__. Osteocephalus buckleyi (p. 170) 46 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Not as described 76 Diferente a lo descrito 76 76. Dorsum cream or yellow with red- dish brown flecks and broad lateral stripe Hyla rhodopepJa (p. 159) Not as described 77 76. Dorso crema o amarillo con pecas y amplia banda lateral marron-rojiza Hyla rhodopepla (p. 159) Diferente a lo descrito 77 77. Thighs dark brown or black with one to three cream or yellow spots on dorsal surface 78 Thighs not so patterned 80 78. Venter gray and white with bright orange spot on proximal ventral sur- face of shank; one white suborbital bar Hyla parviceps (p. 156) Venter creamy white; no orange spot on shank; two white suborbital bars 79 77. Muslos marron-oscuros o negros, con una a tres manchas amarillas sobre la cara dorsal 78 Muslos con otro diseiio 80 78. Vientre gris y bianco con mancha anaranjado brillante sobre la parte proximal de la cara ventral de la canilla; una barra suborbital blanca Hyla parviceps (p. 156) Vientre blanco-cremoso; sin mancha anaranjada en la canilla; dos barras suborbitales blancas 79 79. Anterior surface of thigh unifomi dark brown Hyla hrevifrons (p. 135) Anterior surface of thigh dark brown with one large yellow spot ____ Hyla bokermanni (p. 133) 80. Webbing and hidden surfaces of thighs orange or red 81 Webbing and hidden surfaces of thighs brown or yellow 84 81. Webbing and hidden surfaces of thighs red (pink at night) Hyla trianguhim (p. 167) Webbing and hidden surfaces of thighs orange 82 82. Dorsum dark brown with a network of cream lines Hyla favosa (p. 142) Not as described 83 79. Cara anterior del muslo marron-os- cura uniforme Hyla hrevifrons (p. 135) Cara anterior del muslo marron- oscura con una mancha grande ama- rilla Hyla bokermanni (p. 133) 80. Membranas y caras escondidas de los muslos anaranjadas o rojas . 81 Membranas y caras escondidas de los muslos marrones o amarillas . 84 81. Membranas y caras escondidas de los muslos rojas (rosadas por la noche) __.. Hyla triangulum (p. 167) Membranas y caras escondidas de los muslos anaranjadas 82 82. Dorso marron oscuro con un enre- jado de lineas cremas Hyla favosa (p. 142) Diferente a lo descrito 83 EQUATORIAL HERPETOFAUNA 47 83. Dorsolateral light bands straight- edged, continuing onto and usually fusing on rump, enclosing hour- glass-shaped dark brown dorsal mark; dorsal surfaces of shanks pre- dominately pale with or without one narrow dark crossbar HyJa leucophyllata (p. 152) Dorsolateral light bands irregular, extending diagonally to point be- hind amis; dorsal surfaces of shanks brown with broad, irregular, pale crossbar Hyla sarayactiensis (p. 165) S3. Bandas dorsolaterales tenues con hordes rectos, continuandose y cor- rientemente fusionandose en la nal- ga, encerrando a una mancha mar- ron-oscura en fonna de reloj de arena; cara dorsal de la pierna pre- dominantemente palida con o sin una delgada barra cruzada oscura Hyla leucophyllata (p. 152) Bandas dorsolaterales no acentuadas irregulares, extendiendose diagonal- mente hasta detras de los brazos; cara dorsal de la pierna marron con una barra cruzada ancha, irregular, palida Hyla sarayacuensis (p. 165) 84. Distinct light spots or transverse lines above anus and on heels 85 No light spots or lines above anus and on heels 86 84. Manchas claras resaltantes o lineas tranversales por encima del ano y de los talones 85 Sin manchas claras o lineas por en- cima del ano y de los talones 86 85. Dorsum dark brown with light top of snout, dorsolateral stripes, and round spots — one on rump and each heel Hyla hifurca (p. 130) Dorsum tan with brown blotches narrowly outlined with cream; cream line above anus and on each heel Hyla mimifa (p. 155) 85. Dorso marron oscuro con la punta del hocico clara, bandas dorso-lat- erales, y manchas redondeadas — una sobre el anca y otra en cada talon _-__ Hyla hifurca (p. 130) Dorso marron claro con manchas ir- regulares marrones delimitadas por crema; linea crema por encima del ano y sobre cada talon Hyla minuta (p. 155) 86. Three white spots on upper lip; dor- sum tan with dark brown blotch or chevrons Hyla riveroi (p. 161) Upper lip unmarked; dorsum tan with dark brown Hecks or irregular lines Hyla rossallejii (p. 162) 86. Tres pintas blancas sobre el labio superior; dorso marron claro con mancha marron o en bandas diag- onales fomiando una V Hyla riveroi (p. 161) Labio superior sin marcas; dorso marron con pecas marrones oscuras o con lineas irregulares Hyla rossalleni (p. 162) 48 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Key to the Turtles Clave de la Tortugas 1. Neck folds laterally to lie along mar- gin of shell 2 Neck withdraws straight back into shell ^ 5 1. Cuello plcgado lateralmente fuerca de la concha 2 Cuello se retrac directamente dentro la concha 5 2. Head broad, flat, with fleshy probos- cis and many long barbels; carapace broad, depressed, with ridges of keeled scutes Chelus fimhriatus (p. 192) Not as described 3 2. Cabeza ancha, chata, con proboscis carnosa y muchas barbillas; cara- pacho ancho, deprimido, con las salientes de los escutelos quillados CJielus fim])riatns (p. 192) Diferente a lo descrito 3 3. Two rounded, longitudinal ridges on carapace; lateral edges of carapace upturned Platemys platycephah (p. 193) Not as described 4 Dos crestas longitudinales redon- deadas en el carapacho; margenes laterales del carapacho dispuestos hacia arriba Platemys platycephala (p. 193) Diferente a lo descrito 4 4. Margin of posterior plastral indenta- tion concave (Fig. 27a); plastron dark; throat dusky Mesoclemmys gibha (p. 192) Margin of posterior plastral indenta- tion convex ( Fig. 27b ) ; plastron pale with or without dark spots; throat cream with black spots Phrynops geoffroanus tuherosus (p. 193) 4. Margcn de la identacion plastral posterior concava (Fig. 27a); plas- tron oscuro; region gular terrosa Mesoclemmys gihJ)a (p. 192) Margen de la identacion plastral posterior convexa (Fig. 27b); plas- tron palido con o sin manchas as- curas; region gular crema con man- chas negras Phrynops geoffroanus tuherosus (p. 193) a \ / \ / b Fic. 27. — Posterior margins of plastra: A. Mesoclemmys gibba, B. Phrynops geojfroaniis. EQUATORIAL HERPETOFAUNA 49 5. Carapace high, domed; feet short, stubby ^ Geochelonia dcnficulafa (p. 194) Carapace rounded, with three longi- tudinal ridges; feet broad, webbed - Kinosfcrnon scorpioidcs (p. 194) 5. Carapacho alto, en forma de cupula; pies cortos, gruesos _. GeocJwIonia denticulafa (p. 000) Carapacho redondo, con tres crestas longitudinales; pies anchos palmea- dos _- Kinosternon scorpioides (p. 194) Key to the Crocodilians 1. A transverse bony ridge anterior to orbits; two or three transverse rows of occipital scales (Fig. 2Sa) Caiman crocodilus crocodihis (p. 194) No transverse bony ridge anterior to orbits; one transverse row of greatly enlarged, strongly keeled occipital scales (Fig. 28b) Paleosuchus trigonotus (p. 195) Clave de los Cocodrilos 1. Una cresta osea transversal anterior a las orbitas; dos o tres hileras trans- versales de escamas occipitales ( Fig. 28a ) .__. Caiman crocodihis crocodihis _ (p. 194) Sin cresta osea transversal anterior a las orbitas; una hilera transversal de escamas occipitales grandes fuer- temente guilladas (Fig. 28b) Paleosuchus trigonatus (p. 195) Fig. 28. — Dorsal views of head and neck: A. Caiman crocodilus, B. Falcosuchus trigonatus. Key to the Lizards and Amphisbaenians 1. Feet (at least forefeet) present; eye distinct 2 Limbs absent; eye reduced to an in- distinct spot covered with skin _^__ Amphishacna fuhginosa (p. 224) 2. Lower eyelid absent; dorsal scales small, granular, those on top of head same size as scales on middorsum of body 3 Lower eyelid present; scales on top of head not granular, differing from those on middorsum of body 5 Clave de los Lacartos y Anfisbaenas 1. Pies (al menos los anteriores) pre- sentes; ojo distinto 2 Sin extremidades; ojo reducido a una mancha indistinta cubierta de piel . Amphishacna fuhginosa (p. 224) 2. Parpado inferior ausente; escamas dorsales pequeiias, granulares, aquellas en la superficie de la cabeza del mismo tamaiio que aquellas en la mitad dorsal del cuerpo 3 Parpado inferior presente; escamas cefalicas no granulares difiriendo de aquellas en la mitad dorsal del cuerpo 5 50 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 3. Toes having greatly expanded ad- hesive pads in two rows (Fig. 29a) __ Thecadactyhis rapicauda (p. 197) Toes not having adhesive pads 4 3. Dedos con cojinetes adhesivos muy expandidos, dispuestos en dos hil- eras (Fig. 29a) . Thecadactyhis rapicauda (p. 197) Dedos sin cojinete adhesivos 4 Claws covered by scaly sheath ( Fig. 29b); scales on belly pointed pos- teriorly; nuchal collar absent; body dark brown; top of head tan; throat white with brown streaks; tail not banded .... Psetidogonotodes guianensis ........^ (p. 196) Claws not covered by scaly sheath (Fig. 29c); scales on belly broadly rounded posteriorly; narrow white nuchal collar usually present; fe- males brown with dark brown trans- verse marks or reticulations; males with orange head and black and white mottled body; tail usually banded tan and dark brown Gonatodes concmnatus (p. 195) Carras cubiertas por una vaina es- camosa (Fig. 29b); escamas ven- trales orientedas hacia atras; collar nucal ausente; cuerpo marron os- curo; parte superior de la cabeza bronceado; region gular blanca con ray as marrones; cola sin bandas ____ Fseudogonatodes guianensis (p. 196) Garras no cubiertas por una vaina escamosa (Fig. 29c); escamas ven- trales ampliamente redondeadas posteriormente; collar nucal bianco, angosto, generalmente presente; hembras marrones con marcas o re- ticulaciones transversales marron oscuras; machos con la cabeza ana- ranjada y cuerpo moteado de bianco y negro; cola generalmente con ban- das bronceadas y marron oscuras - Gonatodes concinnatus (p. 195) Fig. 29. — Ventral views of digits: A. Thccadactijhis rapicauda, B. Fseudogonatodes guinanen- sis, C. Gonatodes concinnatus, D. Anolis fuscoauratus, E. Plica umbra. EQUATORIAL HERPETOFAUNA 51 Top of head covered with granular or irregular flat scales, the largest usually arranged in semicircles be- tween the eyes (Fig. 30a); lacking median frontal plate between eyes; tongue fleshy, not extensible 6 Top of head with regular flat plates; median frontal plate between eyes (Fig. 30c); tongue extensible, bifur- cate 16 Superficie de la cabeza con escamas planas granulares o irregulares, las mas grandes usualmente dispuestas en semicirculos entre los ojos (Fig. 30a); sin placa media frontal entre los ojos; lengua carnosa, no extensi- ble 6 Superficie de la cabeza con placas chatas regulares; con placa frontal media entre los ojos (Fig. 30c); lengua bifurcada extensible 16 ^<^es -ft Fig. 30. — Dorsal views of heads: A. Anolis transversalis with smooth scales and enlarged supraorbital semicircular scales (stippled on right side), B. AnoJis trachijderma with keeled scales and small supraorbital semicircular scales, C. Kentropyx pclviccps with regular cephalic plates, including a median frontal plate ( stippled ) . 6. Tail having spiny scales arranged in transverse whorls (Fig. 31a, c) 7 Tail not having spiny scales ar- ranged in transverse whorls (Fig. 31b) 9 6. Cola con escamas espinosis dis- puestas en anillos transversales ( Fig. 31a, c) 7 Cola sin escamas espinosas dis- puestas en anillos transversales ( Fig. 31b) 9 Tail cylindrical, covered with rings of small scales; a middorsal crest of long, spiny scales (Fig. 32a) 8 Tail flattened in horizontal plane, short; all caudal scales large and spiny (Fig. 31c); body and tail black; head orange Uracentron flaviceps (p. 208) 7. Cola cilindrica, cubierta con anillos de escamas pequenas; una cresta media dorsal de escamas espinosas grandes (Fig. 32a) 8 Cola achatada en piano horizontal, corta; todas las escamas caudales grandes y espinosas (Fig. 31c); cuerpo y cola negros; cabeza anaran- jada -_ Uracentron flaviceps (p. 208) 52 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 31. — Dorsal views of proximal segments of tails: A. Emjalioides laticeps, B. Plica timhro, C. Uracentron flaviceps. Fig. 32. — Lateral views of A. Emjalioides laticeps with middorsal crest (stippled), B. Anolis transversalis with, dewlap (stippled). 8. A dorsolateral row of enlarged spiny scales on body Emjalioides cofanorum (p. 203) No dorsolateral row of enlarged scales Emjalioides laticeps festae (p. 204) 9. Toes dilated and flattened (Fig. 29d); gular dewlap present at least in males (Fig. 32b) 10 Toes cylindrical or laterally com- pressed (Fig. 29e); gular dewlap absent 15 8. Con hilera dorsolateral de escamas espinosas dilatadas en el cuerpo Emjalioides cofanorum (p. 203) Sin hilera dorsolateral de escamas espinosas dilatadas en el cuerpo Emjalioides laticeps festae (p. 204) 9. Dedos dilatados y aplanados (Fig. 29d); abanico gular presente, al menos en los machos (Fig. 32b) 10 Dedos cilindricos o lateralmente comprimidos (Fig. 29c); abanico gular ausente 15 EQUATORIAL HERPETOFAUNA 53 10. Supraorbital scales large, smooth (Fig. 30a); other dorsal head scales smooth, or only scales on snout keeled; ventral scales large, smooth or weakly keeled 11 Supraorbital scales small, strongly keeled (Fig. 30b), or median supra- orbitals enlarged, keeled; all other dorsal head scales keeled; ventrals keeled, smooth, or granular 13 10. Escamas supraorbitales grandes, li- sas (Fig. 30a); las otras escamas dorsales de la cabeza lisas o sola- mente quilladas en el hocico; es- camas ventrales grandes, lisas o li- geramente c^uilladas 11 Escamas supraorbitales pequenas, fuertemente quilladas (Fig. 30b), o las supraorbitales medianas dilata- das, quilladas; todas las otras es- camas dorsales de la cabeza quilla- das; ventrales quilladas, lisas o gran- ulares 13 11. All head scales smooth: dorsum green or tan with bold diagonal transverse dark brown lines or bars; dewlap pale yellow with black bars AnoUs tronsversalis (p. 202) Scales on tip of snout keeled; colora- tion not as described 12 11. Todas las escamas de la cabeza lisis; dorso verdoso o bronceado con lineas o barras diagonales transver- sales marron oscuras; abanico gular amarillo palido con barras negras _ Anolis transversaJis (p. 202) Escamas en la punta del hocico quil- ladas; coloracion diferente a lo des- crito 12 12. Ventral scales weakly keeled; dor- sum green (blue-gray in preserva- tive) with bluish white flecks; scales around eye yellow; dewlap orange- yellow with white scales Anolis punctatus botdengeri (p. 201) Venter scales smooth; dorsum tan or gray, mottled with brown; venter creamy tan; dewlap orange with red streaks AnoUs ortonii (p. 200) 13. Ventral scales large, keeled; dorsum brown with diagonal dark brown streaks and/or cream middorsal line; belly cream with brown streaks; dewlap red peripherally, blue with white scales medially Anolis chnjsohpis scijpheus (p. 198) Ventral scales smooth; coloration not as described 14 12. Escamas ventrales ligeramente quil- ladas; dorso verdoso (azul grisaceo en preservative) con puntos blanco- azulados; escamas alrededor del ojo amarillas; abanico gular anaranjado amarillento con escamas blancas ____ ____ Anolis punctatus boulengeri (p. 201) Escamas ventrales lisas; dorso bron- ceado o grisaceo, moteado de mar- ron; vientre cremoso bronceado; abanico gular anaranjado con rayas rojas Anolis ortonii (p. 200) 13. Escamas ventrales grandes quilla- das; dorso marron con rayas diag- onales marron oscuro y/o linea me- diodorsal crema; vientre crema con rayas marrones; abanico gular rojo en la periferia, azul con escamas blancas en el centre Anolis chrysolepis scijpheus (p. 198) Escamas ventrales lisas; coloracion diferente a lo descrito 14 54 MISCELLANEOUS PUBLICATION 14. Ventral scales smaller, nearly granu- lar; throat and belly white, heavily flecked with brown or gray; dewlap rose-pink with white scales and white margin Anolis fuscoauratus fuscoauratus . .. (p. 199) Ventral scales larger, flat; belly dark gray laterally, yellow or creamy white medially; chin dark with trans- verse yellow line; dewlap orange- red with black scales __i AnoUs trochyderma (p. 201) 15. Middorsal crest present; tail less than twice length of body -. Plica umhra ochrocoUari.s (p. 206) Middorsal crest absent; tail more than twice length of body Polychrus marmoratus (p. 207) MUSEUM OF NATURAL HISTORY 14. Escamas ventrales mas pequeiias, casi granulares; region gular y vien- tre blancos, profusamente moteado de marron o gris; abanico gular ro- sado con el margen y las escamas blancas . _ Anolis fuscoauratus fuscoauratus (p. 199) Escamas ventrales mas grandes, planas; vientre lateralmente gris os- curo, amarillo o bianco cremoso en el centro; menton oscuro con una linea transversal amarilla; abanico gular rojo-anaranjado con escamas negras Anolis tracJujdernui (p. 201) 15. Cresta media dorsal presente; cola menos de dos veces la longitud del cuerpo .. Plica umhra ochrocoUaris (p. 206) Cresta media dorsal ausente; cola mas de dos veces la longitud del cuerpo . Polychrus marmoratus (p. 207) 16. Dorsal scales heterogeneous; large keeled scales mixed with small gran- ular scales (Fig. 33a) 17 Dorsal scales homogeneous or nearly so 18 16. Escamas dorsales heterogeneas; es- camas grandes quilladas mezcladas con las escamas granulares pequeiias (Fig. 33a) 17 Escamas dorsales homogeneas o casi homogeneas 18 i mm rwi Fig. 33. — Dorsal views of body scutellation: A. Neiisticunis ecpleopus, B. Kentiopyx pelvi- ceps, C. Mabutju mahouya, D. Iphi.sa eJegans. EQUATORIAL HERPETOFAUNA 55 17. Small (total length less than 200 mm); median ventral scales smooth; snont pointed; black spot with light center above insertion of arm Neiisticurus ecpleopus (p. 218) Large ( total length to S50 mm ) ; me- dian ventral scales slightly keeled; snout blunt; no black spot above in- sertion of anil Dracaena guianensis (p. 214) 17. Animales pequeiios (longitud total menor de 200 mm ) ; escamas ven- trales medianas lisas; hocico puntia- gudo; mancha negra con el centro claro sobre la union del brazo Neiisticurus ecpleopus (p. 218) Animales grandes (longitud total hasta 850 mm); escamas ventrales medianas ligeramente quilladas; hocico redondo sin mancha negra sobre la union del brazo Dracaena guianensis (p. 214) 18. Dorsal scales granular, much smaller than ventrals (Fig. 33b) 19 Dorsal scales not granular, as large as, or only slightly smaller than ven- trals 21 18. Escamas dorsales granulares mas pe- queiias que las ventrales (Fig. 33b) 19 Escamas dorsales no granulares tan grandes como o solo ligeramente mas pequeiias que las ventrales _ 21 19. Ventral scales in more than 20 longi- tudinal rows; dorsal scales smooth Tupinamhis tequixin (p. 223) Ventral scales in fewer than 17 lon- gitudinal rows; dorsal granules small, rounded or keeled 20 19. Mas de 20 hileras de escamas ven- trales longitudinales; escamas dor- sales lisas Tupinamhis tequixin (p. 223) Menos de 17 hileras de escamas ven- trales longitudinales; granulos dor- sales pequenos, redondos o quillados 20 20. Ventral scales keeled Kentropijx pelviceps (p. 215) Ventral scales smooth _ „ Ameiva ameiva petersii (p. 210) 21. Hind limbs normal 22 Hind limbs reduced to small stubs ._ Bachia trinasale trinasale (p. 213) 20. Escamas ventrales quilladas Kentropijx pelviceps (p. 215) Escamas ventrales lisas _ .. Ameiva ameiva petersii (p. 210) 21. Extremidades posteriores normales 22 Extremidades posteriores reducidas a pequeiias yemas Bachia trisanale trisanale (p. 213) 22. Dorsal scales smooth 23 Dorsal scales keeled 24 22. Escamas dorsales lisas 23 Escamas dorsales quilladas 24 23. All dorsal body scales about equal in size (Fig. 33c) Mahuija mabouija (p. 208) 23. Todas las escamas dorsales del cuerpo casi del mismo tamaiio ( Fig. 33c) — ^ Mahuija mahoiii/a (p. 208) 56 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Dorsum with two longitudinal rows of greatly enlarged scales ( Fig. 33d) Iphisa elegans (p. 215) Dorso con dos hileras de escamas longitudinales considerablemente di- latadas (Fig. 33d) Iphisa elegans (p. 215) 24. Scales around body in distinct trans- verse rows (Fig. 34a) 25 Scales around body not in distinct transverse rows (Fig. 34b) 28 24. Escamas alrededor del cuerpo en hileras transversales regulares (Fig. 34a) 25 Escamas alrededor del cuerpo no en hileras transversales ( Fig. 34b ) ._ 28 25. Distinct lateral zone of granular scales between dorsals and ventrals (Fig. 34c) -.- 26 Dorsal scales meeting or nearly meeting ventrals; no more than two longitudinal rows of granules later- ally 27 25. Zona lateral distinta de escamas granulares entre las dorsales y ven- trales (Fig. 34c) 26 Escamas dorsales en contacto o casi en contacto con las ventrales; no mas de dos hileras longitudinales de granulos lateralmente 27 nei/es 'fZ Fig. 34. — Lateral views of midbody scutellation (anterior to left): A. Arthrosaum reticulata with scales in transverse rows, B. Lcposoma parictale with scales in diagonal rows, C. Priono- chictijhts arguhis with granular lateral scales. 26. Frontonasal single ( Fig. 35a ) ; Hanks uniform dark brown; white ventro- lateral stripe extending from lip to hind limb .. Prionodactijlus manicatus (p. 221) Frontonasal longitudinally divided ( Fig. 35b ) ; flanks brown with light- centered ocelli; ventrolateral stripe absent or indistinct — Prionochictylus argulus (p. 220) 26. Frontonasal entero (Fig. 35a); flan- cos uniformemente marron oscuro; cinta ventrolateral blanca extendien- dose desde el labio hasta la extremi- dad posterior _ Prionodactyhis manicatus ( p. 221 ) Frontonasal dividido longitudinal- mente (Fig. 35b); flancos marrones con ocelos mas claros en el centro; cinta ventrolateral ausente o indis- tinta Prionodactyhis argulus (p. 220) EQUATORIAL HERPETOFAUNA 57 Fig. 35. — Dorsal views of heads: A. Prionodac- tijlus manicatus with single frontonasal (stip- pled), B. Prioitodactylus ar^ultts with di\ided frontonasal ( stippled ) . Fig. 36. — Ventral views of chins showing rela- tive lengths of chinshields (stippled): A. Ar- throsaura reticulata, B. Ptychoglossus brevi- frontalis. 27. Three supraoculars; suture between second pair of chinshields much longer than suture between first pair (Fig. 36a); dorsal caudal scales weakly keeled, keels not extending posteriorly beyond margins of scales Arthrosaura reticulata (p. 213) Four supraoculars; suture between second pair of chinshields shorter than suture between first pair ( Fig. 36b); caudal scales moderately keeled, keels extending posteriorly beyond margins of scales ..__ Ptychoglossus brevifrontaUs (p. 222) 27. Tres supraoculares; sutura entre el segundo par de escudos mentales mucho mas grande que la sutura entre el primer par (Fig. 36a); es- camas dorsales caudales ligeramente quilladas, las quillas no sobrepasan los margenes posteriores de las es- camas Arthrosaura reticulata (p. 213) Cuatro supraoculares; sutura entre el segundo par de escudos mentales mas corto que la sutura entre el primer par ( Fig. 36b ) ; escamas cau- dales ligeramente quilladas, las quil- las sobrepasan los margenes poste- riores de las escamas Ptychoglossus brevifrontaUs (p. 222) 28. Ventral scales keeled 29 Ventral scales smooth Alopoglossus copii (p. 210) 28. Escamas ventrales quilladas _— 29 Escamas ventrales lisas Alopoglossus copii (p. 210) 29. Head scales having a few rounded ridges; venter cream, becoming black in adult males -._ Alopoglossus atriventris (p. 209) Head scales having many keels; ven- ter cream or orange Leposoma parietale (p. 217) 29. Escamas cefalicas con pocas crestas redondas; vientre crema, negro en los machos adultos __-_ Alopoglossus atriventris (p. 209) Escamas cefalicas con muchas quil- las; vientre crema o anaranjado Leposoma parietale (p. 219) 58 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Key to the Snakes Clave de las Culebras 1. A deep pit between nostril and eye (Fig. 37a) 2 No deep pit between nostril and eye (Fig. 37b) 5 1. Una fosa profunda entre el nostrilo y el ojo (Fig. 37a) 2 Sin fosa profunda entre el nostrilo y el ojo (Fig. 37b) 5 ^1U£B 'f3 Fig. 37. — Sides of heads: A. BotJirops atrox with facial pit (stippled) between eye and nostril, B. Clelia clelia with normal coKihrid scutellation, C. CoraUus caniniis with labial pits ( stip- pled), D. Anilius scijtale with eye beneath single scale. 2. All dorsal scales uniformly keeled (Fig. 38a); two rows of scales be- tween orbit and upper labials 3 Scales in middorsal rows having knobby keels ( Fig. 3Se ) ; four or five rows of scales between orbit and upper labials Lachesis muta muta (p. 265) 2. Todas las escamas dorsales uni- fonnemente quilladas (Fig. 38a); dos hileras de escamas entre la 6r- bita y las supralabiales 3 Escamas de las hileras media dor- sales con quillas nudosas ( Fig. 38e ) ; cuatro o cinco hileras de escamas entre la orbita y las supralabiales _ ^ Lachesis muta muta (p. 265) Dorsum green with cream or yellow stripe on first row of dorsal scales _.__ -_. Bothrops bilineatus smara(i,dimis ..-.__. (p. 264) Dorsum with brown or black blotches; no yellow stripe of first row of dorsal scales 4 3. Dorso verdoso con ray a crema o amarilla en la primera hilera de es- camas dorsales - . Bothrops bilineatus smaraf^dinus (p. 264) Dorso con manchas marrones o ne- gras sin raya amarilla en la primera hilera de las escamas dorsales 4 EQUATORIAL HERPETOFAUNA 59 Fig. 38. — Dorsal scales ot niidhody: A. BoiJirops atrox with uniformly keeled scales, B. Imantodes ccnchoa with enlarged middorsal row; C. Clelia clclia with smooth scales ( metliod of counting dorsal scales stippled), D. Chironius carinatus with scales in even-numbered oblique rows, E. Lachesis muta with knob-like middorsal scales. Dorsum greenish yellow with nar- row chocolate brown blotches; ven- ter dark brown with yellow flecks ____ Bothrops castelnaudi (p. 264) Dorsum tan, brown, or gray with large darker rhomboidal blotches; venter mottled cream and brown or black Bothrops tarox (p. 262) Dorsal scales in more than 40 rows at midbody; top of head covered with many small scales ( Fig. 39a ) ; large scales present or not on snout (Fig. 39b) 6 Dorsal scales in fewer than 30 rows at midbody; top of head covered with large plates (Fig. 39c) 10 6. Scales on top of snout small, same size as those on top of head (Fig. 39a); single median longitudinal dark stripe on top of head Boa constrictor constrictor (p. 225) 4. Dorso amarillo verdoso con manchas angostas marron chocolate; vientre marron oscuro con moteados ama- rillos Bothrops castelnaudi (p. 264) Dorso bronceado, marron o gris con grandes manchas romboidales os- curas; vientre moteado de crema y marron o negro Bothrops atrox (p. 262) 5. Mas de 40 hileras de escamas dor- sales en medio del cuerpo; parte su- perior de la cabeza cubierta con muchas escamas pequeiias ( Fig. 39a); escamas grandes presentes o no en el hocico ( Fig. 39b ) 6 Menos de 30 hileras de escamas dor- sales en medio del cuerpo; parte su- perior de la cabeza cubiarta con placas grandes (Fig. 39c) 10 6. Escamas sobre el hocico pequefias, del mismo tamaiio que aquellas so- bre la cabeza ( Fig. 39a) ; raya media longitudinal oscura sencilla sobre la cabeza Boa constrictor constrictor (p. 225) 60 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 39. — Tops of heads: A. Boa constrictor with uniformly small scales, B. Epicrates cenchria with small scales and enlarged intemasals (stippled), C. CicJia cJcIia with normal colubrid scutellation, D. Helicops au^iiJatus witli single internasal (stippled), E. Xenopliolis scalaris with single prefrontal (stippled). Scales on top of snout consisting minimally of one pair of enlarged internasals (Fig. 39b); dorsal head pattern not as described 7 Escamas sobre el hocico reducidas a un par de internasales grandes ( Fig. 39b); diseno cefalico dorsal difer- ente a lo descrito 7 7. No pits in labials; top of head dark brown with broad tan or orange postorbital stripe Eunectes murimis murinus (p. 227) Pits present in labials (Fig. 37c); dorsal head pattern not as described 8 7. Sin fosa en las labiales; sobre la ca- beza marron oscuro con raya post- orbital ancha bronceada o anaran- jada Eunectes murimis murinus (p. 227) Fosas presentes en las labiales ( Fig. 37c ) ; diseno cefalico dorsal no como el descrito 8 8. Upper labials entering orbit; paired internasals bordered posteriorly by large scales (Fig. 39b); one loreal; dorsal head pattern consisting of narrow median and postorbital dark stripes; dorsal body pattern consist- ing of black circles with brown cen- ters Epicrates cenchria cenchria (p. 226) Upper labials separated from orbit; paired internasals bordered poste- riorly by small scales; two or more loreals; coloration not as described 9 8. Supralabiales en contacto con la 6r- bita; internasales pareadas, bordea- das posteriormcnte por escamas grandes (Fig. 39b); una loreal; di- seiio cefalico dorsal compuesto de rayas medias angostas y postorbi- tales oscuras; diseiio dorsal del cuerpo con circulos negros con rayas marrones Epicrates cenchria cenchria (p. 226) Supralabiales separadas de la orbita; internasales pareadas, l)ordeadas postcriormente por escamas pe- quefias; dos o mas loreales; colora- cion diferente a la descrita 9 EQUATORIAL HERPETOFAUNA 61 9. Dorsal scales in fewer than 55 rows; two or three loreals; one preocular; nasals in contact medially; interna- sals large; pits absent in anterior upper labials; body brown or gray with dark brown or black saddles bordered l)y cream — CoraUiis enydris enydris (p. 226) Dorsal scales in more than 55 rows; four loreals; two or three preoculars; nasals broadly separated by small internasals; pits present in all upper labials; body red or yellow in juve- niles, green in adults, with narrow vertical white bars Corallus caninus (p. 225) 9. Menos de 55 hilcras de escamas dor- sales; dos o tres loreales; una pre- ocular; nasales en contacto en la linea media; internasales grandes; fosas ausentes en la parte anterior de las supralabiales; cuerpo marron o gris con dibujos en fomia de mon- tura de color marron oscuro o negro bordeadas de crema .... Corallus enydris enydris (p. 226) Mas de 55 hileras de escamas dor- sales; cuatro loreales; dos o tres pre- oculares; nasales separadas por in- ternasales pequenas; fosas presentes en todas las supralabiales; cuerpo rojo o amarillo en juveniles, verde en los adultos, con barras angostas verticales blancas Corallus caninus (p. 225) 10. Ventral scales much wider than long; eye bordered by at least four scales; minimally six dorsal head shields (Fig. 39c) 11 Ventral scales only slightly wider than long; eye contained in a single large scale ( Fig. 37d ) ; enlarged head shields consisting of small paired internasals, large paired pre- frontals, and small diamond-shaped frontal Aiiilius scytale scytale (p. 224) 11. Dorsal scales in even number of rows (Fig. 38d) 12 Dorsal scales in odd number of rows (Fig. 38c) 15 12. Dorsal scales in 12 rows anterior to midbody; anal plate divided (Fig. 40a) 13 Dorsal scales in 10 rows anterior to midbody; anal jilate single (Fig. 40b) 14 10. Escamas ventrales mucho mas an- chas que largas; ojo bordcado al menos por cuatro escamas; sies es- cudos sobre la cabeza como minimo (Fig. 39c) 11 Escamas ventrales solo ligeramente mas anchas que largas; ojo bordeado por una sola escama grande (Fig. 37d); escudos sobre la cabeza for- mados por las internasales pareadas pequefias, prefrontales pareadas grandes, y frontal pequefia^n forma del diamante Anilius scytale scytale (p. 224) 11. Escamas dorsales en hileras pares (Fig. 38d) 12 Escamas dorsales en hileras impares (Fig. 38c) 15 12. Doce hileras de escamas dorsales en la parte anterior al media cuerpo; placa anal dividida (Fig. 40a) ._ 13 Diez hileras de escamas dorsales en la parte anterior al media cuerpo; placa anal entera (Fig. 40b) 14 62 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 40. — Ventral views of anal regions: A. Lcptodeira annulata with divided anal plate (stippled) and divided subcaudals, B. Fscudohoa coronata with single anal plate (stippled) and single subcaudals. 13. Ventrals fewer than 155 Cliironius carinatiis (p. 231) Ventrals more than 155 Chiromns mulfiventris (p. 233) 13. Menos de 155 ventrales Cliironius carinatus (p. 231) Mas dc 155 ventrales Cliironius miiltivcntris (p. 233) 14. Paravertebral rows of scales keeled (weakly or not at all in juveniles); ventrals fewer than 155; caiidals more than 117; juveniles grayish tan with olive-brown transverse blotches; adults olive-brown; venter yellow anteriorly Cliironius fuscus (p. 232) Paravertebral rows of scales smooth; ventrals more than 150; caudals fewer than 118; juveniles green above and below; adults black or dark olive-brown with rust or orange spots; venter rust or orange-tan an- teriorly Cliironius scurrtihis (p. 234) 15. Some or all dorsal scales keeled (Fig. 38a) 16 All dorsal scales smooth (Fig. 38c) 21 14. Hileras de escamas paravertebrales quilladas (poco notorio en juve- niles); menos de 155 ventrales; mas de 117 caudales; juveniles, broncea- do-grisaceos con rayas transversales marron-oliva; vientre amarillo an- teriormente Cliironius fuscus (p. 232) Hileras de escamas paravertebrales lisas; mas de 150 ventrales; menos de 118 caudales; juveniles, verde arriba y abajo; adultos, negros o marron-oliva oscuro con manchas marron-rojizas o anaranjadas; vien- tre marron-rojizas anaranjado-bron- ceado antoriormente Cliironius scurrulus (p. 234) 15. Algunas o todas las escamas dorsales quilladas (Fig. 38a) 16 Todas las escamas dorsales lisas (Fig. 38c) 21 16. Internasals paired; nostrils directed laterally (Fig. 39c) 17 Internasals fused into single scale; nostrils directed dorsolatcrally (Fig. 39d) 20 16. Internasales pareadas; nostrilos ori- entados lateralmente (Fig. 39c) 17 Internasales fusionadas en una sola escama; nostrilos orientados dorsal- mente (Fig. 39d) 20 17. Dorsal scales in 17 rows at midbody 18 17. 17 hiknas de escamas dorsales a medio cuerpo 18 EQUATORIAL HERPETOFAUNA 63 Dorsal scales in 21 rows at midbody 19 18. Anal plate single (Fig. 40b) Dendrophidion de7idrophis (p. 236) Anal plate divided (Fig. 40a) Drymobius rhombifer (p. 240) 19. Dorsal scales in 12 to 15 rows one head length anterior to anus; loreal quadrangular; one preocular; head and body dull olive-tan; no contrast- ing pale band across head Pseustes sulphureus sulphureus (p. 255) Dorsal scales in 21 rows one head length anterior to anus; loreal elon- gate; no preocular; unifomi dark gray with pale band across head — . Ninia hudsoni (p. 250) 20. Dorsal scales in 19 rows at midbody Helicops anguJatus (p. 242) Dorsal scales in 21 rows at midbody Helicops peter si (p. 243) 21 hileras de escamas dorsales a medio cuerpo 19 18. Placa anal entera ( Fig. 40b ) Dendrophidion dendrophis (p. 236) Placa anal dividida (Fig. 40a) Drymobius rhombifer (p. 240) 19. De 12 a 15 hileras de escamas dor- sales a una distancia de una cabeza de largo anterior al ano; loreal cuad- rangular; una preocular; cabeza y cuello bronceado-oliva opaco; sin contraste de banda palida sobre la cabeza Pseustes sulphureus sulphureus (p. 255) 21 hileras de escamas dorsales a una distancia de una cabeza de largo anterior al ano; loreal alargada; sin preocular; gris oscuro unifomie con una banda palida sobre la cabeza . Ninia hudsoni (p. 250) 20. 19 hileras de escamas dorsales a medio cuerpo Helicops angulatus (p. 242) 21 hileras de escamas dorsales a medio cuerpo Helicops petersi (p. 243) 21. Vertebral scale row noticeably wider than other rows at midbody (Fig. 38b); body slender, laterally com- pressed; pupil vertically elliptical .— 22 Vertical scale row same width as others at midbody (Fig. 38c), or, if wider, body pattern not consisting of blotches; body not laterally com- pressed; pupil round 28 22. Dorsal scales in 13 rows at midbody 23 Dorsal scales in 15 or more rows at midbody 25 21. La hilera de escamas vertebrales no- tablemente mas ancha que las otras hileras a medio cuerpo (Fig. 38b); cuerpo delgado, lateralmente com- primido; pupila vertical eliptica - 22 La hilera de escamas vertebrales de igual anchura que las otras en medio del cuerpo (Fig. 38c); o si mas anchas el cuerpo, carece de man- chas; cuerpo no comprimido lateral- mente; pupila redonda 28 22. 13 hileras de escamas dorsales a medio cuerpo 23 15 o mas hileras de escamas dorsales a medio cuerpo 25 64 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 23. Loreal present (Fig. 41a) Dlpsas cateshyi (p. 237) Loreal absent (Fig. 41b-c) 24 23. Loreal presente (Fig. 41a) Dipsas cateshyi (p. 237) Loreal ausente (Fig. 41b, c) 24 Fig. 41. — Lateral views of heads showing loreal and preocular scales (stippled): A. Dipsas cateshyi, B. Dipsas indica, C. Dipsas pavonina. 24. Top of head brown with dark brown streaks; body tan or gray with dark brown blotches wider ventrally than dorsally; venter dark brown with cream ventrolateral spots Dipsas indica eciiadorensis (p. 238) Top of head black with white trans- verse line on snout; body brown with broad black blotches wider dorsally than ventrally; posterior venter uniform tan Dipsas pavonina (p. 239) 24. Parte superior de la cabeza marron con rayas marron-oscuras; cuerpo bronceado o gris con manchas mar- ron oscuras mas anchas en el vientre que en el dor so; vientre marron- oscuro con manchas ventrolaterales crema Dipsas indica eciiadorensis ^ (p. 238) Parte superior de la cabeza negra con linea transversal blanca sobre el hocico; cuerpo marron con manchas negras dilatadas mas anchas en el dorso que ventralmente; vientre uni- forme bronceado posteriormente — . Dipsas pavonina (p. 239) 25. Dorsal scales in 15 rows at mid- body hnantodes lentiferus (p. 245) Dorsal scales in 17 or more rows at midbody 26 26. Dorsal scales in 17 rows at midbody hnantodes cenchoa cenchoa (p. 244) Dorsal scales in 19 rows at midbody 27 25. 15 hileras de escamas dorsales a medio cuerpo hnantodes lentiferus (p. 245) 17 o mas hileras de escamas dorsales a medio cuerpo 26 26. 17 hileras de escamas dorsales a medio cuerpo hnantodes cenchoa cenchoa (p. 244) 19 hileras de escamas dorsales a medio cuerpo 27 EQUATORIAL HERPETOFAUNA 65 27. Anal plate divided (Fig. 40a); one anterior temporal (Fig. 42a); body blotches extending laterally to fourth or fifth row of scales ... Leptodeira annulata annulata (p. 247) Anal plate single (Fig. 40b); two anterior temporals ( Fig. 42b ) ; body pattern consisting of narrow dark crossbars extending laterally to first or second row of scales ____ Tripanurgos compressus (p. 258) 27. Placa anal dividida (Fig. 40a); una temporal anterior (Fig. 42a); man- chas corporales extendidas lateral- mente hasta la cuarta o quinta hilera de escamas Leptodeira anntdata anmdata - (p. 247) Placa anal entera ( Fig. 40b ) ; dos temporales anteriores (Fig. 42b); disefio del ceurpo formado por bar- ras transversales oscuras extendidas lateralmente hasta la primera o se- gunda hilera de escamas .... Tripanurgos compressus (p. 258) a b c Fig. 42. — Lateral views of heads showing number and position of anterior temporals (stip- pled): A. Leptodeira annulata, B. Tripanurgos compressus, C. Micrurus surinamensis. 28. Dorsal scales in 15 rows at midbody 29 Dorsal scales in 17 or more rows at midbody 40 29. Color pattern consisting of red and black rings completely encircling body, or at least evident ventrally; yellow or white rings present or not 30 Color pattern not consisting of red and black rings encircling body 35 28. 15 hileras de escamas dorsales a medio cuerpo 29 17 o mas hileras de escamas dorsales a medio cuerpo 40 29. Disefio de coloracion consiste de anillos rojos y negros circundando completamente el cuerpo, o al menos ventralmente evidentes; anil- los amarillos o blancos presentes o no 30 Diseiio de coloracion diferente a la descrita 35 30. Anal plate single (Fig. 40a); one pair of chinshields; one postocular (Fig. 43b) -_ Atractus elaps (p. 227) Anal plate divided (Fig. 40a); two pairs of chinshields; two postoculars (Fig. 43a) 31 30. Placa anal entera (Fig. 40b); un par de placas mentales; una postocular (Fig. 43b) Atractus elaps (p. 227) Placa anal dividida (Fig. 40a); dos pares de placas mentales; dos post- oculares (Fig. 43a) 31 66 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 43. — Lateral views of heads showing loreal (stippled) between nasal and preocular or orbit and number of postoculars (stippled): A. En/tlirolamprus aesculapii, B. Atractiis elaps, C. Siphlophis cervinus. 31. Two pairs of secondary temporals (Fig. 42a) Erytlirolamprus aesculapii aesculapii (p. 242) One pair of secondary temporals (Fig. 42c) 32 31. Dos pares de temporales secundarias (Fig. 42a) Erijthrolamprus aesculapii aesculapii (p. 242) Un par de temporales secundarias (Fig. 42c) 32 32. Head broad, depressed; nostrils di- rected dorsally; eye small; fourth upper labial barely entering orbit; frontal small; head shields black with reddish brown mark on each scale Micrurus surinamensis surinamensis (p. 262) Head not depressed; nostrils di- rected laterally; eye normal; fourth upper labial broadly entering orbit; frontal large; head pattern not as described 33 32. Cabeza ancha, deprimida; nostrilos orientadas dorsalmente; ojo pe- queho; cuarta supralabial escasa- mente entra en la orbita; frontal pequeiia; escudos de la cabeza ne- gros con una marca marron rojiza en cada escama Micrurus surinamensis surinamensis (p. 262) Cabeza no deprimida, nostrilos ori- entados lateralmente; ojo normal; cuarta supralabial penetra bien en la orbita; frontal grande; diseiios cefalicos diferentes a los descritos arriba 33 33. Black rings in triads ( red-black-yel- low-black-yellow-black-red ) 34 Black rings not in triads; 36-38 black rings, incompletely separated from red rings by white scales, on body Micrurus langsdorffi lan' with a creamy white to pale pink head. In some individuals the tip of the tail is pale gray. No other caecilian in the area has a contrastingly colored head. The length of the body is 40-48 times the diameter. S 235 mm; 9 206 mm. 80 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Occurrence. — One individual was found under a log in primary forest at Santa Cecilia; the large series from Lago Agrio was obtained by following bull- dozers clearing primary forest. Life History. — Seven of nine females collected from 29 April to 24 May con- tained small ova; the other two showed no evidence of reproductive acivity. Oscaecilia bassleri (Dunn) Caecilia bassleri Dunn, 1942, Bull. Mus. Comp. Zool., 91:518 [Type locality.— Rio Pastaza, Ecuador]. Oscaecilia bassleri — Taylor, 1968, Caecilians World: 600. Material. — Lago Agrio, 5. Identification. — This long, slender caecilian can be confused with Caecilia disossea; the length of the body is 90-140 times the diameter in O. bassleri. All other caecilians, except C. disossea, have much heavier bodies. In Oscaecilia the eye is visible under a thin layer of bone, whereas in Caecilia disossea the eye is in a socket and not covered by bone. The animal is uniform dull bluish black dor- sally and slightly paler below, i 864 mm; 9 774 mm. Occurrence. — All were obtained dur- ing clearing of primary forest by bull- dozers. Life Hi.stonj. — One of two females obtained in May contained small ova. Siphonops annulatus (Mikan) Caecilia annuata Mikan, 1820, Delectus Floriae Faiuia Brasil, pi. 11 [Type locality. — "Se- bastianopolis" ( =Rio de Janeiro), Brasil]. Sip]}onoi)s annulatus — Wagler, 1830, Isis von Oken, 21:742. Material. — Puerto Libre, 1. Identification. — This is a stout-bodied caecilian with a body length 32 times the diameter. The head and body are dark bluish black with narrow pale bluish white rings corresponding to the primary folds on the body. No other caecilian in the area is so colored, i 454 mm. Occurrence. — The specimen was in a trench in early morning; presumably it fell in during the night. Life History. — No data are available. SALAMANDERS PLETHODONTroAE Bolitoglossa equatoriana Brame and Wake Bolitoglossa equatoriana Brame and Wake, 1971, Contr. Sci. Los Angeles Co. Mus., 219:23 [Type locality. — Limoncocha, Provincia Napo, Ecuador]. Material. — Santa Cecilia, 3. Identification. — This is the largest sal- amander in the area. By comparison with B. peruviana, this species is not only larger but proportionately more robust. FurtheiTOore, the venter in B. equatori- ana is dark with large white spots, where- as the venter in B. peruviana is gray with small pale flecks. In B. equatoriana the dorsum is reddish brown; the proximal parts of the hind limbs are dull yellow. The venter is gray with white flecks clumped into irregular spots. The iris is bronze. $ 44+44 mm, 9 52+47 mm. Occurrence. — The three individuals were found at night on low vegetation in secondary forest. One is from August and two from September. Life History. — The single female ob- tained in September had small ovarian eggs. Food. — All specimens contained ants; two also contained small beetles, and one had a small wasp. Bemarks. — Our specimens confomi to the description given by Brame and Wake ( 1971 ) . Formerly the species was known only from Limoncocha, about 65 km SE of Santa Cecilia. Bolitoglossa peruviana (Boulenger) ( Figure 48 ) Spelcrpcs peruviana Boulenger, 1883, Ann. Mag. Nat. Hist., (5)12:165 [Type locality.— Mo- yobamba, Departamento San Martin, Peru]. Bolitoglossa peruviana — Taylor, 1944, Univ. Kansas Sci. Bull,, 30:219. EQUATORIAL HERPETOFAUNA 81 Fig. 48. — Bolifoglossa peruviana. Upper: ju- venile, 22 mm SVL (KU 124125); lower: ju- venile, 24 mm SVL (KU 124126). Material. — Puerto Libre, 100; Santa Cecilia, 19. Identification. — This small relatively slender salamander differs from the larg- er, more robust B. equatoriana by having scattered small pale flecks, instead of flecks clumped into spots, on the venter. In B. peruviana the dorsum is tan, with or without a median brown streak at night; by day the dorsum is reddish brown or dull tan with dark brown streaks or flecks and (in some individ- uals) a brown triangle on the head. The venter is gray with scattered sihery white to gold flecks. The iris is tan with an orange ring around the pupil. $ 32+27 mm, 9 43+38 mm. Occurrence. — All of the specimens from Puerto Libre were collected in July. Three specimens were obtained by day; two were in leaf litter in primary forest, and one was under a fallen palm frond in secondary forest. The others were found at night in secondary forest. Most of the salamanders were on leaves of low plants, principally ferns between 5 and 30 cm above the ground, but a few were found as high as 1 m above the ground, and two were at heights of more than 1.5 m. Five were in primary forest, one on a banana plant, and the rest in secondary forest. Life History. — Thirt\--eight females contained 6-13 (x=8.39) ovarian eggs; 32 of these females were obtained in July at Puerto Libre. The other six are from Santa Cecilia; one was obtained in each month (April-September). Of 20 addi- tional females taken in Jul\' at Puerto Libre, seven contained immatiu-e ovarian .eggs, and 13 showed no evidence of re- productive activity'. At Santa Cecilia ju- veniles ( < 25 mm SVL ) were found in June, July, and February; no salaman- ders were found in October-January. The presence of juveniles from different times of the year and of gra\'id females from every month in which females were found is indicative of a lengthy breeding season or perhaps acyclic breeding in this species — an hypothesis supported by the absence of size classes in the large series obtained at Puerto Libre in Julv (Fig. 49). The smallest gra\id females have snout-vent lengths of 30 mm and ovarian complements of 6, 8, and 9 eggs. The two largest females have snout- vent lengths of 37 and 39 mm; each has an ovarian complement of 10 eggs. The largest ovarian complement, 13, is in a female ha\'ing a snout-vent length of 33 mm. In the series of 38 gravid females, only a slight positive correlation r= 0.19) exists between ovarian complement and snout-vent length. Food. — The stomachs of 14 adults contained primarily ants, accounting for 172 of 190 prey items. Other prey includ- ed 13 beetle larvae, 1 beetle, 2 spiders, 1 mite, and 1 leaf hopper (CicadelHdae). Taxonomic Comments. — Two small species of Bolitoglossa are known from the Amazon Basin - B. peruviana Bou- lenger and B. alfamazonica Cope. Iden- tifications of series of specimens obtained in Ecuador and Brasil have been con- finned by David B. Wake. A series of B. altamazonica from the vicinity of Belem, Brasil, was compared with the series of B. peruviana from Puerto Libre, by Mar- 82 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY O c CD 13 cr (D Lj_ 12 10 8 6 4 Juv. D 9 m^: l\ > • • • • "• •■.", ,' H*M I. |¥M"! «!»•»•»' I^ijifi; 1 ■I'Xya:':-:-:' m* H4f i+H !^ WA ????: SS ft«* Fig. 49.— 18 20 22 24 26 28 30 32 34 36 38 Snout-vent Length (nnm) Frequency distribution of snout-vent lengths of 100 Bolitoglossa peruviana collected in July 1968 at Puerto Libre, Ecuador. tha L. Crump, who generously provided the results of her analysis ( Table 4 ) . Al- though average differences are noted in several morphometric characters, the principal structural differences between the samples are that B. peruviana is smaller with a proportionately shorter tail, larger eye, and fewer maxillary and vomerine teeth. Although the coloration of the two species is similar, B. altamazonica is more variable. The dorsal ground color is dark gray to reddish brown, and the venter is gray, flecked with white in many indi- viduals. In some specimens the dorsum is unicolor, whereas in others the dorsum is mottled or has a middorsal stripe, light- er or darker than the ground color. All individuals have a dark brown triangular mark with the base between the eyes and the apex posteriorly. Some individuals have white flecks on the limbs, flanks, and lateral surfaces of the tail. Brame and Wake ( 1971 ) recognized B. altamazonica and B. peruviana as the only members of an Amazonian species group of Bolitofijossa. The former is known from Colombia, Peru, Bolivia, and extreme western Brasil in the upper Amazon Basin and from the vicinity of Belem, Brasil, near the mouth of the Amazon. Bolitoglossa peruviana is known from one locality in Peru and three in Ecuador. The two species have not been found at the same locality ( see Brame and Wake, 1963, for distribution map). FBOGS PiPlDAE Pipa pipa (Linnaeus) Rana pipa Linnaeus, 1758, Syst. Nat., Stock- holm, Ed. 10, 1:210 [Type locality.— Suri- nam]. Pipa pipa — Barbour, 1923, Proc. New England Zool. Club, 9:3. Material. — Santa Cecilia, 8. Identification. — This aquatic frog cannot be confused with any other from the area. The body is depressed, and the EQUATORIAL HERPETOFAUNA 83 -a c c > CD S *-' a si ?> o Co o > o CD y c C CT! •S O 0) C rt J-T -C o U t « >T3 C ti C 03 ■■^ S 13 +1 TO i s O ■" O "73 o '^^ ^% c a O 11 O 03 I. H u i-i m < S CH- CQ o K Ot 2t- 05 U o o V o o o o o o o o V V V V ■^_ iq P p P P rH 00 oooooo—Jo +1 +1 +1 +1 +1 +1 +1 +1 00 in ^ p in ci t^ ;i ^ ^ M* CO <0 (M CO I I ' I I I I I ■-HcipioocooqiX) o o in i-i oq c4 ""^ CO c O CO •^ in CO C-I in oi c>i CO oi oi 111 o in t- o d ^ o o V o V i> o in i— I o o V V 02 -* p tM p I— ( I— ; in p >— I d d d d d ■-< >— i +1 +1 +1 +1 +1 +1 +1 +1 in I--; p Cl p p '^ 1>; CO CD CD c-i CO CO i-^ d TP CO CM ^ o d OT CO CO C-1 CO ^ I> CI CO ^ CO i> iri in ci ci CO CO in CO cq c^ CI ^ 05CD'— !■— (i-Hi— (Ot^ +1 II +1 +1 +1 +1 +1 +1 '-ioqinpt-;ini>:P d ci in cq o4 CO d ^' CO CO I— I 1— ( in o ■<* C head is broad, flattened, and triangular with a truncate snout and dermal folds at the corners of the large mouth. The eyes are minute and dorsal in position. The toes are fully webbed; the long, un- webbed fingers terminate in star-shaped structures. The dorsum is dull brown with, or without, small scattered darker brown spots; the venter is dull tan with creamy flecks and blotches. The skin on the dorsum is finely tuberculate. All specimens from Santa Cecilia are juve- niles; the largest has a snout-vent length of 78 mm. Adults in excess of 150 mm snout- vent length are known from other localities. Occurrence. — Pipa pipa was found only in pennanent ponds in the forest. Life History. — No observations were made on this species; eggs are carried on the back of the female and develop di- rectly into small frogs ( Rabb and Snedi- gar, 1960; Rabb and Rabb, 1960, 1963). Mating, Call. — The call is a series of clicks produced underwater by move- ment of the arytenoid bones within a bony laryngeal structure fonued by hy- oid elements (Rabb, 1960). Food. — A specimen 78 mm in length contained two fish - an Erythriniis erij- thrinus (Erythrinidae) 30 mm long and a somewhat smaller pimelodid catfish. Remarks. — Apparently this is the first report of the genus from Ecuador. Dunn (1948:11) noted its widespread occur- rence in the Amazon Basin and the ab- sence of records from Ecuador. Leptodactylidae Adenomera andreae (Miiller) ( FlGLTRE 50 ) Adenomera andreae Miiller, 1923, Zool. Anz., Leipzig, 57:40 [Type locality. — Peixeboi, Estado do Para, Brasil]. Heyer, 1973, Contr. Sci. Los Angeles Co. Mus., 251:29. Adenomera andreae — Heyer, 1974, Contr. Sci. Nat. Hist. Mus. Los Angeles Co., 253:4. Material. — Lago Agrio, 3; Libre, 32; Santa Cecilia, 43. Puerto 84 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 50. — Adenomera andreae, 9 , 26 mm SVL (KU 119323). Identification. — This small species differs from Leptodactyliis in the area by having the first and second fingers equal in length. It is most easily confused with small individuals of Leptodactylus disco- dactijlus and L. wap^ncri, both of which have the first finger longer than the sec- ond and have lateral fringes on the toes. Other species of Leptodactylus in the area have dorsolateral dermal folds, which are absent in Adenomera and in L. discodactylus and wafineri. However, elongate tubercular ridges are present laterally in Adenomera: in most individ- uals a prominent ridge extends diago- nally from the scapular region to the groin. The absence of expanded tips on the digits distinguishes Adenomera from most Eleiitherodactyhis, and the absence of webbing distinguishes it from Rana. Adenomera andreae superficially resem- bles Eleiitherodactyhis nigrovittatus, which has smooth skin dorsally, has a dark anal patch, and lacks an interorbital triangular mark. The dorsum is dull olive-brown to grayish brown with darker brown mark- ings consisting of an interorbital triangle with apex posteriorly; short, broad longi- tudinal bars behind tympanum, in groin, and posteriorly from scapular region; diagonal bar in groin; round spots on flanks; and tran.sverse bars on hind limbs. The upper lip is marked by two dark brown suborbital bars diverging from the eye. A short creamy white bar extends from the tympanum to the base of the arm, which is orange-tan. The posterior surfaces of the thighs are brown. The venter is creamy white with brown flecks peripherally on the chin and laterally on the belly and thighs in some individuals. The iris is dull bronze with black reticu- lations. Nuptial pads and spines are absent in breeding males, i 25 mm; 9 27 mm. Occurrence. — This small terrestrial frog is active by night and day through- out the year, although about 60 percent of the frogs were found at night. Ten individuals were hidden beneath logs by day, and two were resting on low vege- tation at night. Eighty percent of the frogs were in leaf litter in primary or secondary forest; only one individual was in water. Eleven individuals were found in clearings at night. Life History. — No males were heard calling. Females having mature ovarian eggs were found in February, March, May, July, August, and September; fe- males having immature ovarian eggs were found in February-May and July- October. Five or more females were found in each of three months; the per- centage of females having mature ovar- ian eggs in each of those months is May (62%, N=S), July (70%, N=20), August (40%, N=5). The data are insufficient to determine with certainty the seasonal na- ture of reproduction in this species. The species of Adenomera deposit terrestrial foam nests, in which the larvae complete their development (Heyer, 1973). A foam nest, presumably, pro- duced by A. andreae, was found beneath a log in primary forest in March 1967. Twelve females contained 6-16 (x=8.7) unpigmented eggs having a maximum diameter of 3.0 mm. Of the three small- est juveniles (snout-vent lengths of 13-14 mm), one is from June and two from July. Mating CaU. — The call is unknown. Food. — Of 11 specimens containing food, ants were present in five. Roaches EQUATORIAL HERPETOFAUNA 85 (Blattidae) and beetles were present in two stomachs, whereas each of the fol- lowing food items was present in only one individual: Dermaptera, Isoptera, Heniiptera, Homoptera, Chilopoda, Ara- nea, and Annelida. Taxonomic Comments. — Most of the specimens from the Santa Cecilia region were utilized by Heyer (1973) in his systematic analysis of the Lcptodoctyltis marmorata group. Heyer resurrected the name L. andreae for an Amazonian spe- cies which had been confused with L. JiylaedacfyJiis; Heyer (1974a) resurrect- ed the generic name Adenomera Fitz- inger for members of the Leptodocfyhis marmoratus group. Ceratophrys cornuta (Linnaeus) (Figure 51 ) Raua cornuta Linnaeus, 1758, Syst. Nat., Stock- holm, Ed. 10, 1:212 [Type locality.— "Vir- ginia"]. Ccmtophn/s cornuta — Peters, Monatsb. Akad. Wiss. Berlin, 1872:204. Material. — Santa Cecilia, l-j-l. Identification. — This large distinctive frog has a broad head ( > 50% of snout- vent length) and demial "horns" on the eyelids. The only other frog in the area with large demial "horns" on the eyelids is Hemiphractiis prohoscideiis, which has a fleshy proboscis and a depressed body, in marked contrast to the robust body of Ceratophrys. The dorsum is brown and tan. A broad middorsal tan stripe begins on the eyelids, narrows in the occipital region, and extends to the vent. The stripe is bordered by dark brown; dorsolaterally there is a complex design of brown and tan. The side of the head is grayish tan with diagonal brown marks. The limbs are brown with dark brown transverse bars. The posterior surfaces of the thighs are brown with indistinct yellow mot- tling. The posterior part of the flanks, anterior surfaces of thighs, and inner surfaces of shanks are brown mottled with yellow. The belly is mottled brown and gray. The throat and ventral sur- FiG. 51. — Ceratophrifs cornuta, S, 80 mm SVL (UMMZ 129287). faces of the hands and feet are dark brown. The iris is creamy bronze with a median horizontal red streak. The single specimen from Santa Cecilia (UMMZ 129287) is a male having a snout- vent length of 80 mm, but Ronn Altig has a living male from Santa Cecilia with a snout- vent length of 110 mm. Occurrence. — The single specimen was sitting on the ground in secondary forest on the night of 15 May 1969. Life History. — No data arc available other than the presence of tadpoles in a pond in May. Tadpoles. — The following description is based on a tadpole (KU 158500) in stage 35 having a body length of 19 mm and a total length of 56 mm. The body is elongately o\'oid, wider than deep, and widest just anterior to the eyes. The snout is round. The eyes are small, close together, and directed anterolaterally. The nostrils are directed laterally at a point about one-third of the distance be- tween the eyes and the tip of the snout. The opening of the sinistral spiracle is directed posterolaterally on the ventro- lateral surface of the body at about mid- length of the body. The cloacal tube is short and dextral. The caudal muscula- ture is moderately robust proximally and gradually tapers to a slender tip. The fins are equal in depth and deepest at the midlength of the tail, where each fin is slightly deeper than the caudal muscula- ture. The fins extend to the tip of the 86 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY pointed tail, and the dorsal fin does not extend onto the body. The tail comprises about 66 percent of the total length. The large mouth is anterior with expanded protruding lips, each bearing a single row of large pointed papillae. The beaks are massive; the lower beak has a medial spine that fits into a notch in the upper beak. There are nine upper and nine lower rows of denticles, plus some short rows lateral to the beaks. The three in- nermost rows of l)oth series are inter- rupted medially. The body is dull brown; the tail is tan with a brown stripe later- ally on the proximal one-third of the tail. The entire caudal musculature and dorsal fin bear fine brown reticulations; the ven- tral fin is unpigmented. Mating Call. — The call is unknown. Food. — The specimen contained an adult frog, Edalorhina perezi, one spider, and four orthoptcrans. Edalorhina perezi Jimenez de la Espada (Plate 1) Edalorhina perezi Jimenez de la Espada, 1871, J. Sci. Acad. Lisboa, 3:58 [Type locality. — hanks of the Rio Napo, Ecuador]. Material. — Lago Agrio, 3; Santa Ce- cilia, 5. Identification. — This frog with a de- pressed body, cluster of tubercles on the eyelids, tubercle on snout, dorsolateral dermal fold, and black and white belly is distinct from all other frogs in the area. Eleutherodactijlus orphnolaimus has tu- bercles on the eyelid but lacks dorsolat- eral demial folds; Ceratophnjs cormita and Hemiphractus proboscideus have single dermal "horns" on the eyelids but difi^cr in structure and coloration. Eda- lohrina perezi is unique in having a white venter with large black spots laterally. The skin on the dorsum bears 5-7 longitudinal tubercular folds or two or three transverse folds; diagonal tubercu- lar folds are present on the dorsal sur- faces of the limbs. The dorsum is brown with tan streaks, or tan with a pale green suff^usion on the snout, eyelids, flanks. knees, and heels. There is a dark brown interorbital bar and transverse bars on the limbs. The flanks and ventrolateral surfaces are black. A yellow-orange spot is present in the groin. The anterior and posterior surfaces of the thighs, ventral surfaces of shanks, and inner surfaces of the feet are black and yellow or orange. The throat and belly are white, except for large black areas laterally on the belly. The iris is bronze with radiating grav-brown bars. £ 33 mm; $ 36 mm. Occurrence. — All were found active by day amidst leaf litter on the ground in primary forest. The frogs moved by series of short hops and hid amongst dead leaves, where they were effectively camouflaged. Life Hi.story. — One female collected in May contained 69 ovarian eggs, a second showed no evidence of reproduc- tive activity. Three juveniles from April and May have snout-vent lengths of 16 and 17 mm. No tadpoles were found, nor was any breeding activity observed. Mating Call. — The call is unknown. Food. — Four individuals contained food - Diptera (2), Hymenoptera (1), Orthoptera ( 1 ) . Taxonomic Comments. — Dunn (1949) noted the variation in the presence or absence of prevomerine teeth and poly- morphism in dorsal tubercular folds (longitudinal or transverse). Of six adults, one has longitudinal folds and no teeth; two have longitudinal folds and teeth, and three have transverse folds and no teeth. The only specimens having streaked brown and tan dorsal surfaces are those having longitudinal folds. In other aspects of coloration, the morphs are identical. Eleutherodactylus acuminatus Shreve ( Figure 52 ) Eletithcrodaciijliis acuminatus Shreve, 1935. Occas. Papers, Boston Soc. Nat. Hist., 8: 217 [Type locality. — Canelos, Provincia Pastaza, Ecuador]. Material. — Lago Agrio, 3; Puerto Libre, 6; Santa Cecilia, 26. EQUATORIAL HERPETOFAUNA 87 Fig. 52. — Eleutherodactijlus acuminatus, 9 , 31 mmSVL (KU 148720). Identification. — This is one of three species of green Eleutherodactylus in the area; the others are E. p send o acuminatus and pauJulus, both of which are smaller species having an external t>-mpanum. The tympanum is not visible in E. acumi- natus, which has a black stripe from the snout through the eye and extending to a point above the arm. The only other species of Eleutherodactylus in the area lacking an external tympanum are E. croceoinguinis, martiae, and £. qua- quaversus, which are predominantly brown frogs lacking a black canthal stripe. Other small green frogs in the area are three species of Centrolenella, two species of Sphaenorhynchus, Hyla granosa and H. punctata; all of these have tympana and extensive webbing between the toes. Webbing is absent in E. acuminatus. The snout is pointed and projects be- yond the margin of the lip. The skin is smooth on the dorsum and granular on the belly; dorsal tubercles and folds are absent. The discs on the fingers are broadly expanded. The dorsum and the anterior and posterior surfaces of the thighs are pale green; the venter and flanks are white, and the throat is green- ish yellow. In some individuals a few small black spots are present dorsolater- ally on the anterior part of the flanks; these extend posteriorly from the end of the black stripe extending from the tip of the snout, through the eye, to a point above the axilla. The iris is pale bronze. S 21 mm; ? 31 mm. Occurrence. — With the exception of one individual found amidst leaf litter on the forest floor by day and one on the ground in a clearing at night, all indi- viduals were found in bushes and trees. Seven were in bromelaids by day; all others were sitting on leaves of bushes and trees at night. Most frogs were on leaves of bushes less than 1.5 m above the ground in secondary forest, but three were higher, and six were in primary forest. Life History. — zA.lthough adult fe- males were found in all months except February, March, November, and De- cember, females with mature ovarian eggs were found only in May (3), July ( 2 ) , and August ( 1 ) . The six gravid females were 27.0-31.0 (X=28.8) mm in snout-vent length and contained 12-21 (X=17.2) eggs, the largest of which was 2.4 mm in diameter. An amplectant pair was found on a bush on 23 May 1972. Possibly eggs arc deposited in brome- liads, where development is direct. The smallest juveniles ( 14-0-15.0 mm), found in June and July, probably are not recent hatchlings. The data are insufficient to determine the extent of the breeding season in E. acuminatus: I suspect that breeding occurs throughout a much long- er period than that suggested by the dates of collection of gravid females. Mating Call — The mating call is a short, high whistle repeated infrequently. Food. — Of 24 stomachs examined, 23 contained 165 ants; other food items in- cluded two small beetles and one small homopteran. Eleutherodactylus altamazonicus Barbour and Dunn ( Figure 53 ) Eleutherodactyhis altamazonicus Barbour and Dunn, 1921, Proc. Biol. Soc. Washington, 34:161 [Type locality. — probably Nauta, Departamento Loreto, Peru]. Material— L^go Agrio, 21; Puerto Libre, 53; Puerto Ore, 2; Santa CeciHa, 136. Identification. — This small species of 88 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 53. — EJeutherodactijhis altamazonicus, 9 , 26mmSVL (KU 110777). Eleuthcrochictijhis with a partially con- cealed tympanum has black and cream, bluish white, or rose pink mottling in the groin and on the posterior surfaces of the thighs. The only other species hav- ing similar coloration is the larger E. diodematus. The latter has bluish white or pale gray and diagonal black bars in the groin. The two species also can be distinguished by the shape of the snout (pointed tip in E. altamazonicus and round in E. diadematus), and ventral coloration (brown or gray with white flecks in E. altamazonicus and cream with brown reticulations in E. diadema- tus), textin-e of the skin on the dorsum (tubercular in E. altamazonicus and smooth with scattered pustules in E. diadematus, which has a prominent tym- panum ) . Other species having pale spots on the posterior surfaces of the thighs either have smooth skin on the venter (£. conspicillatus and E. lanthanites) or short unexpanded digits (E. sidcatus). Eleutherodactijlus altamazonicus super- ficially resembles another small species with tubercular skin on the dorsum (E. croceoinguinis) , which is readily distin- guished by the presence of a large deep yellow to orange spot in the groin. The snout is nearly truncate in dorsal view except for a projecting tip, which provides a pointed profile. The tips of the fingers are expanded about thrice the width of the digit and are truncate. An exterior tympanum is rarely present. The skin on the dorsum is tubercular, and that on the belly is granular. The dorsum varies from tan to reddish brown. Many individuals are tan to pale reddish brown dorsally with the side of the head and flanks dark brown. In others the dorsum is brown with a pale tan inter- orbital bar and W-shaped mark in the scapular region, each bordered posterior- ly by a corresponding dark brown mark. The lips are barred; canthal and post- orbital stripes are absent. The flanks are dark brown or tan or gray with broad diagonal dark brown marks. The dorsal surfaces of the limbs are brown with dark brown transverse bars; some individuals have pale tan heels. The inguinal region, proximal anterior surfaces of the thighs, and the posterior surfaces of the thighs are mottled or barred with l:)lack or dark brown and cream, bluish white, or rose- pink. The ventral surfaces are gray brown to dark brown with minute white flecks. The iris is reddish copper with a gray suffusion, c^ 22mm; 9 34 mm. Occurrence. — Two individuals were found on the ground at night in clear- ings, and one was on a banana plant at night. All others were in primary forest (9%) or secondary forest (91%). Of those in the forest, 15 were on the forest floor by day, and 147 were on vegetation at night; two of the latter were on tree trunks, and the others were on leaves less than 1.5 m above the ground. Life History. — Of 85 adult females, 44 were gravid; these were found throughout the year, except December thru February. Two non-gravid females were found in February. Twenty gravid females had snout-vent lengths of 27.0- 31.5 (X=28.1) mm and contained 12-25 (X=18.2) ovarian eggs, the largest of which had a diameter of 2.5 mm. Am- plectant pairs were found on low vegeta- tion at night from May through Novem- ber; four of the 13 pairs were found in August. Three pairs laid eggs in the laboratory on 25 and 26 August and 24 November 1971. The clutches consisted of 17, 19, and 20 unpigmented eggs hav- ing diameters of 3.1, 2.3, and 3.0 mm, respectively. None of the eggs hatched. The four smallest juveniles had snout- EQUATORIAL HERPETOFAUNA 89 vent lengths of 8.0-10.0 (X=9.0) mm and were found in July. Evidently re- production occurs throughout the year. Presumably the eggs are deposited amidst leaf litter or other debris on the forest floor. Mating Call — The mating call is a soft "cluck", repeated once or twice in succession. Food. — Twenty-six individuals had eaten a variet)^ of small arthropods, among which orthopterans were the most frequent prey (34.6% of stomachs), fol- lowed by ants (26.9%), beetles (15.4%), spiders (15.4%), dipteran larvae (11.5%), and beetle larvae ( 7.7% ) . Eleutherodactylus conspicillatus (Giinther) ( Figure 54 ) Htjlodes conspicillatus Giinther, 1859, Cat. Batr. Sal. Brit. Mus., London: 92 [T>ioe locality.— Andes of Ecuador]. Eleutherodactylus conspicillatus — Rivero, 1961, Bull. Mus. Comp. Zool., 126:63. Material. — Lago Agrio, 2; Puerto Libre, 19; Santa Cecilia, 100. Identification. — Eleutherodactylus conspicillatus is one of three species of Eleutherodactylus in the area that has smooth skin on the venter. This species and E. lanthanites have dorsolateral der- mal folds, and broadly expanded termi- nal discs on the fingers and toes. Eleu- therodactylus lanthanites has a gray throat with a median white streak, a pale canthal stripe bordered below by a brown stripe, and pale creamy tan flecks on brown ground color on the posterior surfaces of the thighs, whereas E. con- spicillatus has a creamy white throat, a distinct narrow creamy white canthal stripe with the side of the head black below the stripe, and bright orange or red flecks on a dark brown ground color on the posterior surfaces of the thighs. The only other species having smooth skin on the venter is E. nigrovittatus, which lacks dorsolateral folds and ex- panded discs. Three other species (E. altamazonicus, diadematus, and sulca- FiG. 54. — Eleutherodactylus conspicillatus, $ , 27mmSVL (KU 110784). tiis) have pale spots on the posterior sur- faces of the thighs, but in these species the spots are large, and the skin on the belly is granular. In coloration E. con- spicillatui superficially resembles the much la:ger Hyla lanciformis, which has granular ventral skin and extensive web- bing between the toes. The snout is narrow and rounded and projects slightly beyond the margin of the lip. The skin is finely granular on the dorsum and smooth on the venter; dorsal tubercules are absent, but narrow, low dorsolateral dennal folds are present. The discs on the fingers are expanded and round, about twice the width of the digit. An external tympanum is present. The dorsum and flanks vary from dull olive-green to pinkish tan with darker olive or brown markings, each usually narrowly outlined with cream. The markings consist of a broad interorbital bar and two or three chevrons on the back; the apex of each chevron is an- terior and the lateral ends tenninate on the flanks. The dorsal surfaces of the limbs are marked with diagonal dark bars about half as wide as the inter- spaces. The side of the head is black, and a black postorbital stripe angles ven- trally just posterior to the tympanum. The canthal stripe is creamy yellow to white and begins on the tip of the snout. In most individuals the narrow dorsolat- eral folds are cream. The groin and an- terior surfaces of the thighs are pinkish tan to orange; the posterior surfaces of the thighs are dark brown with orange or red flecks. The belly is white, with 90 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY or without gray flecks; the ventral sur- faces of the hmbs are pale gray. The iris is deep bronze with a median horizontal brown streak. $ 31 mm; 9 44 mm. Occurrence.— Eleufherodactylus con- spicillatus is a conspicuous forest inhabi- tant. Individuals are active on the forest floor by day and on the ground or low vegetation at night. The following anal- ysis is based on data on 107 individuals: 56 in primary forest, 51 in secondary forest; 42 on ground by day, 7 at night, and 58 on vegetation ( < 1.5 m ) at night. The frogs are active both night and day. During the day they are most active on the ground early in the morning or im- mediately after showers. At night they perch on leaves or branches; all seemed to be awake at night. Life History.— Only 12 gravid fe- males were found: April (3), May (2), July (5), August (1), September (1); non-gravid females were found in those months and in February, June, and No- vember. Gravid females had snout-vent lengths of 40.0-44.5 (X=:42.3) mm and contained 27-62 (X=42.8) ovarian eggs. The largest eggs had diameters of 3.5 mm. One juvenile with a snout-vent length of 13.0 mm was found in May, and three (10.0, 12.0 and 13.0 mm) were found in July. Although the data are in- conclusive, i suspect that E. conspicil- latus breeds throughout the year with the possible exception of periodic dry times. Presumably the eggs are depos- ited amidst leaf httcr on the forest floor. Mating Call.— The call consists of a series of low-pitched soft notes. Poor/.— Although several kinds of arthropods were present in 27 stomachs, orthopterans (74% of the stomachs) and spiders (26%) were the most abundant food items. Eleuthrodactylus croccoinguinis Lynch ( Figure .55 ) Eleidhcrodaciiihis croccoinguinis Lynch, 1968, J. Ilerp., 2:13.3 [Type locality.— Santa Ce- cilia, Provincia Napo, Ecuador]. Fig 55. — Eletithcrodactt/his croceoingitinis, 9, 20mmSVL (KU 123454). Materials.— La.go Agrio, 1; Puerto Libre, 24; Puerto Ore, 4; Santa Ceciha, 115. hlentification.-This small species of EleutherodactyJus differs from all other species in the area, except E. variabilis, in having a bright yellow spot in the groin and on the proximal part of the anterior surface of the thigh. In E. cro- ceoinguinis the spot is deep yellow to pale orange, and the spots do not ap- proach the midline, whereas in E. vari- abilis the spots are pale yellow, large, narrowly bordered with black, and con- fluent, or nearly so, midventrally. More- over, E. croceoinguinis has a more robust body and tubercular skin on the dorsum, and E. variabilis has a more slender body and finely granular skin on the dorsum. EJeutherodactyhis croceoinguinis super- ficially resembles some individuals of E. niartiae, which is readily distinguished from E. croceoinguinis by the absence of yellow inguinal spots. EJeutherodactyhis altamazonicus is like E. croceoinguinis in having tubercular skin dorsally, but the groin and posterior surfaces of the thighs are barred or molded with black or dull red. The snout is round in dorsal view and in profile. The tympanum is concealed. The skin on the dorsum is tubercular, and that on the venter is granular. The discs on the fingers are about thrice the width of the digits and are truncate. The dorsal ground color is tan or brown with EQUATORIAL HERPETOFAUNA 91 dark brown to olive-brown markings con- sisting of an irregukir interorbital bar, a transverse or shallow W-shaped mark on the scapular region, short transverse or diagonal marks on the body, and trans- verse bars on the limbs. Canthal and postorbital stripes are absent. A narrow middorsal cream or orange stripe is pres- ent in some individuals. Dark brown diagonal bars are present on the lips. The flanks are gray or tan. A large deep yel- low to orange spot in the inguinal region is confluent or not with another spot on the proximal anterior surface of the thigh. The venter is gray with white flecks. The iris is dull bronze with dense black reticulations. 6 16 mm; 9 23 mm. Occurrence. — This species is a forest dweller. Thirteen individuals were found on the forest floor by day, whereas 114 were on leaves and branches of bushes less than 1.5 m above the ground at night. Eleiitherodacfyhis croceoin^iiinis seems to be primarily nocturnal; all mating was observed at night. Those individuals found by day were inactive until disturbed. Life History. — One hundred and three gravid females were found through- out the year; only 12 non-gravid females were found - March (1), May (1), July (6), August (2), September (1), De- cember (1). Eighty-six gravid females had snout-vent lengths of 18.0-23.0 (X= 20.3) mm and contained 2-7 (X=4.8) ovarian eggs no larger than 2.8 mm in diameter. Eleven amplectant pairs were found in March (2), April (3), May (3), June (2), and October (1). Two pairs deposited clutches of 6 and 4 unpig- mented eggs in the laboratory on 25 Oc- tober 1971 and 22 March 1972. The eggs were 3.4 mm in diameter. Juveniles hav- ing snout- vent lengths of 12.0-13.0 mm were found in March, April, July, and August; these seem to be too large to be recent hatchlings. Breeding obviously occurs throughout the year. Eggs prob- ably are deposited on the ground. Mating Call. — The call consists of a short click-like sound. Food. — Identifiable food items were found in 24 stomachs. Spiders were the most common item (45.8% of the stom- achs), followed by orthopterans (41.6%), beetles (20.8%), and ants (16.7%). Eleutherodactylus diadematus (Jimenez de Espada) ( Figure 56 ) Hylodes diadematus Jimenez de la Espada, 1875, Vert. Viaj. Pacif. Verif. Batr., Lisboa: pi. 3, fig. 3 [Type locality. — Unknown]. ElciitherodactyJiis diadematus — Gorham, 1966, Das Tieireich, Berlin, 85:69. Material. — Lago Agrio, 17; Puerto Libre, 5; Santa Cecilia, 13. Identification. — This species of Eleu- therodactylus has diagonal dark brown or black markings on a yellowish tan to a bluish white ground color on the flanks and inguinal region. In this manner it resembles the smaller E. altamazonicus, a species differing from E. diadematus by having a pointed snout, tubercular skin on the dorsum, and dark brown ven- ter; E. diadematus has a round snout, smooth skin with scattered pustules on the dorsum, and a pale venter with dark reticulations. Some individuals of E. diadematus have orange flecks on the posterior surfaces of the thighs. Other species having flecks or spots on the pos- terior surfaces of the thighs either have no dorsal pustules ( E. conspicillatus and E. lanthanites) or have non-expanded tips of the fingers (E. sulcatus). The snout is round in dorsal view and profile; an external tympanum is present. The discs on the fingers are expanded to thrice the width of the digits and are truncate. The skin on the dorsum is smooth with scattered pustules, and that on the venter is granular. The dorsum is tan, gray, or rusty brown with darker brown markings consisting of longitud- inal marks extending posteriorly from the eyelids to the scapular region, an irregu- lar transverse mark in the scapular re- gion, and diagonal marks extending pos- terolaterally onto the flanks. The groin 92 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY > Fig. 56. — Elcutherodacttjlus diadematus, $ , 26 mm SVL; Cordillera del Due, Provincia Napo, Ecuador (KU 123764). and the proximal anterior and dorsal sur- faces of the highs are bluish white to yel- lowish tan with diagonal dark brown bars in the groin and transverse bars on the limbs. The posterior surfaces of the thighs are dark brown. Canthal and post- orbital stripes are absent; dark brown bars are present on the lips. The venter is creamy white with a greenish tint posteriorly; brown dashes or mottling are present on the throat and the belly. The iris is greenish bronze with a median horizontal red streak. S 28 mm; 9 45 mm. Occurrence. — EJeuthcrodactijliis dia- dematus is an arboreal species occurring in primary (60%) and secondary (40%) forests. One was found on the forest floor by day; 11 were obtained by day when trees were felled, and 20 were perched on bushes and branches at night. Life History. — Only nine adult fe- males are in the collection. Three of six found in May and two of three in July were gravid. These had snout-vent lengths of 40.0-45.0 (X=42.4) mm and contained 43-108 (X=61.6) ovarian eggs, the largest of which had a diameter of 3.0 mm. One amplectant pair was found on a bush in July. Two juveniles having snout-vent lengths of 13.0 and 14.0 mm were found in July and August. One pair deposited eggs in the laboratory on 9 May 1969; the 18 unpigmented eggs were 3.9 mm in diameter. Reproduction may be cyclic in this species. Because of its arboreal habits, eggs likely are to be deposited in bromeliads or amidst other epiphytes. Mating Call. — The call is unknown. Food. — Only 11 stomachs contained food. Roaches were by far the most com- mon food item, appearing in 36.4 percent of the stomachs. These were followed by beetles (27.3%), spiders (18.2%) and ants (18.2%). Eleutherodactylus lacrimosus (Jimenez de la Espada) ( Figure 57 ) Cycloccpliahts lacrimosus Jimenez de la Espada, 1S75, Vert. Viaj. Pacif. Verif. Batr., Lisboa: pi. 3, fi^. 5 [Type locality. — Unknown]. Eleutherodactylus lacrimosus — Gorham, Das Tierreich, Berlin, 85:80. Material — Lago Agrio, 2; Puerto Libre, 14; Santa Cecilia, 43. Identification. — This small species with a flat head and proportionately large eyes has rounded expanded discs on the digits and can be mistaken for a Hijla. The absence of webbing in E. lacrimosus immediately distinguishes it from any hylid with which it might be confused. Labial bars are absent in E. lacrimosus, actiniinatus, pseudoacumina- tus, and quac/uaversus. Both E. aciimi- natus and quacjuaversus have concealed tympana; E. lacrimosus has dorsal dark markings, which are absent in E. pseudo- acuminattis. The snout is round in dorsal view and truncate in profile. An external tym- panum is present. The terminal discs on the fingers are round and about twice the width of the digit. The skin on the dor- sum is smooth; that on the belly is granu- lar. The dorsum is yellow, tan, olive green, or reddish brown with darker brown markings consisting of an inter- orbital stripe and one or more marks on the body. In some individuals these con- sist of transverse bars or chevrons. In others there is a large, elongate, irregu- EQUATORIAL HERPETOFAUNA 93 Fig. 57. — Eleutherodactijlus lacrimostis, $ , 25 mm SVL: Mera, Provincia Pastaza, Ecuador - (KU 119513). lar, niiddorsal mark. Narrow canthal and postorbital .stripes are present or absent. Labial bars are absent. Faint transverse bars are evident on the limbs in some individuals. The flanks and anterior and posterior surfaces of the thighs are tan. Some individuals have the snout, rump, or heels creamy yellow, orange, or pale green. The venter is creamy yellow. The iris is bronze with a median horizontal red streak, which can diff^use to give the iris a copper color. S 23 mm; 9 29 mm. Occurrence. — With the exception of one individual on the forest floor by day, all specimens of E. lacrimostis were found on bushes and trees at night; only 4 percent were above 1.5 m. Fifteen in- dividuals were in primary forest; 37 were in secondary forest. Life History. — Gravid and non-gravid females were found only in Februar)' and April through September. Eleven females had snout-vent lengths of 20.0- 29.0 (X=23.5) mm and contained 7-16 (X=9.5) ovarian eggs. The largest eggs were 2.5 mm in diameter. Amplectant pairs were found on low vegetation in June (1) and September (2). One pair deposited seven eggs, 4.5 mm in diam- eter, in the laboratory on 23 September 1971. Reproductivity activity is evident through most of the year; thus, I suspect that E. locrimosus breeds throughout the year, with the possible exception of No- vember-January. Oviposition sites are unknown, but because of the arboreal habits, it is likely that E. lacrimostis de- posits eggs in bromeliads. Mating Call — The call consists of a single peep, repeated at intervals of a minute or more. Food. — Among 17 somachs contain- ing food, small beetles were the most common food item (41.2% of stomachs), followed by orthopterans (29.4%) and spiders (23.5%). Eleutherodactylus lanthanites Lynch ( Figure 58 ) Eleutherodactylus lanthanites Lynch, 1975, Contr. Sci. Los Angeles Co. Mus., 272:10 [Type locality. — Santa Cecilia, Provincia Napo, Ecuador]. Material. — Lago Agrio, 9; Puerto Libre, 50; Santa Cecilia, 286. Identification. — This is one of three species of EletitJierodactyhis in the area having smooth skin on the venter. This species and E. conspicillatiis have ex- panded terminal discs on the digits and dorsolateral dermal folds; E. conspicil- lattis, has a black face (side of head an- terior to orbit), a creamy white throat, and orange or red flecks on a dark brown ground color on the posterior surfaces of the thighs, whereas E. lanthanites has a pale canthal stripe bordered below by a narrow brown stripe and a pale brown face, a gray throat with a median white streak, and pale creamy tan flecks on brown ground color on the posterior sur- faces of the thighs. The only other spe- cies having smooth skin on the venter is E. nigrovittattis, which lacks dorsolateral folds and expanded discs. Three other species (E. altamazoniciis, diadematus, and sulcatus) have pale spots on the pos- terior surfaces of the thighs, but in these species the spots are large, and the skin on the belly is granular. The snout is narrowed anteriorly and rounded, barely projecting beyond the margin of the jaw. The skin is finely tu- berculate above and smooth below; dor- sal tubercules are absent, but narrow, low dorsolateral demial folds are present. The discs on the fingers are expanded, truncate, and about two and one-half times the width of the digit. An external 94 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 58. — Eleiithcrodact\ihis lanthanitcs, 9 , 39 mmSVL ( KU 146144). tympanum is present. The dorsal ground color is olive-tan, grayish-tan, or pinkish brown with darker brown markings. The markings are highly variable but usually consist of an interorbital bar, a chevron or X-shaped mark in the scapular region, and two or three chevrons or transverse marks on the posterior part of the body. There usually is a pair of small black spots in the scapular region. The limbs are marked with brown transverse bars that are slightly narrower than the inter- spaces. A narrow creamy white or yel- lowish tan canthal stripe usually is evi- dent; this is bordered below by a narrow dark brown stripe; a narrow dark brown or black postorbital stripe usually termi- nates above the tympanum. In most in- dividuals the lips are barred tan or gray and dark brown, but some have a creamy white labial stripe. The dorsolateral folds are yellowish or pinkish tan; the flanks are paler than the dorsum and marked with tan or olive-gray vertical or diag- onal marks, which may or may not be confluent with the dorsal markings. The groin and anterior surfaces of the thighs are pale orange to pale rose, whereas the posterior surfaces are brown with small, irregular yellowish tan ( nearly orange in some individuals) flecks. The throat is gray with white flecks and a broad me- dial white stripe; the belly is creamy white with gray flecks, and the ventral surfaces of the thighs are greenish cream with brown flecks. The iris is bronze, clear above and with radiating black streaks below and a median horizontal red streak. $ 29 mm; 9 46 mm. Occurrence. — Eleiitherodacfyhis lan- thanites is the most common and con- spicuous forest-dwelling Eleufherodacty- liis in the area and is about equally abun- dant in primary and secondary forest. Ninety-eight individuals were found on the forest floor by day; one was in a small stream by day. Three individuals were on the ground at night, and 171 were on vegetation at night; of these, only 11 were at heights of more than 1.5 m. This species is active on the ground by day and on low vegetation at night. Life History. — One hundred and five females containing mature ovarian eggs were found throughout the year, except September, when only two non-gravid individuals were found. Sixty-eight fe- males had snout-vent lengths of 34.0- 46.0 (X=38.7) mm and contained 20-52 (X=30.4) ovarian eggs, the largest of which had a diameter of 3.2 mm. One amplectant pair found on a bush on 2 April 1972 laid 30 eggs having diameters of 4.0 mm in the laboratory. The three smallest juveniles (10.0-11.0 mm in snout- vent length) were found in June and July. Certainly the species breeds throughout the year. Prol:)ably eggs are deposited on the ground as they are in other members of the Eleutherodactylus fitzingeri group. Mating Call. — The call is unknown. Food. — In 25 stomachs examined, orthopterans were present in 92 percent; beetles and spiders each were present in 20 percent. One fly, one ant, and one millipede completed the diets. Eleutherodactylus martiae Lynch ( Figure 59 ) Elcutherodactijlus martiae Lynch, 1974, Occas. Papers Mus. Nat. Hist. Ihiiv. Kansas, 31:2 [Type locality. — Santa Cecilia, Piovincia Napo, Ecuador]. Material. — Lago Agrio, 7; Puerto Libre, 51; Santa Cecilia, 57. Identification. — This species is like E. acuminatus, E. croceoinguinis, and E. quaquaversus in not having a visible EQUATORIAL HERPETOFAUNA 95 Fig. 59. — Eleittherodactyhis niartiae, amplec- tant pair, S, 18 mm SVL (KU 123815); 9, 23mmSVL (KU 123816). tympanum. Among these species, E. actiminatus is readily distinguished by its unifonn green dorsum and bold black canthal stripe. Eleutherodactyliis mar- tiae has scattered pustules on the dor- sum and low dorsolateral dennal folds; the lips are barred, and tubercles are absent on the heels. In contrast, E. qua- quaversus has a smooth dorsum without dorsolateral folds, no bars on the lips, and a small, pointed tubercle on each heel. EJeutherodactyJus croceoinguinis has tubercular skin on the dorsum and bold yellow or orange spots in the groin. Eleutherodactylus martiae superficially resembles E. ockendeni and E. pauhdus, both of which have an external tym- panum; E. ockendeni usually has an H- or W-shaped arrangement of dennal ridges in the scapular region, and E. paididus has smooth skin without pus- tules on the dorsum. The snout is acutely rounded in dor- sal view and round in profile. The discs on the fingers are expanded, about twice the width of the digit, and broadly rounded. The skin on the belly is granu- lar. The dorsum is dark brown with ir- regular transverse or longitudinal tan marks. In some individuals the heels are tan. The dorsal markings are highly vari- able. Although most individuals have a dark brown interorbital bar, in some it is tan, and others lack an interorbital mark- ing. Narrow dark brown bars are present on the limbs. The groin and the anterior and posterior surfaces of the thighs are pale orange to tan, with or without faint brown bars. A narrow, diflFuse, dark brown canthal stripe is evident in some individuals. Dark brown bars are pres- ent on the lips, and a narrow black post- orbital stripe expends posteroventrally from the eye. The ventral surfaces are gray or graying brown with small white to pale orange flecks. The iris is bronze with a median horizontal brown streak. S 21 mm; 9 25 mm. Occurrence. — Eleutherodactylus mar- tiae is a nocturnal forest inhabitant that seeks diurnal shelter amidst leaf litter. All individuals were in the forest - 24 in primary forest and 79 in secondary for- est. Seven were in leaf litter on the forest floor by day, and 96 were on low vegetation (< 1.5 mm) at night. Life History. — Females containing mature ovarian eggs were found in every month, except April and December, months in which no females were found. Eighteen females having snout-vent lengths of 19.0-23.0 (X=21.7) mm con- tained 5-13 (X=7.8) ovarian eggs, the largest of which was 2.0 mm in diameter. Ten amplectant pairs were found on low vegetation in seven months (January, March, May- August, November). Four pairs deposited eggs in the laboratory; clutches contained 8, 8, 9, and 10 unpig- mented eggs having diameters of 3.2, 3.5, 3.2, and 3.7 mm, respectively. Two of eight eggs deposited on 4 June hatched on 30 June; hatchlings had snout-vent lengths of 5.0 mm. This species obvious- ly breeds throughout the year. Presum- ably the egg clutches are placed amidst leaf litter on the forest floor. Mating Call— The call consists of a series of short clicks. Food. — A diversity of small arthro- pods was present in 25 stomachs. Orthop- terans were present in 48.0 percent of the stomachs, followed by beetles (16.0%), flies (16.0%), homopterans (12.0%), and lepidopteran larvae (12.0%). 96 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 60. — Eleutherodactylus ni^rovittatus, 5 , 26mmSVL (KU 123503). Eleutherodactylus nigrovittatus Andersson ( Figure 60 ) Eleutherodactylus ni^iovittafus Andersson, 1945, Ark. Zool., 37A(2):42 [Type local- ity.— Abitagua, Provincia Pastaza, Ecuador]. Material. — Lago Agrio, 1; Puerto Libre, 5. Identification. — This is one of three species of EJcutlicrodactyhis in the area having smooth skin on the venter. The other two species (E. conspicillatus and E. Janthanites) have dorsolateral dermal folds and long fingers with expanded terminal discs, whereas E. nigrovittatus has short fingers without expanded ter- minal discs and lacks dorsolateral folds. The only other species of Eleutherodac- tylus having short fingers without ex- panded discs is E. sulcatus, a larger spe- cies with a broad head ( > 50% of snout- vent length), numerous longitudinal ridges on the dorsum, tubercles on the eyelids, and granular skin on the venter. Eleutherodactylus nigrovittatus can be easily confused with Adenomera an- dreae; the latter differs from E. nigro- vittatus by having pustular skin on the dorsum, more pointed snout in dorsal view and profile, absence of a dark anal patch, and presence of a dark triangular mark on the occiput. Eleutherodactylus nigrovittatus has smooth skin on the dor- i sum, a dark anal patch, and no triangular mark on the occiput. The snout is broadly rounded in dor- sal view and truncate in profile. The skin is smooth on the dorsum and venter; dorsal tubercles and folds are absent. The digits are short and lack expanded discs; an external tympanum is present. The dorsum is reddish tan to pale brown with dark brown transverse bars on the limbs and dark brown or black longitud- inal dashes dorsolaterally on the body. Canthal and postorbital stripes are ab- sent. The upper lips are barred with dark brown and white or pale tan. The forelimbs and flanks are orange-tan; creamy white mottling is present on the flanks in one individual. A dark brown inguinal spot is present in most speci- mens. The subanal region is dark brown; distally the posterior surfaces of the thighs are tan. The ventral surfaces are pale gray with minute white flecks. The iris is dull bronze suffused with black, giving an overall gray appearance. $ 19 mm; 9 22 mm. Occurrence. — The limited data on six specimens agree with more extensive ob- servations on this species in the Cordil- lera del Due to the west of Santa Cecilia. The species is active by day on the forest floor. One was found in primary forest, five in secondary forest; one was sitting on the ground at night, and the others were active by day. Life History. — Four adult females were found, one in May, and three in July; the latter all had snout-vent lengths of 22.0 mm and contained 9-10 (Xr=9.3) ovarian eggs 2.5 mm in diameter. Data are insufficient to determine the season- ality of breeding. Presumably eggs are deposited on the ground. Mating Call. — The call is unknown. Food. — Four individuals had eaten 9 ants, 2 spiders, 2 beetles, 2 orthopter- ans, one beetle larva, and one annelid worm. EQUATORIAL HERPETOFAUNA 97 Fig. 61. — EleutherodactyJus ockendeni. Upper: ?, 27 mm SVL (KU 126213). Lower: S, 20 mm SVL ( KU 126214). Eleutherodactylus ockendeni (Boulenger) ( Figure 61 ) Hijlodes ockendeni Boulenger, 1912, Ann. Mag. Nat. Hist., (8)10:187 [Type locality.— La Union, Rio Huacamayo, Carabaya, Departa- mento Puno, Peru]. Eleutherodactyhis ockendeni — Gorham, 1968, Das Tierreich, 85:89. Material. — Dureno, 1; Lago Agrio, 6; Puerto Libre, 22; Puerto Ore, 6; Santa Cecilia, 127. Identification. — Most individuals of of this species have W- or H-shaped tu- bercular ridges in the scapular region; otherwise the skin on the dorsum is smooth. Some individuals of E. croceo- infiuinis have a shallow W-shaped series of folds in the scapular region; else- where the skin on the dorsum is tubercu- lar. Many short ridges are present on the back of E. sulcatiis, and in some individ- uals the ridges form an H-shapcd pattern in the scapular region, but E. suJcafus has short slender digits and a broad head (>50% of snout- vent length), whereas E. ockendeni has long digits with ex- panded discs and a narrow head ( ± of snout- vent length). Young individ- uals of E. ockendeni or specimens having poorly developed scapular patterns can be confused with E. paidulu.s, which lacks a scapular arrangement of tubercu- lar ridges but has dark canthal and post- orbital stripes and a pale gold iris with a black horizontal stripe. Canthal and postorbital stripes are absent in E. ocken- deni, in which the iris is metallic green above and red below or green with a median horizontal red streak. The snout is round in dorsal view and truncate in profile. An external tym- panum is present. The tenninal discs on the fingers are expanded about two and one-half times the width of the digit and are truncate. The skin on the belly is granular; with the exception of an H- or W-shaped arrangement of tubercules in the scapular region, the skin on the dor- sum is smooth. The dorsum varies from tan to reddish brown with dark brown markings consisting of H- or W-shaped marks corresponding to the tubercules in the scapular region, one or two narrow transverse marks post-sacrally, and trans- verse bars on the limbs. Some individ- uals have a brown interorbital bar, and others have a pair of small dorsolateral spots in the scapular region. Canthal and postorbital dark stripes are absent. Dark brown bars are present on the lips, but these are faint in some individuals. The flanks are paler than the dorsum, plain, or marked with irregular brown spots. The anterior and posterior surfaces of the thighs are brown to rose-red. The venter is pale gray. In most indi\'iduals the iris is metallic green; the lower half is red or a median horizontal red streak is present. In some individuals the upper part of the iris is bronze. S 21 mm; $ 29 mm. Occurrence. — Data on 141 specimens reveal that this strictly forest-inhabiting frog is active on the forest floor by day and on low vegetation at night. About half (53%) of the frogs were in primary forest. Twenty-seven frogs were found on the forest floor by day, and three 98 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY were on the ground at night. Of 111 found on vegetation at niglit, only three were at heights of more than 1.5 m. Life History. — Thirty-six females con- taining mature ovarian eggs were found throughout the year, except September and November through January. Non- gravid females were obtained in every month except January. Twenty-nine gravid females had snout-vent lengths of 23.0-29.0 (X=26.4) mm and contained 5-18 (X=13.0) ovarian eggs. The larg- est eggs were 2.8 mm in diameter. Four clasping pairs were found in April (2), May ( 1 ) , and June ( 1 ) . Apparently breeding takes place throughout the year, with the possible exception of No- vember-January. Eggs proljably are de- posited amidst leaf litter on the forest floor. Mating Call — The call consists of two notes, "aah-aah." Food. — In 25 stomachs, orthopterans were found in 68.0%, followed by spiders (24.0%), beetles (12.0%), and ants (8.0%). Eleutherodactylus orphnolaimus Lynch ( Figure 62 ) Eleutherodactylus orphnolaimus Lynch, 1970, Proc. Biol. Soc. Washington, 83:221 [Type locality. — Lago Agrio, Provincia Napo, Ecuador]. Material. — Lago Agrio, 2. Identification. — This is one of two species in the area having tubercles on the eyelid. The other species, E. sulca- tus, has a broad head ( > 50% of snout- vent length), demial ridges on the dor- sum, and short fingers lacking expanded discs. Eleutherodactylus orphnolaimus has a much narrower head, no dennal ridges on the dorsum, and long fingers with expanded discs. The snout is pointed in dorsal view and in profile; the tip of the snout pro- jects well beyond the margin of the lip. An external tympanum is present. The discs on the fingers are expanded, two and one-half times the width of the digit, Fig. 62. — Eleutherodactylus orphnolaimus, S , 24 mm SVL (KU 125332). and truncate. The skin on the dorsum is smooth, and that on the belly is granular. The dorsum is tan to grayish tan, with or without a sufi^usion of pale green, with a narrow brown interorbital bar and two narrow brown chevrons on the body. Faint brown labial bars and narrow transverse bars on the limbs are evident. Dark brown canthal and postorbital stripes are present. The flanks are brown. The groin and anterior and posterior sur- faces of the thighs are reddish orange. The belly is brown with white flecks, and the throat is brown with tan streaks. The iris is pale bronze, with or without a pinkish suflPusion dorsally, and a me- dian horizontal reddish brown streak. S 24 mm; 9 34 mm. Occurrence. — Both specimens were obtained on the ground immediately af- ter large trees had been felled in primary forest. These observations and the en- larged tenuinal discs on the toes suggest that the species is arboreal. Life History. — No data are available. The single female, obtained in May, showed no evidence of reproductive ac- tivity. Mating Call. — The call is unknown. Eleutherodactylus paululus Lynch ( Figure 63 ) Eleutherodactylus paululus Lynch, 1974, Occas. Papers Mus. Nat. Hist. Univ. Kansas, 31:6 [Type locality. — Lago Agrio, Provincia Napo, Ecuador]. Material. — Lago Agrio, 4; Puerto Libre, 1; Santa Cecilia, 5. EQUATORIAL HERPETOFAUNA 99 Fig. 63. — Eleutherodactylus paululus, $ , 16 mmSVL (KU 126208). Identification. — This small species has smooth skin on the dorsum, expand- ed discs on the digits, and a dull green dorsum. Two other Eleutlierodactylus in the area are green dorsally; E. acumi- natus has a concealed tympanum and a black canthal stripe, and E. pseudoacu- minatus has a narrow snout and narrower digits than E. paululus. The species re- sembles small individuals of E. ocken- deni, most of which have H- or W- shaped tubercular ridges in the scapular region. Furthermore, E. paululus has dark canthal and postorbital stripes ( ab- sent in E. ockendeni) and a pale gold iris with a median horizontal black stripe (metallic green and red in E. ocken- deni). The coloration of E. paululus is similar to that of E. martiae, which dif- fers by having a concealed tympanum and scattered pustules on the dorsum. Eleutherodactylus corceoinguinis super- ficially resembles E. paululus but differs by having tubercular skin on the dorsum and large yellow spots in the groin. The latter character is shared with E. varia- bilis, which has smooth skin on the dor- sum. The snout is round in dorsal view, shallow and truncate in profile. An ex- ternal tympanum is present. The discs on the fingers are expanded, about twice the width of the digits, and truncate. The skin on the dorsum is smooth, that on the belly granular. The dorsum is dull green or greenish gray, with or without a tan suff^usion laterally, with reddish brown or grayish black markings consisting of a narrow interorbital stripe, narrow can- thai and postorbital stripes, faint labial bars, a blotch or W-shaped mark in the scapular region, one or two chevrons or pair of spots posteriorly on the body, and narrow transverse bars on the limbs. The flanks are pale gray or tan, plain or marked with the posterolateral ends of dorsal chevrons. The anterior and pos- terior surfaces of the thighs are grayish tan. The proximal posterior surfaces of the thighs are brown. The venter is pale green with small white flecks on the belly. The iris is pale bronze to gold with a median horizontal brown or black streak. $ 16 mm; $ 21 mm. Occurrence. — Four individuals were amidst leaf litter on the ground in pri- mary forest by day; four were on low bushes in secondary forest, and one was on a bush in a clearing, all at night. These limited data suggest that the spe- cies is nocturnal and a forest inhabitant. Life History . — Adult females were found in March- May and July- August. One each from March and May were gravid. Each had a snout-vent length of 19.0 mm and contained five ovarian eggs 2.0 mm in diameter. One female from each of March and May was in amplexus. The data are too scanty for conclusions regarding seasonality of breeding. Pre- sumably eggs are laid on the forest floor. Mating Call. — The call is unknown. Food. — Eight individuals had eaten 7 flies, 2 beetles, 1 homopteran, and 1 spider. Eleutherodactylus pseudoacuminatus Shreve (Figure 64) Eleutherodactylus pseudoacuminatus Shreve, 1935, Occas. Papers Boston Soc. Nat. Hist., 8:218 [Type locality. — Sarayacu, Provincia Pastaza, Ecuador]. Material. — Lago Agrio, 9; Puerto Libre, 7; Puerto Ore, 1; Santa Cecilia, 23. 100 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 64. — Eleutherodactijlus pseudoacuminatus, $,21mmSVL (KU 126164). Identificotion. — This is one of three species of green Eleutherochictyhis in the area; E. acuminatus is a larger species having the tympanum covered with skin and having a black stripe from the snout through the eye. In E. pseudoacumi- natus and pauhdus the tympanum is visi- ble externally, and a black canthal stripe is absent. Eleutherodactijlus pauhdus differs from E. pseudoacuminatus by having a broader snout and broader discs on the fingers. Other small green frogs in the area are centrolenids and hylids, all of which have extensive webbing be- tween the toes; webbing is absent in E. pseudoacuminatus. The snout is narrow and rounded. The skin on the dorsum is smooth; that on the belly is granular; dorsal tubercles and folds are absent. The discs on the fingers are slightly expanded (1.5 X digit) and round. Usually the dorsum and the anterior and posterior surfaces of the thighs are uniform pale green to grayish green. Some individuals have a tan suffusion, especially middorsally. In a few individuals, a narrow creamy white canthal stripe is present. Two specimens had creamy yellow dorsolateral stripes, and one had a cream snout. Small black flecks are present on the dorsum in some individuals. The venter and flanks are white. The iris is pale bronze with a median, horizontal red stripe. $ 19 mm; ? 23 mm. Occurrence. — Frogs of this species are found in about equal abundance in primary and secondary forest. Three were obtained from bromeliads by day, and two were on the ground at night. All others were on vegetation at night; only four of those were at heights of more than 1.5 m. Life History. — Fourteen gravid fe- males were obtained throughout the year: February (3), March (1), May (3), July (4), August (2), December (1). Twelve gravid females having snout- vent lengths of 18.0-22.0 (X= 20.8) mm contained 4-11 (X=7.3) ovarian eggs having diameters of 3.0 mm. Four of these females were in amplexus (February, March, May and December). One female deposited 7 unpigmented eggs 3.5 mm in diameter in the labora- tory on 17 December. Two of the eggs hatched on 15 January; both hatchlings had snout- vent lengths of 5.0 mm. Ap- parently E. pseudoacuminatus breeds throughout the year. Oviposition sites are unknown, but the fact that this or- boreal species inhabits bromeliads, sug- gests that eggs may be deposited in bromeliads. Mating Call — The mating call is a soft, buzz-like peep. Food. — Only 11 stomachs contained food. Small beetles were found in 36.4 percent of the stomachs, followed by flies (27.3%), ants (18.2%), homopterans (18.2%), and spiders (18.2%). Eleutherodactylus quaquaversus Lynch ( Figure 65 ) Eleutherodactylus quaquaversus Lynch, 1974, Occas. Papers Miis. Nat. Hist. Univ. Kansas, 31:9 [Type locality. — South slope of Cordi- llera del Due, above Rio Coca, 1150 m, Provincia Napo, Eucadorl. Material — Puerto Libre, 3; Santa Ce- cilia, 10. Identification. — This is one of three species of Eleutherodactyhis in which the tympanum is not visible. Among the three species, E. acuminatus is readily distinguished by its unifomrly green dor- sum and bold black canthal stripe. Eleu- therodactylus quaquaversus has smooth EQUATORIAL HERPETOFAUNA 101 Fig. 65. — Eleutherodactyhis quaqtiaversus. Up- per: S, 23 mm SVL; Cordillera del Due, Provincia Napo, Ecuador (KU 123746). Mid- dle: 5, 31 mm SVL; Cordillera del Due, Provincia Napo, Ecuador (KU 123749). Low- er: $, 22 mm SVL; 17.5 km NNE Santa Rosa, Provincia Napo, Ecuador (KU 143441). skin on the dorsum, no bars on the hps, and a pointed tubercle on the heel. By comparison, E. martiae has scattered pus- tules dorsally, dorsolateral dermal folds, bars on the lips, and no tubercle on the heel. The combination of a covered tym- panum and a tubercule on the heel im- mediately distinguishes E. quaquaversus from other Eleutherodactyhis in the area. The snout is round in dorsal view and in profile. The discs on the fingers are about twice the width of the digits and truncate. The skin on the dorsum is smooth; that on the belly is granular. The dorsal ground color is dull red, tan, brown, or gray with irregular darker markings consisting of an interorbital bar, longitudinal dashes or transverse marks on the body, and transverse bars on the limbs. In darker colored individ- uals a narrow cream canthal stripe is evi- dent. Although dark vertical bars are not present on the upper lips, one or two small dark spots are present in some in- dividuals; others have one or two small white spots below the eye. A few indi- viduals have a broad cream interorbital bar, and one has a cream snout. A post- orbital stripe usually is absent. The flanks are tan or pale gray; small dark flecks are present in some individuals. The groin and anterior and posterior sur- faces of the thighs are purplish red with small white flecks. The venter is bronze- white, with small scattered black flecks in most individuals. The iris varies from silvery cream to green with a red tint below. $ 22 mm; 9 29 mm. Occurrence. — This species is noc- turnal in the forest; 9 of 13 individuals were in secondary forest. All were on low vegetation at night. These observa- tions are corroborated by more extensive data from the Cordillera del Due and Cordillera Umbaqui to the west of Santa Cecilia, where most of the specimens of this species have been collected. Life History. — One gravid female from September, 29.0 mm in snout-vent length, contained 22 ovarian eggs 2.6 mm in diameter. Mating Call — The mating consists of a single high-pitched note. Food. — A variety of small arthropods have been eaten by 13 individuals. Bee- tles were present in 35.3 percent of the stomachs, followed by orthopterans (29.4%), spiders (23.5%), lepidopteran larvae (11.8%), and ants (11.8%). 102 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 66. — Eleutherodactylus sulcatus, 2 , 49 mmSVL (KU 123586). Eleutherodactylus sulcatus (Cope) (Figure 66) Hylodes sulcatus Cope, 1874, Proc. Acad. Nat. Sci. Philadelphia, 26:126 [Type locality. — Nauta, Departamento Loreto, Peru]. Eleutherodactylus stdcatus — Gorham, 1966, Das Tierreich, Berlin, 85:103. Material. — Lago Agrio, 6; Puerto Libre, 3; Santa Cecilia, 22. Identification. — This toad-hke Eleu- therodactylus has a broad depressed head ( > 50% of snout-vent length ) with cranial ridges, short digits lacking ex- panded terminal discs, granular skin on the venter, tubercles on the eyelids, and longitudinal demial ridges on the doi"- sum. The only other species of Eleu- therodactylus in the area having short digits without expanded terminal discs is E. nigrovittatus, which has a narrower head, smooth skin on the venter, and no dennal ridges on the dorsum. The only other species having tubercles on the eyelids, E. orphnolaitnus, has smooth skin on the dorsum and long lingers with expanded discs. Ischnocnema quixensis also has a short, squat body, but it differs from E. sulcatus by having a narrower head, tubercular skin on the dorsum, and small light flecks on the posterior sur- faces of the thighs. All other species of Eleutherodactylus with pale spots on the posterior surfaces of the thighs have ex- panded discs on the fingers. True toads (Bufo) have paratoid glands. The snout is broad and rounded in dorsal view, depressed and round in profile. The skin on the venter is granu- lar; that on the dorsum is smooth with scattered small tubercles and short, lon- gitudinal dermal ridges, which tend to form an H-shaped pattern in the scapular region in most individuals. An external tympanum is present. The dorsum is dull gray or reddish tan with or without irregular small dark brown marks or larger markings consisting of an inter- orbital bar, a blotch in the scapular re- gion and one or two chevrons or trans- verse bars on the posterior part of the body. Narrow dark brown transverse bars are present on the limbs. The lips usually are boldly marked with dark brown bars. Canthal and postorbital stripes are absent. The flanks are pale gray or pinkish tan. The groin and the anterior and posterior surfaces of the thighs are dark brown to black with large cream spots. The venter is grayish cream; the throat is noticeably darker in some individuals. The ventral surfaces of the hands and feet are dull orange. The iris is greenish silver, reticulated with black. S 35 mm; 9 54 mm. Occurrence. — This large, terrestrial Eleutherodactylus occurs only in forests, where individuals have been found ac- tive or amidst leaf litter by day. Three were found on the forest floor by day. Ten of 26 specimens were taken in pri- mary forest. Life History. — Of eight adult females, only one (collected in July) was gravid; it had a snout-vent length of 49.0 mm and contained 32 ovarian eggs 2.5 mm in diameter. One juvenile having a snout- vent length of 11.0 mm was found in March, and two having lengths of 13.0 mm were found in May. Presumably eggs are deposited on the ground. Mating, Call. — The call is unknown. Food. — A variety of terrestrial arthro- pods was included in the diets of 15 in- dividuals. Orthopterans were found in EQUATORIAL HERPETOFAUNA 103 53.3 percent of the stomachs, beetles in 20.0 percent, centipedes in 20.0 percent, and milhpedes in 13.3 percent. Eleutherodactylus variabilis Lynch (Figure 67) Eleutherodactylus variabilis Lynch, 1968, J. Herp., 2:129 [Type locality. — Limoncocha, Provincia Napo, Eucador]. Material. — Lago Agrio, 2; Puerto Libre, 49; Santa Cecilia, 268. Identification. — This is one of two species of Eleutherodactylus in the area having large yellow spots in the groin. In E. variabilis the spots (one on each side ) are pale yellow, narrowly bordered by black, and confluent, or nearly so, midventrally. The other species, E. cro- ceoinguinis, has deep yellow or orange spots that do not approach the midline. Furthermore, in E. variabilis the body is more robust, and the skin on the dorsum is finely granular, whereas in E. croceo- inguinis the body is more robust, and the skin on the dorsum is tubercular. The snout is narrow and rounded in dorsal view and in profile; an external tympanum is present. The discs on the fingers are truncate and about twice the width of the digits. The skin on the dor- sum is finely granular ( shagreened ) ; that on the belly is granular. The dorsal col- oration is highly variable, ranging in color from tan to brown, gray, red, or green. Most individuals have a pattern of darker markings consisting of a nar- row interorbital bar and transverse or diagonal markings on the body and limbs; the markings on the body usually extend onto the flanks. In some individ- uals the dorsum is uniform tan, green, or dull red, with or without darker longi- tudinal streaks; in these individuals the flanks are brown or gray. A few individ- uals have a narrow middorsal cream or yellow stripe. Canthal and postorbital stripes are absent or dark brown, when present. Dark brown diagonal bars are present on the lips. A large pale yellow spot extends from the inguinal region Fig. 67. — Eleutherodactylus variabilis. Upper: $, 21 mm SVL (KU 111168). Lower: 5, 23 mmSVL (KU 111140). onto the proximal anterior surface of the thigh. Usually the spot has a narrow black border. The spots on either side are confluent, or nearly so, midventrally. The posterior surfaces of the thighs are dull red, rose-pink, or gray. The ventral surfaces are cream with brown or gray flecks. The iris is bronze with a median horizontal reddish brown streak. $ 19 mm; 5 27 mm. Occurrence. — Eleutherodactylus vari- abilis apparently is a "weed" species most frequently encountered in disturbed areas. Only five individuals were found by day - three on the ground in primary forest (two where trees were being felled), one amidst leaf litter in secon- dary forest, and one on the ground in a clearing after a rain. All others were found at night on low vegetation - sec- ondary forest (210), primary forest (16), swamp (21), clearing (3), banana grove ( 1 ) . Observations made in 1971-72 indi- cate that E. variabilis is especially abun- dant in partially cleared forest and in forest-edge habitats. Life History. — No fewer than six gravid females were found in any given 104 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY month; 195 of the 289 females contained mature ovarian eggs. One hundred and forty-eight gravid females having snout- vent lengths of 22.0-27.0 (X=24.4) mm contained 2-11 (X=6.7) ovarian eggs having diameters up to 3.0 mm. Amplec- tant pairs were found on low vegetation in March (3), April (1), May (1), June (2), August (1), October (1), and No- vember ( 1 ) . Six captive pairs deposited eggs in the laboratory. In clutches of 5, 6, 7, 7, 7, and 11 unpigmented eggs, the diameters of the eggs were 3.8, 4.0, 4.0, 3.9, 3.8, and 4.0 mm, respectively. Ob- viously this species breeds throughout the year. Possibly the eggs are deposited amidst leaf litter or beneath detritus on the forest floor or amidst debris in bushes or low trees. Mating Call. — The call consists of a series of soft clicks. Food. — Orthopterans were found in 38.4 percent of 26 stomachs; ants, flies, beetles, and spiders were each found in 15.4 percent. Hemipterans, homopter- ans, and beetle larvae also were included in the diets. Ischnocnema quixensis (Jimenez de la Espada) (Figure 68) Oreobates quixensis Jimenez de la Espada, 1872, An. Soc. Espan. Hist. Nat., Madrid, 1:87 [Type locality. — "Quito, Ecuador"]. Ischnocnema quixensis — Lynch, 1971, Misc. Publ. Mus. Nat. Hist. Univ. Kansas, 53:156. Material. — Dureno, 1; Lago Agrio, 7; Puerto Libre, 9; Puerto Ore, 1; Santa Ce- cilia, 106. Identification. — This robust toad-like species has large tubercles on all dorsal surfaces of the body. It differs from toads of the genus Bufo by having tu- bercles on the head and by lacking bony cranial crests. Among the many species of Eleutherodactylus in the area, Isch- nocnema can be confused only with E. sulcatus, which has a smooth dorsum with longitudinal dennal ridges. Small individuals of Ischnocnema might be mistaken for Phijsalaemus petersi, anoth- FiG. 68. — Ischnocnema quixensis, $ , 55 mm SVL (KU 146174). er toad-like species with black spots in an orange groin. The snout is acutely rounded in dor- sal view and round in profile. An exter- nal tympanum is present. The fingers and toes are moderately long, unwebbed, not expanded terminally and bear large conical subarticular tubercles. The skin on the dorsum is tubercular; that on the belly is smooth. The dorsum varies from pinkish brown to olive-brown with dark brown to reddish brown markings con- sisting of a broad interorbital bar, a chev- ron or H- or W-shaped mark in the scap- ular region, irregular spots or transverse marks elsewhere on the back, vertical bars on the lips, and broad transverse bars on the limbs. The flanks are brown; some individuals have a pinkish tint in the inguinal region. The anterior and posterior surfaces of the thighs are brown. The ventral surfaces are brown to grayish brown with white mottling. The iris is coppery bronze. S 48 mm; 9 59 mm. Occurrence. — This frog primarily is a noctinnal forest dweller. Of 91 speci- mens, only 24 were obtained by day; of these, six were under logs, 16 were in leaf litter, and two were in a small stream. A juvenile was sitting on a low herb at EQUATORIAL HERPETOFAUNA 105 night; otherwise all other individuals were on the ground. Two of the frogs were in clearings and one in a banana grove; all others were in forest (primaiy 23%, secondary 77%). Life History. — Gravid females were found in every month except January and October. Thirty-four females having snout- vent lengths of 47.0-59.0 (X=52.9) mm contained 15-51 (X=35.1) ovarian eggs having diameters up to 4.1 mm. Juveniles having snout-vent lengths of 11.0-14.0 mm were found in January, March, June, and July. Thus, it seems that this species breeds throughout the year. Previously the life history of Isch- nocnema has been unknown. Although the present data are not conclusive, I believe that it is safe to assume that Ischnocnema quixensis is like Eleuthero- dactijhis in depositing clutches of terres- trial eggs that undergo development di- rectly into small frogs. This assumption is based on the small clutch size, large ovum size, and lack of pigment in the eggs. Mating Call. — The call consists of a series of low-pitched guttiu-al notes. Food. — Examination of 26 stomachs revealed the presence of orthopterans in half of the stomachs. Other major food items were beetles (11.5%) and milli- pedes (11.5%), whereas spiders, beetle larvae, and ants were each present in only two stomachs. Each of the follow- ing were found in onlv one stomach: hemipteran, lepidopteran larva, and gas- tropod. Leptodactylus discodactylus Boulenger (Figure 69) Leptodactylus discodactylus Boulenger, 1883, Proc. Zool. Soc. London, 1883:637 [Type locality. — Yurimaguas, Departamento Lo- reto, Peru]. Material. — Santa Cecilia, 26. Identification. — Leptodactylus disco- dactylus and L. ivagneri differ from the other species of Leptodactylus in the area by having lateral fringes on the toes. Fig. 69. — Leptodactylus discodactylus, $ , 33 mmSVL (KU 143518). These two species differ in that L. disco- dactylus has expanded, round, tenninal discs on the digits and usually a dark throat and cream belly, whereas L. ivag- neri lacks expanded temiinal discs on the digits and has a cream throat and belly, both mottled with brown. Furthermore, L. discodactylus is smaller (< 36 mm) than L. wagneri ( < 78 mm). Leptodac- tylus discodactylus is most easily con- fused with Adenomera andreae, a small- er species ( < 28 mm ) lacking lateral fringes on the toes and having the first two fingers equal in length; the first finger is longer than the second in L. discodactylus. The other species of Lep- todactylus in the area, except L. ivagneri, have distinct dorsolateral demial folds. The presence of short toes with basal webbing distinguishes L. discodactylus from Eleutherodactylus, and the expand- ed discs and little webbing distinguish it from Rana. The dorsum is dark brown, and the flanks are grayish brown or reddish tan, usually with a brown spot in the groin and one or two brown spots on the flank. A narrow pinkish cream or gray inter- orbital bar usually is evident. The side of the head is dark brown, usually with four narrow, diagonal or vertical cream bars ( two below orbit, one below nostril, and one between nostril and orbit); another cream bar is present on the tip of the snout. The limbs are dark brown with darker transverse bars barely evi- dent in some specimens. The posterior 106 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY surfaces of the thighs are brown; black flecks are evident distally in some speci- mens. The throat is dark brown with cream flecks, and the belly and ventral surfaces of the limbs are cream with brown flecks laterally. The iris is bronze with brown reticulations. Some adults of both sexes have bright Nellow glandular areas laterally on the body. Breeding males lack nuptial pads or spines. <^ 29 mm; 9 35 mm. Occurrence. — All individuals of Lep- todactylus discodactylus were found in forest. Two were resting on low vege- tation at night, two were in a swamp at night, and one was on the forest floor at night. Nineteen individuals were found by day; all of these were in swampy areas. Although all but three were near water, only two were in water. The small sample of L. discodactijhis is from March-October with no specimens from May and September and 15 specimens from July. My general impression is that this frog is restricted to swamp forest, where it probably is active throughout the year. Life History. — Six females having 148-420 (X=234.8) mature ovarian eggs were found in April (2), June, July (2), and October; the maximum diameter of the eggs is 1.2 mm. Heyer and Bcllin (1973) noted that at Limoncocha, Ecua- dor, male L. discodactylus called day and night after heavy rains and that they did not call during rainless periods or times of light rain. The smallest juve- niles (7.3-7.7 mm in snout-vent length) were found in July. These limited data suggest that L. discodactylus may be a sporadic opportunistic Inecder, whose breeding activity is associated with pe- riods of heavy rainfall. At such times depressions in the forest are filled with water and form suitable breeding sites. Tadpoles. — No tadpoles of this spe- cies were found at Santa Cecilia, but tadpoles are available from Mera, Pro- vincia Pastaza, Ecuador, where they were obtained in a shallow forest-edge swamp. The following description is based on KU 121362 (tadpoles in stage 30). The body length is 10 mm, and the total length is 25 mm. The body is ovoid, wid- est and deepest posteriorly; the snout is bluntly rounded in dorsal view and acutely rounded in profile. The eyes are small, close together, and directed dorso- laterally. The nostrils are about two- fifths the distance from the tip of the snout to the eyes. The spiracle is sinis- tral; the opening is directed posterodor- sally at a point just below the midline at midlength of the body. The mouth is ventral. The lips have lateral folds; the median part of the upper lip is bare; elsewhere there is a single row of labial papillae. There are two upper and three lower rows of denticles; all are complete. The beaks are slender and finely serrate. The dorsal and ventral fins are equal in depth and slightly shallower than the caudal musculature at midlength of the tail. The dorsal fin does not extend onto the body and is deepest at about two- fifths of the length of the tail. The tip of the tail is acutely rounded. The body is brown with white flecks ventrally. The caudal musculature is dull brown, and the fins are tan. Mating, Call. — A recording of Lepto- dacfyliis discodactylus made at Limon- cocha, Ecuador, generously was provid- ed by W. R. Heyer. The call consists of a series of short notes "wheeet, wheeet, wheeet" produced at a rate of 44 notes per minute. The dominant frequency of the notes is at about 3000 Hz. Food. — Of eight stomachs containing food, five contained eight beetles com- prising 41 percent of the bulk. Two con- tained one homopteran each, represent- ing S.7 percent of the bulk. Other food items were represented by one individual each and included Diptera, Orthoptera, formicid, Hemiptera, Isopoda, and An- nellida. Ta .V o n o m ic Co m ments. — Heyer (1974b) suggested that this species is generically distinct from Leptodactylus and proposed for it the generic name Vanzolinius. EQUATORIAL HERPETOFAUNA 107 Fig. 70. — Leptodactyhis miistaceus, juvenile, 17 mmSVL (KU 122555). Leptodactylus mystaceus (Spix) (Figure 70) Raua mijstacca Spix, 1824, Spec. Testud. Ran. Biasil., Monaco: 27 [T\npe locality. — Baliia and Solimoes, Brasil]. Leptodactylus mijstaceus — Mehely, 1904, Ann. Mns. Hungar., Budapest, 2:219. Materia]. — Lago Agrio, 2; Puerto, 1; Santa Cecilia, 114+21. Identification. — This species of Lep- todactyhis is readily distinguished from other frogs in the area by its acuminate snout and distinct creamy white labial stripe. The only other Leptodactylus having a pale labial stripe is L. rhodo- mystax, in which the stripe is pinkish tan. Also, L. rhodomystax has a blunt snout and a dark venter; the venter in L. mystaceus is creamy white. Leptodac- tylus mystaceus is like other members of the genus, except L. discodactylus and wagneri, in lacking lateral fringes on the toes. Leptodactyhis mystaceus has dorsolateral dermal folds, a charac- ter shared with L. pentadactyhis, L. rho- domystax, L. rhodonotus, and Rami pal- mipes. The white labial stripe and cream belly immediately distinguish L. mysta- ceus from all of these frogs, and basal webbing between the toes serves as an additional difference from Rami pal- mipcs, which has extensive webbing. Leptodactyhis mystaceus frequently is mistaken as an Eleutherodactylus, but no species of Eleutherodactylus in the area (except variants of E. lanthanites, which have expanded discs on the digits) has a white labial stripe and dorsolateral folds. The dorsum is gray, olive-tan, or red- dish tan with darker brown or grayish brown blotches consisting of a triangular mark with the apex posteriorly on the head and usually three crude chevron- shaped marks middorsally. The hind limbs are colored like the body and have dark transverse bars; the forelimbs are paler than the body and usually have faint transverse bars on the forearms. The posterior surfaces of the thighs are mottled brown and black with a longi- tudinal cream stripe distally. The dorso- lateral fold is dark brown with a cream streak on the medial edge posteriorly. The flanks are pale gray or tan. The side of the head, including the tympanic region, is black, sharply defined above by a narrow bronze-tan canthal stripe. A broad silvery white or creamy white labial stripe extends from the tip of the snout to the base of the ami. The venter is creamy white with brown flecks lat- erally on the throat and belly. The iris is bronze above and dark metallic red- dish brown below. Breeding males lack nuptial pads or spines. 6 56 mm; 9 60 mm. Occurrence. — Leptodactyhis mysta- ceus was found throughout the year in primary forest (15%), secondary forest (50%), forest edge (5%), and clearings ( 30%) . About one-third of the frogs were found by day in leaf litter or under logs. At night they were on the ground in forest or clearings, but 10 were found on low bushes ( < 1.5 m), and two were in water in swamps. Most (84%) individ- uals were not near water. Life History. — Males call throughout the year from secluded sites ( under logs, under tree roots, in holes in the ground). Females bearing mature ovarian eggs were found in every month except April, June, September, and December; no fe- males were found in June. Apparently 108 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY L. mystaceus breeds throughout the year. The unpigmented eggs are deposited in a foam nest; one nest about 100 mm in diameter and 50 mm in deptli was in a cavity under a rock in a marshy clearing; others were found in weedy shallow wa- ter. Twenty-two females having snout- vent lengths of 52.0-60.0 (X=54.8) mm contained 171-425 (X=280.5) eggs, the largest having a diameter of 2.4 mm. Nine recently metamorphosed young have snout-vent lengths of 14.0-15.5 (X=: 14.7) mm. Tadpoles. — The following description is based on KU 109168, stage 27, having a snout-vent length of 9 mm ^md a total length of 26 mm. The bodV is ovoid, widest medially and flat ventrally. In dorsal view the snout is bluntly round- ed, whereas in profile it is inclined an- teronventrally and rounded below. The eyes are moderate in size, well separated from one another, and directed dorso- laterally. The nostrils are midway be- tween the tip of the snout and the eyes. The spiracle is sinistral; the opening is directed posterodorsally at a point just below the midline at about two-thirds of the length of the body. The mouth is ventral and has lateral folds. The me- dium part of the upper lip is bare; else- where these is one row of labial papillae. There are two upper and three lower rows of denticles; the second upper row is interrupted medially. The beaks are slender and finely serrated. The dorsal and ventral fins are equal in depth and shallower than the caudal musculatiu-e. The dorsal fin does not extend onto the body and is deepest at midlength of the tail. The tip of the tail is pointed. The body is olive-tan; in large individuals ( > stage 37 ) a dark brown dorsolateral stripe and an interorbital bar are evident. The venter is gray with gold flecks an- teriorly. The tail is olive-tan with dull green and orange-yellow flecks and dark brown reticulations. The iris is tiiolden bronze. Tadpoles have been found in shallow grassy pools in clearings, in a IlcUconia- choked swamp in forest, and in a par- tially shaded pond at the edge of the forest. All tadpoles were observed in shallow water ( < 25 cm ) , where they took refuge in grasses or amongst leaves J on the bottom. Mating Call. — The call consists of a series of notes "oit, oit, oit." Analysis of one recording made at Santa Cecilia re- veals that the note repetition rate is 48 notes per minute with the dominant frequency at about 1500 Hz. and an em- phasized harmonic at about 6000 Hz. Food. — Sixteen of 26 frogs examined contained orthopterans, and 10 of these contained only orthopterans, which make up 81 percent of the bulk of the food. Individual beetles were present in six stomachs and represent 8 percent of the bulk. The other food items were single examples of a variety of small arthropods. Leptodactylus pentadactylus (Laurenti) (Figure 71 ) Rana pcntadacttjia Laurenti, 1768, Synops. Rept., Wien: 32 [Type locality. — "Indiis"]. Leptodactylus pentadactylus — Nieden, 1923, Das Tieneich, Bedin, 46:472. Material. — Dureno, 1; Lago Agrio, 2; Puerto Libre, 1; Santa Cecilia, 30+13. Identification. — This large Leptodac- tylus is like L. mystaceus, L. rhodomy- stax, L. rhodonotus, and Rana palmipes in having distinct dorsolateral dermal folds. Leptodactylus pentadactylus usu- ally has a reddish brown dorsum and has the toes webbed basally in contrast to Rana palmipes, which is grayish green dorsally and has extensive webbing be- tween the toes. Of the aforementioned species of Leptodactylus, L. mystaceus and L. rhodomystax have pale labial stripes, whereas L. pentadactylus and L. rhodonotus have dark vertical bars on the lips. These two species differ in col- oration and in the disposition of dermal folds. The dorsum in L. pentadactylus usually has distinct darker brown or red- dish brown transverse blotches, whereas the dorsum in L. rhodonotus is unifomi EQUATORIAL HERPETOFAUNA 109 Fig. 71. — Leptodactylus pentadactyJus. Upper: 5, 154 mm SVL (KU 146176). Lower: juve- nile, 55 mm SVL (KU 122576). dusky brown. The limbs in L. pentadac- tylus have distinct broad, transverse dark bars, but in L. rhodonotus only indis- tinct, narrow, transverse lines are pres- ent. Usually the posterior surfaces of the thighs are black with creamy yellow flecks in L. pentadactyJus and unifomi brown in L. rhodonotus. The dorsolat- eral fold in L. rhodonotus is continuous with the supratympanic fold, but in L. pentadactyhis the dorsolateral fold ex- tends posteriorly from the posterior edge of the eyelid and is distinct from the supratympanic fold, which extends diag- onally onto the flank. In most individuals the dorsum is orange-tan or grayish tan with five or six broad, transverse brown or reddish brown blotches between the dorsolateral folds. The dorsal surfaces of the limbs are tan or gray with distinct, broad, transverse dark brown bars. A narrow black stripe extends from the snout through the nostril and eye to the upper edge of the tympanum. The lips are tan with four or five short, black vertical bars on the margin of the lip. The flanks are paler than the dorsum and marked by a few black spots. The posterior sur- faces of the thighs are black with cream flecks. The venter is cream with bold dark brown mottling, especially on the belly and hind limbs. In some juveniles the dorsum is pale green with dark brown or black markings; the posterior surfaces of the thighs arc black, and the belly is dark grayish brown with or with- out cream flecks. The iris is dark bronze. Breeding males have large nuptial spines on the prepollex and on the chest. S 169 mm; 2 181 mm. Occurrence. — Leptodactyhis penta- dactyhis was found in primary forest. Most individuals were active on the ground at night, but two were in water. Juveniles were found amidst leaf litter or under logs by day. Life History. — Males call sporadical- ly throughout the year. Although some were observed to call from various ter- restrial sites in the forest, most sounded as though they were calling from near the edge of the largest lake. Only ten adult females were found; none was gravid. Tadpoles were obtained in March-July. Small juveniles were taken in June, July, and November. These data are inconclusive with respect to seasonal- ity of breeding. Tadpoles. — The following description is based on KU 146771 (stage 32, body length 18.2 mm, total length 61.5 mm). The body is ovoid, widest at midlength and deepest posteriorly. The snout is bluntly rounded in dorsal view and round in profile. The eyes are moderate- ly large, widely separated, and directed dorsolaterally. The nostrils are midway between the tip of the snout and the eyes. The spiracle is sinistral and di- rected posterodorsally at a point on the 110 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY midline at about three-fifths the length of the body. The mouth is anteroventral and lacks lateral folds. The median part of the upper lip is bare; elsewhere there is one row of labial papillae. There are two upper and three lower rows of den- ticles; the second upper row is broadly interrupted medially, and the third lower row is much shorter than the others. The tail is long (70% of total length), pointed terminally, and has low fins. The dorsal fin does not extend onto the body, is one- third deeper than the ventral fin, and three-fourths of the depth of the caudal musculatine. The body is grayish brown with a brown interorbital bar; the venter is gray. The tail is brown with a creamy orange tint to the edge of the dorsal fin. The iris is bronze. Tadpoles have been found in shallow temporary pools in clearings and in swamps in forests. Mating Call. — The call consists of a series of loud notes, "whooop, whooop, whooop." Some calls consist of only a few notes, whereas others consist of 20- 30 notes, in which case the note repeti- tion rate is 21 notes per minute. In one recording that was analyzed the domi- nant frequency was at about 3300 Hz. Food. — These large frogs have a pro- pensity for eating large arthropods. One contained the remains of a giant tarantula having a chelicera 9 mm in length; another contained a millipede 83 mm in length. Small frogs, crustaceans, and molluscs are taken in addition to a wide variety of insects (Table 5). Taxonomic Comments. — Heyer (1972) named LeptoclactijJus knudscni on the basis of 33 juveniles from Limoncocha, Provincia Napo, Ecuador. The species was diagnosed as differing from L. pen- tadactyhis by having green and black dorsal markings and uniformly dark pos- terior surfaces of the thighs. The largest specimen, the juvenile female holotype, has a snout- vent length of 63 mm. I compared the holotype and six para- types with specimens from Santa Cecilia. Like Heyer, I found no morphological Table 5. — Food Analysis of Leptodactijlus pentadactijlus. ( N:=27 ) Frequency Frequency Percent Prey Item Frogs Prey Bulk Orthoptera 20 25 42.4 Coleoptera 12 24 21.7 Diplopoda 6 9 10.7 Arachnida 3 3 6.8 Cmstacea 1 2 3.3 Anura _- 1 2 3.3 Hemiptera 2 4 3.2 Isopoda 1 2 2.4 Lepidoptera (Iar\'ae) 2 2 2.2 Coleoptera (larvae) 2 2 1.9 Hynienoptera (formicids) -_ 2 5 1.2 Demiaptera 1 1 0.4 Mollusca - 1 1 0.4 Isoptera 1 1 0.1 differences between L. knudseni and L. pentadactijlus. Three individuals from Santa Cecilia (snout-vent lengths of 37, 40, and 52 mm ) have a pale green dorsal ground color with dark brown to black transverse blotches and black flanks and posterior surfaces of the thighs. Among 10 knudseni-MVe frogs, seven have uni- formly grayish brown venters, whereas small cream spots are present on the venters of the other three. Of seven juveniles having the pentadactyhis-\ike dorsal coloration, three have spots on the venter, and four are uniformly grayish brown. Also, three of these seven have uniformly dark posterior surfaces of the thighs. Because of the various combinations of color characters occurring in the ju- veniles, the absence of morphological differences in differently patterned speci- mens, and the absence of adults having the green and black coloration, I am reluctant to recognize Leptodactijlus knudseni as a species distinct from L. pentadactijlus: instead it seems as though two color morphs occur in juvenile L. pentadactijlus. A taxonomic revision of the Leptodactijlus pentadactijlus group is beyond the scope of this paper, espe- cially since there is no biological evi- dence to support the recognition of more EQUATORIAL HERPETOFAUNA 111 than one species of L. pentadoctylus-like frogs at Santa Cecilia. Leptodactylus rhoclomystax Boulenger (Figure 72) Leptodactylus rhodomijstax Boulenger, 1883, Proc. Zool. Soc. London, 1883:637 [Type locality. — Yurimaguas, Provincia Loreto, Peni]. Material. — Lago Agrio, 1; Puerto Libre, 1; Santa Cecilia, 3+2. Identification. — This moderately large species resembles Leptodactylus mysta- ceus, L. pentadactylus, L. rhodonotus, and Rana palmipes in having distinct dorsolateral demial folds. The presence of extensive webbing between the toes distinguishes Rana palmipes from L. rho- domystax, which has only basal webbing. Leptodactylus pentadactylus and L. rho- donotus have dark bars on the lips and thereby differ from L. mystaceus and L. rhoclomystax, both of which have pale labial stripes. Leptodactylus rhodomy- stax has a blunt snout, pinkish tan labial stripe, dark venter, and pale flecks on the posterior surfaces of the thighs; L. mystaceus has an acuminate snout, creamy white labial stripe, cream venter, and a pale stripe on the posterior surface of each thigh. The dorsum is reddish brown to gray- ish brown; the dorsolateral folds are dark brown. The flanks are orange-tan. The groin and posterior surfaces of the thighs are dark grayish brown with small pale greenish yellow spots. The side of the head, including the tympanic region, is dark brown. A pale pinkish tan labial stripe extends from the snout to the base of the ami. The throat is dark brown; the chest, belly, and ventral surfaces of the limbs are cream, heavily suffused with brown. The iris is dull bronze above and reddish bronze below. Breeding males have nuptial spines on the prepol- lex and chest. $ 90 mm. Occurrence. — Three adult males were found at night in swamps at Santa Ce- cilia in March. One adult male was amidst leaf litter in mature forest at Fig. 72. — Leptodactylus rhodomystax, mmSVL (KU 122581). ?, 60 Lago Agrio in May, and one small female was beneath a stone at the edge of a clearing at Puerto Libre in July. Life History. — Except for one male calling in March, nothing is known. Tadpoles. — Tadpoles tentatively iden- tified as this species by W. R. Heyer were found in two swamps in forest in June, 1968. The following description is based on KU 124147 (stage 25, body length 6 mm, total length 21.3 mm ) . The body is ovoid, widest anteriorly and deepest posteriorly. The snout is trun- cate in dorsal view and round in profile. The eyes are large, widely separated, and directed laterally. The nostrils are about one-third the distance from the tip of the snout to the eyes. The spiracle is sinistral; the opening is directed pos- terodorsally at the midline and about two-thirds of the length of the body. The mouth is anteroventral and has no lateral folds. The median part of the upper lip is bare; elsewhere there is a single row of labial papillae. There are two upper and three lower rows of denticles; the second upper row is widely separated medially, and the third lower row is much shorter than the others. The tail is 72 percent of the total length and terminates in a filamentous tip. The dorsal fin is shal- low, one third the depth of the ventral fin, deepest at midlength of the tail, where it is about equal to the depth of the caudal musculature, and does not extend onto the body. The body and 112 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY caudal musculature are brownish gray, and the tail is marked with irregular darker flecks. Mating Call. — Unfortunately the call was neither recorded nor described. Food. — Three individuals contained two roaches (Blattidae), one grasshop- per ( Orthoptera ) , and one millipede ( Diplopoda ) . Leptodactylus stenodema Jimenez de la Espada ( Figure 73 ) Leptodactylus stenodema Jimenez de la Espada, 1875, Vert. Viaje Pacifico, p. 64 [Type lo- cality.— San Jose de Moti, Provincia Napo, Ecuador]. Pyburn and Heyer, 1975, Copeia, 3:585. Material. — Puerto Libre, 1. Identification. — This species has a ro- bust body and short limbs. It is like L. mystaccus, L. pcntadactijlus, L. rhodo- mystax, and Rana pahnipes in having distinct dorsolateral dermal folds. Lep- todactylus stenodema has a reddish brown dorsum and the toes basally webbed, in contrast to Rana paJmipes, which has a grayish green dorsum and the toes extensively webbed. Leptodac- tylus stenodema and L. pentadactylus have dark labial bars, whereas L. mysta- ceus and L. rhodomystax have pale labial stripes. Leptodactylus stenodema differs from L. pentadactylus in coloration and in the disposition of the dorsolateral folds. In the former the dorsum is uni- form dull reddish brown, and the venter is grayish tan, whereas in L. pentadac- tylus the dorsum usually is orange or reddish tan with distinct transverse dark brown blotches, and the venter is cream mottled with l^rown. The dorsolateral fold is continuous with the supratym- panic fold in L. stenodema; the dorsolat- eral fold is medial to the supratympanic fold in L. pentadactylus. The dorsum is dull reddish brown, becoming orange-brown just medial to the black dorsolateral fold. The dorsal surfaces of the limbs are dark brown with indistinct narrow black transverse Fig. 73. — Leptodactylus stenodema, $ , 94 mm SVL (KU 122582). lines. The posterior surfaces of the thighs are black. The side of the head is brown, and the lip is orange-brown with four triangular black marks. The flanks are dull brown above and orange-brown below. The throat is dark brown, and the other ventral surfaces are grayish tan. The iris is bronze with a median horizon- tal red streak. Breeding males have nup- tial spines on the prepollex. i 94 mm. Occurrence. — The single adult male was sitting on the ground in secondary forest at night. Life History. — Nothing is known. Mating Call. — According to Pyburn and Heyer ( 1975 ) , the call is "a 'whoop' emitted while the frog sits in a burrow somewhat removed from the entrance." Leptodactylus wagneri (Peters) ( Figure 74 ) PIcctiomantis wagneri Peters, 1862, Monatsb. Akad. Wiss. Berlin, 1862:232 [Type local- ity.— West side of Andes of Ecuador]. Leptodactylus wagneri — Nieden, 1923, Das Tierreich, Berlin, 46:479. Material. — Lago Agrio, 6; Puerto Libre, 1; Santa Cecilia, 118-|-3. Identification. — This Leptodactylus is like Adcnomera andreae and L. disco- dactylus in lacking dorsolateral dermal folds; all other species of Leptodactylus in the area have dorsolateral folds. Lep- todactylus wagneri and L. discodactylus differ from Adenomera in having the EQUATORIAL HERPETOFAUNA 113 \^ .^y- -^ Fig. 74. — Lcptodactijhts loagneri. Upper: ju- venile, 19 mm SVL (KU 119343). Lower: $, 45mmSVL ( KU 122584). first finger longer than the second ( equal in Adenomera), and in having lateral fringes on the toes (absent in Adeno- mera). Leptodactylus discodoctyhis dif- fers from L. loagneri by having expand- ed, round, terminal discs on the digits and usually a dark throat and cream belly, whereas L. wap^neri lacks expand- ed terminal discs and has a cream throat and belly, both mottled with brown. Furthermore, L. luagneri is larger ( > 78 mm ) than L. discodactylus ( < 36 mm ) . The snout is rounded. The skin on the dorsum has numerous small spicules and many short elongate ridges laterally. The dorsum and flanks vary from dull olive-green to brown or gray with dark brown to black irregular spots or mot- tling. Usually there is a crudely triangu- lar-shaped mark, with the apex directed posteriorly, on the head. This is narrow- ly bordered by cream or pale gray. The lips are barred with cream and dark brown. The dorsal surfaces of the limbs are colored like the body and have in- distinct, irregular, darker transverse bars. The posterior surfaces of the thighs are black with greenish yellow to yellowish tan flecks or spots. The venter is dull creamy white with brown or dark gray mottling, most prominent in larger indi- viduals. The iris is dull bronze with a median horizontal grayish brown streak. Some adults of both sexes have promi- nent yellow glandular areas ventrolat- erally, and breeding males have two nup- tial spines on the prepollex. 6 59 mm; 9 77 mm. Occurrence. — LeptodactyJus wagneri was found in all months except Septem- ber and December through February. Although it was not found commonly in any given habitat, it occurred in a wide variety of habitats, exclusive of the river edge. One-third of the specimens were found in clearings, usually at night; other places of occurrence were marshes and swamps (25%), primary forest ( 12%), secondary forest (10%), streams (10%), and lakes (3%). Of eight specimens found by day, three were in small streams in forest and five were under cover in clearings. Life History — Calling males were heard in a forest swamp in July. Gravid females were found in March, May, and June at which times 40, 50, and 50 per- cent, respectively, of captured females were gravid. Non-gravid females were found in April, July, and August. Seven gravid females 51.0-77.0 (x=67.3) mm in snout- vent length contained 1000-3150 (x=l58S.6) ovarian eggs. The lightly pigmented eggs are deposited in foam nests in temporary ponds and ditches. Tadpoles. — The following description is based on KU 146773 ( stage 27, body length 12.0 mm, total length of 34.5 mm). The body is ovoid, and widest and deepest at midlength. In dorsal 114 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY view and in profile the snout is round. The eyes are moderately large, broadly separated, and directed dorsolaterally. The nostrils are about midway between the tip of the snout and the eyes. The spiracle is sinistral; the opening is di- rected posterodorsally at a point just be- low midline at about midlength of the body. The mouth is anteroventral and has shallow lateral folds. The median part of the upper lip is bare; there is one row of labial papillae laterally and two rows ventraly. There are two upper and three lower rows of denticles; the second upper row is narrowly interrupted me- dially. The beaks are moderately slender and weakly serrated. The dorsal and ventral caudal fins are equal in depth; each is about one-half the depth of the caudal musculatiue at midlength of the tail, the point of greatest depth of the fins. The dorsal fin does not extend onto the body, and the tip of the tail is acute- ly rounded. The body is dark brown with white flecks laterally; the tail is cream with brown flecks on the caudal musculature, dorsal fin, and anterior half of ventral fin. Tadpoles were found amidst dense vegetation in a shallow, weed-choked pond in March, and in a scmi-peiTnanent pond in May. Mating Call — The call consists of a series of soft, high-pitched notes: "oit, oit, oit." Analysis of one recording male at Belem, Brasil, reveals that the note repetition rate is 68 notes per minute with the dominant frequency at about 2700 Hz and emphasized harmonics at about 5400 and 8100 Hz. Food. — Twenty-five individuals con- tained a variety of insects, one gastropod, one chilopod, one crustacean, one spider, and one frog (Ht/la hrevifrom). Ants were eaten by 12 frogs, orthopterans by 11, and beetles by 9; these items consti- tuted 12, 27, and 14 percent of the bulk, respectively. The largest prey item was a centipede 47 mm long in a frog having a snout- vent length of 69 mm. Another individual having a snout-vent length of 54 mm had eaten a catepillar 33 mm long. Lithodytes lineatus (Schneider) ( Plate 1 ) Rana lineata Schneider, 1799, Hist. Amphib., 1:138 [Type locality. — Guyana]. Lithodytes lineatus — Fitzinger, 1843, Syst. Rept., Wien, 1:31. Material. — Lago Agrio, 3; Puerto Libre, 8; Santa Cecilia, 2. Identification. — This brightly colored frog with a black dorsum and yellow to tan dorsolateral stripes can be con- fused only with some dendrobatids, all of which have scutes on the dorsal sur- faces of the tenninal segments of each digit. Of these dendrobatids, Dendro- hates pictus and FhijUohates resemble L. lineatus in having dorsolateral light stripes, but both have mottled venters; in D. pictus the dorsolateral stripes are narrow, and a lateral white stripe is pres- ent in P. femoralis. Lithodytes has a large red inguinal spot and another red spot on the posterior surface of the thigh; red spots are lacking in the dendrobatids. The dorsum and flanks are black. A pair of broad dorsolateral stripes origi- nate on the tip of the snout and extend along the edge of the eyelid to the groin; the stripes are creamy yellow in juveniles and golden tan in adults. The dorsal sur- faces of the limbs are pale brown with dark brown transverse marks. A large red spot is present in the inguinal region and proximal anterodorsal surface of the thigh; another large red spot is present on the posterior surface of the thigh. The venter is grayish brown with creamy yel- low flecks laterally. The iris is deep bronze with brown bars. S 45 mm; $ 56 mm. Occurrence. — The terrestrial species apparently is active by day and night. Three were in primary forest, nine in secondary forest, and one at the edge of the forest. Eight were active on the ground by day and four by night; one was sitting on a log at night. EQUATORIAL HERPETOFAUNA 115 Life History. — Two females (May and July ) contained 195 and 230 ovarian eggs. The number of eggs is excessive in comparison with EJeutherodactyhis of similar body size; thus it seems unlikely that Lithodytes has terrestrial eggs with direct development. Perhaps they con- struct aquatic foam nests and have aquatic larvae. Mating Call. — The call is unknown. Food. — Nine individuals contained a variety of small insects. Ants were pres- ent in seven stomachs, fly larvae in three, coleopterans, hemipterans, and orthop- terans in two; and termites in one. Remarks. — The coloration of Litho- dytes sufficiently resembles that of the poisonous FhyUohates femoralis that the former could be considered a mimic and the latter the model. The resemblance is particularly striking in juvenile Litlio- dytes. Both species are terrestrial forest inhabitants from the Guianas to Peru; however, Lithodytes is active by day and night, whereas FhyUoJ)ates is diurnal. Physalaemus petersi (Jimenez de la Espada) ( FlGLTRE 75 ) Engijsiomops petersi Jimenez de la Espada, 1872, An. Soc. Espan. Hist. Nat., Madrid, 1:86 [Type locality. — "Oriente," Ecuador]. PJiysalaemus petersi — Lynch, 1970, Copeia, 1970(3):492. Material. — Puerto Libre, 5; Santa Ce- cilia, 13. Identification. — Resembling a small toad with a pointed snout, Fhysalaemtis petersi, difi^ers from Biifo by lacking cranial crests and parotoid glands and from Dendrophryniscus by having tu- bercles on the dorsum and a black and white mottled belly. The skin on the dorsum is bedecked with numerous, small, red or orange tu- bercles that are in contrast to the dull brown ground color. A single dark brown bar traverses the thigh and shank. Some individuals have small, dark brown spots on the dorsum. The flanks are creamy Fig. 75. — Physalaetnus petersi, S , 24 mm SVL; Mara, Piovincia Pastaza, Ecuador ( KU 120- 280). yellow anteriorly, becoming orange pos- teriorly in larger individuals, with large black spots; the anterior and posterior surfaces of the thighs are mottled black and cream. The posterior part of the belly and ventral surfaces of the thighs are white or gray with bold black mot- tling. The throat and chest are dark gray with minute white flecks and, usually, a narrow median cream line; some individ- uals have faint orange mottling on the throat and chest. The iris is dull bronze. 6 26 mm; 9 32 mm. Occurrence. — Although toads of the genus FhysaJaemus usually are nocturn- al, some observations at Santa Cecilia seem to contradict this generality. Eight individuals were found on the forest floor; two of these were amidst leaf litter by day, whereas the others were active at night. Seven individuals (juveniles and subadults) were found (sleeping?) on leaves of low herbs ( < 30 cm ) at night. All calling males (three) were found at night. M. L. Crump (pers. com.) found juveniles on low vegetation but adults breeding at night at Belem, Brasil, and J. D. Lynch (pers. com.) found males calling at night at Mera, Ecuador. The limited observations sug- gest a possible difference in diel activity cycles between adults and juveniles; at 116 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY least adults are active at night. About equal numbers of individuals were found in primary and secondary forest. Life History. — One gravid female col- lected in July contained 340 eggs. Males were heard calling only in July from shalow water in temporary ponds. Pre- sumably this species, like other Phijsa- loemtis, constructs a foam nest on the surface of ponds. Tadpoles. — No tadpoles were found at Santa Cecilia; the following descrip- tion is based on specimens (KU 121357) from Mera, Provincia Pastaza, Ecuador. A typical tadpole in developmental stage 28 has a body length 13.7 mm and a total length of 31.5 mm. The body is robust, widest posteriorly and about as deep as wide. The snout is narrow and rounded. The eyes are small, close together, and directed dorsolaterally. The nostrils are closer to the eyes than to the tip of the snout. The spiracle is directed postero- dorsally; the spiracular opening is at a point just below midline about one-third the length of the body. The caudal fins are deeper than the moderately robust caudal musculature, deepest at mid- length of the tail, and rounded at the tip; the dorsal fin does not extend onto the body. The mouth is anteroventral, about as wide as the anterior part of the body. The upper lip is bare; elsewhere there is one row of blunt labial papillae. The beaks are broad, slender, and weakly serrate. There are two upper and three lower rows of denticles; the upper rows are slightly longer than the lower rows, and the second upper row is narrowly interrupted medially. The ]:)ody and cau- dal musculatiu-e are pale brown with dark brown markings on the dorsum of the body and large brown spots on the tail. The dorsal markings usually consist of a pair of spots on the snout, a broad blotch posterior to the eyes and a narrow transverse bar posteriorly on the body. The caudal fins are translucent with brown flecks on the dorsal fin; flecks are absent in small tadpoles and most numer- ous in larger ones. Tadpoles have been found in shallow open ponds, where they seek refuge in the mud on the bottom. Matin<^ Call. — Males call while float- ing on the surface of the water. The call is a series of two or three notes: "doo- ing, doo-ing." Food. — Six individuals contained food: 7, 9, 9, 23, 43, and 55 temiites. Remarks. — The range of Physalaemus petersi has been poorly documented. I have examined specimens (KU) from many localities from Departamento Pu- tumayo, Colombia, southward through Ecuador and Peru to Departamento Co- chabamba, Bolivia, and from Belem, Brasil. BUFONIDAE Bufo glaberrimus (Giinther) ( Figure 76 ) Btifo glahcniinus Ciinther, 1868, Proc. Zool. Soc. London, 1868:483 [Type k)ca]ity. — "Bogota," Colombia]. Material. — Puerto Libre, 1; Santa Ce- cilia, 6. Identification. — This toad differs from all other Bufo in the area by lacking cranial crests and tubercles on the dor- sum. These characters, in combination with dark flanks and venter and pale dorsum, immediately distinguish Bufo glaberrimus from other toads and toad- like anurans in the area. The skin on the dorsum is smooth, except for scattered, low pustules in large individuals; the skin is smooth on the throat and chest and weakly granular on the belly and thighs. The parotoid glands are ovoid and about twice the length of the eyelid. The toes are about one-third webbed; the distal two and one-half segments of the fourth toe are not included in the web. The dorsum is orange-tan to olive-tan; the pustules are reddish brown in some individuals. The sides of the head and flanks are dark brown, and the venter is grayish brown with cream spots. The iris is pale gold with black reticulations. S 72 mm. EQUATORIAL HERPETOFAUNA 117 i,'* . _r - . * Fig. 76. — Bufo glaherrimus, 9, 40 mm SVL (KU 120339). Occurrence. — Ecological data are available for only three specimens. An adult male was in a hole in a clearing by day. One juvenile was under a log in a clearing by day, and one juvenile was at the edge of the river at night. I sus- pect that Bufo glaherrimus is like B. gut- tatus and inhabits more open forests than typical of the Santa Cecilia area. Life History. — Nothing is known. Mating Call. — The call is unknown. Food. — Three individuals had eaten 27 ants, eight beetles, one termite, one millipede, and one whip scorpion ( Pedi- palpida). Taxonomic Comments. — Stebbins and Hendrickson (1959:515) and Rivero (1961:20) treated Bufo glaherrimus as a subspecies of Bufo guttatus.. Schneider, but Cochran and Coin (1970:113) con- sidered it to be a full species. Too little material is available from the upper Am- azon Basin to detemiine the taxonomic status of the members of the Bufo gufta- tus complex occurring there. Bufo marinus (Linnaeus) ( FiGtmE 77 ) Rana marina Linnaeus, 1758, Syst. Nat., Ed. 10, 211 [Type locality. — "Americas"]. Bufo marinus — Schneider, 1799, Hist. Aniphib., 1:219. Material — Puerto Libre, 2; Puerto Ore, 3; Santa Cecilia, 49+8. Identification. — Adults of Bufo mari- nus can be distinguished from other toads in the area solely by their enor- mous size. Characteristically juveniles Fig. 77. — Bufo marinus, $, 125 mm SVL; Belem, Estado do Para, Brasil (KU 128735). and subadults have dark spots or mot- tling on the tubercular dorsum. The large trianguloid parotoid gland and low cranial crests distinguish B. marinus from other toads. Cranial crests are ab- sent in B. glaherrimus, as are warts on the dorsum. The supratympanic crests are greatly expanded dorsolaterally in adult B. typlionius. The crest between the eye and the tip of the snout is straight in B. marinus, whereas it extends pos- teriorly from the snout and then angles laterally to the anterior corner of the orbit in B. typhonius. The combination of cranial crests, tubercles on the dor- sum, and webbing between the toes dis- tinguish juvenile B. marimis from Den- drophryniscus minutus and small Isch- nocnema quixensis and Eleutherodacty- lus sulcatus. The skin on the dorsum is tubercular; that on the venter is granular. In adults the tips of the tubercles are keratinized. Adults have distinct, moderately low, keratinized cranial crests and huge, tri- anguloid parotoid glands that are twice the length of the eyelids and extend ventrally to the level of the jaw. The toes are more than one-half webbed; the distal two segments of the fourth toe are free of webbing. The dorsum varies from grayish tan to olive-brown or red- dish brown with or without small darker spots, which are indistinct or absent in lis MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY most large adults. The margin of the ujjper lip is creamy tan. The venter is creamy white with grayish brown spots. The iris is pale green suffused with black. Breeding males have horny nuptial ex- cresences on the thumb and in some indi- viduals the excrescences also occur on the second and third fingers. £ 127 mm; 9 146 mm. Occurrence. — Only two specimens were found in the forest; all others were in clearings and along the Rio Aguarico. Many individuals were observed nightly on the airstrip. By day juveniles were found beneath logs in clearings. Males called from small temporary pools in clearings or pools in the river bank left by the receding river. Life History. — Five of six adult fe- males contained mature ovarian eggs; these were found in March ( 1 ) , July ( 1 ) , August ( 2 ) , and October ( 1 ) . Females having snout- vent lengths of 115, 130, and 146 mm contained 4240, 9100, and 12,700 eggs respectively. Males call throughout the year, but most frequently they are heard on nights when there has been little or no rain. Eggs are deposited in long strings in shallow temporary pools in clearings and in pools in the river bank. Five recently metamor- phosed young from June have snout-vent lengths of 11.9-12.7 (x=12.4) mm. Tadpoles. — The following description is based on tadpoles in developmental stage 30 (KU 109186) having a body length of 10.3 mm and a total length of 26.0 mm. The body is ovoid, widest just behind the eyes and deepest at mid- length. The snout is bluntly rounded in dorsal view and round in profile. The eyes are moderately large, narrowly sep- arated, and directed dorsolaterals. The nostrils are at about 60 percent of the distance from the tip of the snout to the orbits. The sinistral spiracle is directed posterodorsally; the spiracular opening is on the midline at midlength of the body. The mouth is anteroventral and has lateral folds bearing one row of papillae; the upper and lower lips are bare. There are two upper and three lower rows of denticles, of which the second upper row is broadly interrupted medially. The moderately robust beaks are finely serrated. The tail is 60 per- cent of the total length, is acutely round- ed terminally, and has moderately low fins. The dorsal fin barely extends onto the body and is slightly deeper than the caudal musculature or ventral fin. The body and caudal musculature are black with small cream flecks on the tail; the caudal fins are gray. The iris is grayish bronze. Tadpoles were found in a rocky pool in the riverbed and in temporary ponds in clearings. Feeding aggregations of about 30-50 individuals were observed in shallow ( ± 10 cm ) water in a grassy pool. Matins, Call — The call consists of a low, rattling trill; in many cases, the call lasted more than 1 min. Food. — In addition to 7 millipedes (Diplapoda), the stomachs of 25 indi- viduals contained a variety of insects with large ants predominating (68% of stomachs), followed by beetles (Coleop- tera, 64%), orthopterans (40%), hemip- terans (8%), and lepidoptcran larvae (8%). Remarks. — Throughout its broad geo- graphic range, Biifo mar inns either avoids, or occurs in low densities, in trop- ical rainforests. However, it frequently occurs in large populations along river banks and clearings in otherwise forested areas. My observations on the species in Brasil, Colombia, Ecuador, Mexico, and Central America reveal that breeding always occurs in aquatic situations that receive direct sunlight. Perhaps the ther- mal requirements of the embryos are such that the shaded ponds in forests are unsuitable for embryonic develop- ment. Bufo tyijhonius (Linnaeus) ( Figure 78 ) Rana typhonia Linnaeus, 1758, Syst. Nat., 10: 211 [Type locality. — -"Americas"]. EQUATORIAL HERPETOFAUNA 119 Fig. 78.— Biifo ttjphonius, 9 , 83 mm SVL ( KU 123963). Bufo tiiplionius — Schneider, 1799, Hist. Am- phib., 1:207. Material — Lago Agrio, 9; Puerto Libre, 2; Puerto Ore, 0+3; Santa Cecilia, 244+7. Identification. — This moderate-sized toad with a pointed snout has an elon- gate elliptical parotoid gland about one and one-half the length of the eyelid. The dorsum is tuberculate. The cranial crests are high and expanded in adults. A crest extends posteriorly from the snout, angles laterally to the anterior corner of the orbit, and continues pos- teriorly as a supraorbital and supratym- panic crest, ending at the parotoid gland. In large adults the supraorbital crest is expanded dorsally and the supratym- panic crest is Hared dorsolaterally above the tympanum. A row of pointed tuber- cles extends from the posterior end of the parotoid gland posteroventrally to the groin. The shape of the parotoid gland, nature of the cranial crests, and the lateral row of tubercles distinguish this species from other Bufo in the area. Bufo gjaherrimus has smooth skin on the dorsum and lacks cranial crests; B. marinus has low keratinized cranial crests and lacks a lateral row of pointed tubercles. The dorsal ground color is tan, gray, or brown, plain or marked with irregular darker blotches and/or pairs of small chocolate brown spots. In most individ- uals the sides of the head and flanks be- low the row of tubercles are dark brown. Many individuals have a middorsal creamy white or yellow middorsal stripe extending from the tip of the snout to the rump. Some individuals have a nar- row cream labial stripe. The throat, chest, and anterior part of the belly vary from coppery brown to dark brown; the rest of the venter is yellowish or grayish tan with gray or black flecks. The iris is bronze with a narrow greenish gold bor- der to the pupil. Breeding males have horny nuptial excrecences on the thumbs. S 62 mm; $ 87 mm. Occurrence. — Although this terres- trial toad mostly inhabits primary forest (87% of 232 specimens), 26 were in sec- ondary forest, three in forest-edge situa- tions, and one in a clearing. Adults and juveniles are active on the forest floor by day. Of 66 individuals found at night, 62 were on sticks or leaves up to 1.5 m above the ground; many of these ob- viously were asleep. Two were on the ground at night, and two were calling from the ground at edges of a stream and the river at night. Life History. — Gravid females were found throughout the year, except in January, June, and September; however the absence of females from those months evidently is an artifact of sam- pling. Sixteen gravid females having snout- vent lengths of 58-87 (x=73.3) mm contained 765-2500 (x=: 1566.5) ma- ture ovarian eggs. Two others, having snout-vent lengths of 54 and 61 mm, con- tained 430 and 310 eggs, respectively. These small ovarian complements may be due to the small size of the females or they may represent only partial ovarian complements. The pigmented eggs are deposited in strings in pools in streams and in temporary pools in the forest. Six metamorphosing young from June have snout-vent lengths of 7.1-8.1 (x=7.6) mm. Tadpoles. — Tadpoles in development- al stage 41 have body lengths of 8.5 mm and total lengths of 23.5 mm. The body 120 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY is ovoid, widest and deepest at mid- length. The snout is bknitly rounded in dorsal view and round in profile. The eyes are medium-sized, moderately sepa- rated, and directed dorsolaterally. The nostrils are midway between the tip of the snout and the orbit. The spiracle is sinistral; its opening is directed postero- dorsally on the midline at midlength of the body. The caudal musculature is slender. The caudal fins are deeper than the musculature and are rounded termi- nally. The dorsal fin is deeper than the ventral fin and does not extend onto the body. The mouth is anteroventral and has lateral folds bearing one row of papillae. The upper and lower lips are bare. There are two upper and three lower rows of teeth; the second upper row is interrupted medially. The beaks are slender and finely serrate. The body and tail are brown; the fins are translu- cent. Mating Call — The call consists of 3-5 quickly repeated, low-pitched, rattling notes; calls are repeated at intervals of 10-15 seconds. Notes are 0.15-0.20 sec in duration and have a dominant frequency of about 1500 Hz. Food. — Ants occurred in all 25 stom- achs examined; among the 325 ants eat- en, many exceeded 17 mm in length. Thirty-five beetles had been eaten by 12 toads. Other food items were repre- sented by single prey and included an isopod, a millipede, a hemipteran, and an orthopteran. Taxonomic Comments. — Recently it has come to my attention that the toad that is widespread in tropical South America and called Bufo typhonius is a composite of perhaps three species. Through the courtesy of Charles W. My- ers, I have examined Peruvian specimens in the American Museum of Natural His- tory. Some of the Peruvian females have numerous pigmented eggs, whereas oth- ers have few unpigmented eggs. M. S. Hoogmoed (pers. com.) has discerned two morphological types in Surinam, dif- fering in l)ody proportions and the size and shape of the parotoid glands; both types have pigmented eggs. Ronald Al- tig (pers. com.) thinks he can distinguish two mating calls and corresponding dif- ferences in call sites in these toads in Amazonian Ecuador. I have not sub- jected the specimens from Santa Cecilia to a detailed morphometric analysis, but all females from Santa Cecilia have nu- merous pigmented eggs. Dendrophryniscus minutus (Melin) ( Figure 79 ) Atclopus minutu.s Melin, 1941, Goteborgs Ve- tensk Samh. Hand)., 4:18 [Type locality. — Taracua, Territoiio Amazonas, Brasil]. Dendrophryniscus minuius — McDiarmid, 1971, Bull. Los Angeles Co. Mus. Nat. Hist. Sci. 12:40. Material — Santa Cecilia, 157. Identification. — This small toad-like species has a depressed body, pointed snout, and small pointed tubercles on all dorsal surfaces. The absence of parotoid glands and cranial crests distinguish DendropJinjnisciis from Bufo. The ven- ter in Dendrophrijniscus is purplish red, a character that readily distinguishes the species from two other tubercular toad- like species - Physalaemus petersi (black and white venter) and Ischnocnema quixensis (cream venter with brown spots ) . The dorsum is pale brown, and the flanks are reddish tan; both have minute, pale blue flecks. The throat is grayish brown; the chest, belly, and ventral sur- faces of the thighs are purplish red or red with black or brown flecks. A white or pale yellow labial stripe continues onto the anterior surface of the upper arm. The iris is brown peripherally and gold medially. c5 16 mm; 9 24 mm. Occurrence. — Dendrophryniscus mi- nutus is active on the forest floor by day; individuals take refuge amidst leaf litter. At night they sleep on leaves of low herbaceous plants, usually less than 50 cm above the ground. These shade- loving frogs usually are found only in imdisturbed primary forest; only eight EQUATORIAL HERPETOFAUNA 121 Fig. 79. — Dendrophryniscus minutus, ? , 18 mm SVL (KU 146201). individuals were found in secondary for- est. Life History. — Gravid females were found throughout the year, except for October. Fifty-two females having snout- vent lengths of 18-22 (x=19.7) mm con- tained 70-245 (x=: 162.2) pigmented ovarian eggs. Eggs were not collected at Santa Cecilia; however Duellman and Lynch (1969:238) described eggs result- ing from amplectant pairs found at Sara- yacu, Provincia Pastaza, Ecuador. The pigmented eggs were in gelatinous strands. Tadpoles. — The following description is based on tadpoles in stage 31 from Sarayacu, Provincia Pastaza, Ecuador, having body lengths of 5.8 and 6.5 mm and total lengths of 14.1 and 15.4 mm. The body is ovoid, widest posteriorly and slightly wider than deep. The snout is round in dorsal view and in lateral profile. The eyes are moderately large, narrowly separated, and directed dorso- laterally. The nostrils are at a point about 60 percent of the distance from the tip of the snout to the orbits. The sinistral spiracular opening is directed posteriorly on the deadline at about two- thirds the length of the bod)'. The mouth is small and anteroventral with slightly infolded lips that bear a single row of small papillae laterally. The upper and lower lips are bare. There are two upper and three lower rows of denticles, all complete. The beaks are moderately slender and finely serrated. The tail is about 67 percent of the total length, deepest at midlength, and rounded ter- minally. The dorsal fin does not extend onto the body, is slightly deeper than the ventral fin and about equal in depth to the caudal musculatine at the mid- length of the tail. The body and caudal musculature are brown; small black flecks are present on the tail. Mating Call. — The call consists of two or three quickly repeated, short, high-pitched notes. Analysis of one re- cording reveals that the duration of each note is about 0.06 sec, and that a se- quence of three notes was produced in 0.4 sec. The dominant frequency is about 4200 Hz. Food. — The stomachs of 25 individ- uals contained 547 prey items, of which 519 were ants, a prey found in every stomach. Eleven individuals also had eaten 15 small beetles. Other incidental prey included two symphylids, two mites, and one each of the following: beetle larva, homopteran, hymenopteran, millipede, and phalangida. Remarks. — Most specimens from San- ta Cecilia have rather uniform purplish red venters; a few specimens have small creamy yellow spots on the posterior part of the belly. All specimens exam- ined from south of the Rio Napo have large yellow or orange spots on the belly. Dendrobatidae Colostethus marchesianus (Melin) ( Plate 1 ) Pliyllobates maiciicsianus Melin, 1941, Gote- l^oigs Vetensk. Samh. Handl, 4:64 [Type locality. — Taracua, Tenitoiio Amazonas, Brasilj. Colostethus marchesianus — Edwards, 1971, Proc. Biol. Soc. Washington, 84:148. Material. — Puerto Libre, 2; Santa Ce- cilia, 93+3. Identification. — This small frog with 122 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY scutes on the dorsal surfaces of the ter- minal segment of each digit is nearly uni- form brown above with black Hanks bor- dered below by a white stripe. It is most easily confused with Colostethus sauli, a larger frog with dark brown, usually transverse markings on the dorsum and a diagonal yellow stripe on the flank. Other dcndrobatids have black and white or black and blue mottling ventrally. The dorsum is dull brown usually with a reddish or olive tint; dorsolater- ally, the color is paler, giving the indica- tion of an indistinct tan stripe in some individuals. Small dark brown or black flecks are present middorsally in a few specimens. The limbs are pale brown with dark brown transverse markings; in most individuals a pale yellow or yellow- ish tan spot is evident on the proximal upper surface of the arm. The loreal re- gion and flanks are dark brown or black. The upper lip is cream; a narrow white stripe extends from the angle of the jaw to the groin. A narrow area below the stripe is mottled tan and cream. The belly is creamy white to pale yellow, and the ventral surfaces of the limbs are gray- ish white. In males the vocal sac is pale yellow. The iris is black with bronze flecks. $ 18 mm; 9 18 mm. Occurrence. — CoIostefJius marchesi- aniis is equally abundant in primary and secondary forest, where it is active on the ground by day. Periods of greatest activity are early in the morning or late afternoon, especially after showers. The frogs take shelter in crevices under roots of trees, in stilt palms, or amidst leaf litter. This species is not associated with streams, as are most other species of Colosfetlius; instead individuals are widely dispersed in the forest. Males call from the ground or, more frequently, from slight elevations, such as a stick, large leaf, or base of a tree. At night individuals sleep on leaves of low herbs, usually less than 25 cm above the ground. Life History. — Gravid females were found in every month except September, December, and January - all months in which no females were collected. Evi- dently the species breeds throughout the year. Ovarian complements were 7-27 (x=13.4, N=z50). Two males were found carrying tadpoles; one in February had three tadpoles on its back, and one from July had ten. Tadpoles. — Free swimming tadpoles in stage 30 (KU 109314) have body lengths of 7.7 mm and total lengths of 20.7 mm. The body is ovoid, widest me- dially, shallower than wide, and deepest posteriorly. The snout is bluntly round in dorsal view and round in profile. The eyes are moderately large, widely sepa- rated, and directed dorsolaterally. The nostrils are midway between the tip of the snout and the orbits. The opening of the sinistral spiracle is directed dor- sally at the midline at about three-fifths the length of the body. The mouth is directed anterovcntrally and has small lateral folds. The upper lip is bare; else- where around the mouth there is a single row of small labial papillae. There are two upper and three lower rows of den- ticles. The first upper row is on the mar- gin of the upper lip. The second upper row is broadly separated medially. The tail is about 63 percent of the total length. The caudal musculature is ro- bust, its depth at the midlength of the tail being equal to twice the depth of the dorsal fin. The dorsal and ventral fins are about equal in depth, deepest at about midlength of the tail. The dorsal fin does not extend onto the body, and the tip of the tail is round. The body is brown above and below; the tail is cream with brown blotches, and the iris is bronze. The only free swimming tapoles were found in a water-filled segment of a fallen bamboo in March. One metamorphosing individual having a short tail stub had a snout-vent length of 7.0 mm. Mating, Call. — The call consists of a soft single "peep." Food. — In 25 stomachs, ants (25) and small beetles (17) were the most abun- dant prey items, followed by termites EQUATORIAL HERPETOFAUNA 123 (8). Tlie remaining items (Diptera, Hy- menoptera, Oithopteia, and spiders) ac- counted for only 12 percent of tlie items and 20 percent of the volume of food eaten. Colostethus sauli Edwards ( Plate 1 ) Colostethus sauli Edwards, 1974, Occas. Papers Mus. Nat. Hist. Univ. Kansas, 30:10 [Type locality. — Santa Cecilia, Provincia Napo, Ecuador]. Material. — Lago Agrio, 3; Santa Ce- cilia, 19+6. Ide n tificatio n . — T his d u 1 1 - c o 1 o r e d dendrobatid is most easily confused with Colostethus marcliesianus, a smaller frog with a yellow or white venter and a white ventrolateral stripe and no large dark dorsal markings; C. sauli has a gray venter, no white ventrolateral stripe, and dark dorsal marks. Of the other pre- dominately brown dendrobatids in the area, Dendrohates pictus has a narrow white canthal and dorsolateral stripe, and Phyllohafes femoralis has a broad cream canthal and dorsolateral stripe; both have mottled venters. The dorsum is olive-brown to olive- gray with irregular, but usually trans- verse, black markings. The limbs are brown with distinct dark brown trans- verse bars and a pale yellow spot proxi- mally on the upper ami. The side of the head and the flanks are black. A narrow diagonal stripe extends posteroventrally on the flank to the groin; anteriorly the stripe is yellow, changing to bluish gray posteriorly. The anterior and posterior surfaces of the thighs are mottled with shades of gray. The ventral surfaces are uniform pale gray. The iris is grayish bronze. $ 22 mm; 9 26 mm. Occurrence. — Unlike tthe other den- drobatids in the area, this species most frequently occurs near small streams. At two small hillside springs at Santa Ce- cilia, the frogs were found in shallow water and under overhanging banks. Other individuals were found near the streams issuing from the springs, and a few were widely dispersed in the forest. Those individuals in the vicinity of streams took refuge in the water, where they would either sit on the bottom or hide beneath submerged objects. Life History. — Of nine adult females ( March 2, May 2, July 4, and August 1 ) , six contained 6-14 (x=11.3) large ova- rian eggs; one from July had small ovarian eggs, and two from July showed no evidence of reproductive activity. Tadpoles. — Free-swimming tadpoles in stage 36 (KU 109322) have body lengths of 7.8 mm and total lengths of 24.2 mm. The body is elongately ellipti- cal in dorsal view, widest posteriorly, shallower than wide, and deepest pos- teriorly. The snout is round in dorsal view and in profile. The eyes are large, widely separated, and directed laterally. The nostrils are about 60 percent of the distance from the tip of the snout to the orbits. The opening of the sinistral spir- acle is directed posterodorsally on the midline at about three-fifths of the length of the body. The mouth is directed an- teroventrally and has small lateral folds. The upper lip is bare; one row of small labial papillae are present laterally and ventrally. There are two upper and three lower rows of denticles, of which the second upper row is widely separated medially. The beaks are slender and finely serrated; the upper beak is deeply notched. The tail is about 68 percent of the total length. The caudal muscula- ture is robust and at midlength of the tail is about thrice the depth of the dorsal fin. The tip of the tail is acutely rounded. The body is black with white flecks, and the tail is black with white mottling. Tadpoles were found amidst detritus in shallow streams issuing from springs in March and June. One metamorphos- ing young had a snout-vent length of 7.2 mm. Mating Call. — The call consists of a series of two or three quickly repeated peeps. Food. — In eight stomachs, ants (9) and small homopterans (7) were the 124 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY only prey items found more than once; other food items were a small beetle (7 mm), beetle larva (11 mm), wasp, and grasshopper nymph. The largest homop- teran was 8 mm long. Dendrobates parvulus Boulenger ( Plate 1 ) Dendrobates parvulus Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.: 14.5 [Type locality.— Saiayacii, Pro\incia Pa,staza, and Loreto, Piovincia Napo, Ecuador]. Material. — Lago Agrio, 15; Puerto Libre, 17; Puerto Ore, 1; Santa Cecilia, 254+1. Identificatio7i. — This small dendro- batid with a red, granular dorsum and a bright yellow spot at the base of each limb cannot be confused with any other frogs in the area. Although other species of Dendrobates also have black and blue mottled venters, all have some dorsal stripes and none is red. The dorsum of the body is a unifonn dull red; in some specimens there is a suffusion of gray or dull brown, especial- ly posteriorly. The dorsal surfaces of the limbs are bluish black to dark gray with or without a brownish tint; pale blue or white flecks are evident on the limbs in most individuals. The flanks are blue with black mottling, but in some individ- uals the black is so extensive as to give the appearance of a black flank with scattered blue spots. The side of the head is black; a narrow bluish white labial stripe usually extends onto the an- terior edge of the upper arm. Each indi- vidual has a bright yellow spot on the proximal posterodorsal surface of the upper arm and another spot on the proxi- mal anterodorsal surface of the thigh. A bright yellow spot is present on the proximal ventral surface of the shank in most individuals. The throat, belly, and ventral surfaces of the limbs are blue with black spots and mottling. The iris is dark brown to black with minute bronze flecks and a narrow bronze ring around the pupil. S 20 mm; 9 23 mm. Occurrence. — Dendrobates parvulus is one of the most abundant and conspic- uous frogs living on the forest floor; it is especially abundant (or at least most easily observed) in primary forest. The frogs are most active by day — early in the morning or after afternoon showers; diurnal retreats include the leaf litter, cavities beneath logs, and bases of stilt palms. At night the frogs sleep on leaves of herbs and ferns, usually less than 50 cm above the ground. Males call from sticks, logs, or other slight elevations; usually the calling site is hidden beneath plant cover. Life History. — Gravid females were found in every month. Of 169 females, only seven (from five scattered months) showed no evidence of reproductive ac- tivity, and only 10 (March- June) had only small ovarian eggs. Of the 152 fe- males having large ovarian eggs, 78 were from March-June. The ovarian comple- ment is 4-16 (x=9.0, N=:152). two males carrying five and six tadpoles were found in July. Tadpoles. — Four back-riding tadpoles have body lengths of 4.0-4.4 (x=4.2) mm and total lengths of 12.5-13.8 (13.2) mm. The beaks are thin, and no denti- cles are present. One individual was placed in water and preserved 17 days later; it had better developed beaks, one row of labial papillae laterally and ven- trally, and two upper and two lower rows of denticles (second upper row broadly interrupted medially). The body is dark brown, and the tail is tan with dark brown flecks. Mating Call — The call of Dendro- bates parvulus consists of two soft peeps, the second higher pitched than the first. Food. — Ants were present in each of 23 examined; 120 ants comprised 65 per- cent of the volume. The remaining prey items were small beetles (14), beetle larvae ( 2 ) , fly larvae ( 2 ) , mites ( 2 ) , and a termite (1). The largest prey item was a fly larva 8 mm long. Taxonomic Comments. — Silverstone EQUATORIAL HERPETOFAUNA 125 (1975) placed this species in the genus Phyllobates because of the presence of maxillary teeth (see Savage, 1968, for generic definitions). Chares W. Myers (pers. com.) stated that the skin toxins and behavior of parvtihis definitely ally the species with Dendrohates. Dendrobates pictus pictus Dumeril and Bibron ( Plate 1 ) Dendrohates pictus Dumeril and Bibron, 1841, Erpt'tol. Gen., 8:656 [Type locality.— "Chile"]. Dendrohates pictus pictus — Lutz, 1952, Mem. Inst. Oswaldo Cruz, 50:612. Material. — Lago Agrio, 1; Puerto Libre, 4; Santa Cecilia, 116. Idenfificafion. — This dull greenish gray to dark brown dendrobatid with a black and blue venter can be distin- guished from all other dendrobatids and Lifhodytes lineatiis in the area by the presence of a narrow white stripe across the tip of the snout; the stripe continues posteriorly in a dorsolateral position to the groin. FlujUohates fe- moralis and Lifhodytes lineatus have broader, cream or tan dorsolateral stripes; P. femoralis also has white stripe on the flank, and L. lineatus has bright red spots on the thighs and pale gray venter. Dendrohates pictus lacks a stripe on the flank and red spots on the thighs. The dorsum is predominately dark brown with a greenish or grayish tint. Scattered, small black spots usually are evident on the back. The limbs are brown with indistinct dark brown cross- bars. A narrow white or creamy white transverse stripe on the snout is continu- ous with a dorsolateral stripe along the canthus, edge of upper eyelid, and body to the groin. A white labial stripe ex- tends from the lip just anterior to the orbit onto the anterior edge of the upper arm. The side of the head and flanks are black. A small yellow spot is present in the axilla, another on the proximal ven- tral surface of the shank. The throat, belly, and ventral surfaces of the limbs are blue, heavily mottled with black. The iris is dull bronze. ^ 21 mm; 5 24 mm. Occurrence. — Most specimens were found in secondary forest sleeping on leaves of low herbs at night. Others were active by day on the ground in both primary and secondary forest. Eight were sleeping at night on herbs in a marsh adjacent to forest. Life History. — No individuals were found in January or April; gravid females were found in all other months, except December, in which the only female found showed no evidence of reproduc- tive activity. Such individuals comprise only 13 percent of the females and are from seven months from March through December. Ten percent of the females had only small ovarian eggs; these are from FebiTiary, June, July, and October. The evidence overwhelmingly indicates that Dendrohates pictus breeds through- out the year. The ovarian complement is 10-31 (x=18.6, N=53) eggs. Tadpoles. — Tadpoles were described by Lescure ( 1976 ) . Mating CaU. — The call is a soft "tick- tick," or "tick-tick-tick." Food. — Ants were present in all but two of 26 stomachs examined; 110 ants comprised 60 percent of the volume of food. Beetles (16) and temiites (6) were the next most common prey items. Other prey items and their frequency of occur- rence are: Hemiptera (3), fly larvae (3), Diptera (2), small annelids (2), and one each of the following: Dermaptera, Hy- menoptera, coleopteran, larvae, and pseudoscorpion. Taxonomic Comments. — Silvertone (1975) placed this species in PhyUo- hates- 1 retain the species in Dendrohates pending forthcoming biological defini- tions of the genera by Charles W. Myers. Dendrobates quinquevittatus Steindachner ( Plate 1 ) Dendrohates tinctorius var. quinquevittatus 126 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Steindachner, 1864, Verb. Zool.-Bot. Gesell. Wien, 14:260 [Type locality. — "Brasil and Colombia"]. Material. — Lago Agrio, 5. Identification. — This small dendro- batid has a black dorsum with five broad, bright yellow stripes that readily distin- guish it from all other frogs in the area. The dorsum is jet black. Five broad, longitudinal, bright yellow stripes are arranged as follows: middorsal stripe from top of head to midpoint on body; dorsolateral stripes from snout or canthal region, along edge of eyelid to groin; lateral stripes extending along lip, above insertion of arm, and along the flank to the groin. The middorsal and dorsolat- eral stripes are variously interconnected by transverse yellow stripes. The dorsal surfaces of the limbs are yellowish tan with black spots. The throat is yellow, and the belly and ventral surfaces of the limbs are blue. All ventral surfaces are marked with large black spots. The iris is dull brown. S 14 mm; 9 16 mm. Occurrence. — All specimens were found in April and May on the ground in primaiy forest at Lago Agrio. Despite its small size, DenchoJ)ates quinqucvitta- tus should have been found at Santa Cecilia. No explanation is evident for the apparent absence of the species at Santa Cecilia. Life History. — Two females collected in May have 4 and 9 large ovarian eggs. No other data are available. Tadpoles. — Tadpoles are unknown. Mating Call. — The call is unknown. Food. — One individual contained 32 small ants. Phyllobates femoralis (Boulenger) ( Plate 1 ) PrustJicrapis femoralis Boiik'n^cr, 1883, Pioc. Zool. Soc. London, 1883:635 [Type local- ity.— Yurimaguas, Dcpartanicnto Loieto, Peru]. Phyllobates femoralis — Parker, 1935, Proc. Zool. Soc. London, 1935:506. Material. — Lago Agrio, 4; Puerto Ore, 1; Santa Cecilia, 44+2. Identification. — This large dendro- batid has a brown dorsum with cream or tan dorsolateral stripes. The presence of a white stripe on the posterior part of the flank distinguishes P. femoralis from all other dendrobatids in the area, except Colostcthus .sauli. The latter species lacks dorsolateral stripes and has a gray venter; the venter is mottled blue and black in P. femoralis. Superficially, P. femoralis resembles Dendrohates pictiis and Lithodytes lineatus. The former has narrow white dorsolateral stripes and no white stripe on the flank, and the latter has bright red spots on the thighs, no white stripe on the flank, and a gray venter. The dorsum varies from dark brown or nearly black to olive-brown or reddish brown; in some paler individuals bronze flecks are evident. A dorsolateral stripe extends along the canthus and edge of the eyelid to the groin; the stripe varies from silvery white to cream or tan. A narrow white labial stripe extends along the lip to the base of the arm. A longi- tudinal bluish white to pale yellow stripe extends from the midflank to the base of the thigh. The side of the head and flanks are black. The forearm is tan with black spots, and the hind limb is mottled reddish tan and black with bluish white flecks. A pale yellow spot is present on the proximal dorsal surface of the arm, and another is present on the proximal anterior surface of the thigh. The throat and chest are black, and the other ven- tral surfaces are blue heavily mottled with black. The iris is brown. $ 25 mm; 9 29 mm. Occurrence. — Individuals were found in equal abundance in primary and sec- ondary forest, where they are active on the ground by day. The frogs are widely dispersed in the forest and not associated with water. Males call from secluded sites early in the morning and late in the afternoon, especially after showers. Life History. — No females were found in January, February, September, or De- EQUATORIAL HERPETOFAUNA 127 cember. Gravid females were found in all other months. \\'ith the possible ex- ception of the period December-Febru- ary, breeding evidently occurs through- out the year. The ovarian complement is 8-43 (x=22.4, N=15) eggs. One male was carrying tadpoles in May. Tadpoles. — One back-riding tadpole (KU 125893) has a body length of 3.6 mm and a tail length of 7.4 mm. There is a single row of labial papillae laterally and ventrally, and two upper and three lower rows of denticles; the second up- per row is broadly interrupted medially. The dorsum is pale brown with yellow- ish tan Hecks, and the rest of the tadpole is translucent with brown flecks on the tail. Mating, Call. — The call of this species is a series of high-pitched, quickly re- peated pairs of notes: "peep-peep, peep- peep, peep-peep." Food. — Twenty-one individuals had eaten a variety of small insects and spi- ders. Ants were the most common food item retrieved from stomachs, followed by Coleoptera, Hymenoptera, and spi- ders; five other orders ( Lepidoptera, Isoptera, Hemiptera, Diptera, and Orth- optera) also were present. The largest prey taken were beetles; one 8 mm long was eaten by a frog 26 mm in snout-vent length, and one 7 mm long by a frog 27 mm in length. Hylidae Hemiphractus proboscideus (Jimenez de la Espada) (Figure 80) Cciafht/la prohoscidea Jimenez de la Espada, 1870, Jour. Math. Phys. Nat., Lisboa, 3:64 [Type locality. — Siimaco, Pro\incia Napo, Ecuador]. Hemiphractus proboscideus — Tiiieb, 1974, Oc- cas. Pap. Mus. Nat. Hist. Univ. Kansas, 29:38. Material. — Lago Agrio, 1; Puerto Libre, 5; Santa Cecilia, 46. Identification. — This bizarre frog with a fleshy proboscis, demial "horns" on the Fig. 80. — Hemiphractus proboscideus, $ , 58 mmSVL (KU 124139). eyelids, bony posterolateral projections on the head, and depressed body with protruding neural spines of the vertebrae is entirely unlike any other frog known from the area. Any damn fool should be able to identify this species. Cerato- phnjs cornuta is the only other species having demial "horns" on the eyelids, but that species has a robust body and lacks a fleshy proboscis. In dorsal view, the head is triangular with the posterior margin elevated and concave, the posterolateral corners being formed by three converging body ridges : one posterolaterally from the orbit, one dorsally from the angle of the jaw, and one transversely across the back margin of the skull. Bony canthal ridges diverge from a common point just posterior to the nostrils, extend to the orbits and join the supraorbital ridges, which extend posteromesially to the transverse occipi- tal ridge. A laterally compressed, fleshy proboscis projects from the tip of the snout. A single large dermal tubercle, bordered by several small tubercles, is present on each eyelid. A small triangu- lar calcar is present on the heel. The fincfers are not webl:)ed, and the toes are webbed basallv. At night the dorsum is yellowish tan with brown transverse or diagonal marks on the body and trans- verse bars on the limbs; the venter is grayish cream to white. By day the dor- 128 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY sum is brown with darker brown mark- ings and the venter is brown to reddish brown with orange or yellowish blotches on the belly and yellow or white spots on the throat. The discs on the first and second fingers are pale orange, changing to white in preservative. The tongue is orange. The iris is pale gray, with a pinkish tint in some individuals, with difi'use darker gray triangular marks an- teriorly and posteriorly. 9 50 mm; S 67 mm. Occurrence. — Ecological data are available for 39 sj^ecimens, of which 35 were found in primary forest; the others were in secondary forest. One was in leaf litter by day, and the others were perched on low vegetation (usually leaves or palm fronds less than 1.5 m above the ground) at night. Observa- tions on this and other species of Hemi- phractus (except scutatus) indicate that they remain on the forest floor by day, where their coloration and disruptive form provide excellent concealment amidst leaf litter. At night they ascend low vegetation. The presence of both diurnal and nocturnal prey in their stom- achs reveals that they feed by day and night. The distribution of specimens from throughout the year is uneven: March (4), April (5), May (4), June (17), July (15), August (5), November a). Life History. — Females of this species brood eggs on the back, where they un- dergo direct development into small frogs. No brooding females were found. In fact only one gravid female was dis- covered; that individual was found in July, and it contained 26 ovarian eggs 2.5 mm in diameter. Juveniles having snout- vent lengths of 16 mm and 28 mm were found in the same area of forest on 29 March and 15 June 1972, respectively. On 2 April 1972 four juveniles having snout-vent lengths of 18-19 mm were found in an area of about 20m-; on 10 May 1972 another juvenile having a snout-vent length of 23 mm was found at the same place. These five frogs may be part of one brood. Matinfi Call. — Apparently this species does not have a mating call; one male emitted a "grunt" when picked up. Food. — Hemiphractiis prohoscideus is a carnivore with a prodigious appetite. Ten of 13 stomachs contained 15 frogs, including EleutlierodactyJiis croceoin- giiinis (1), E. diademofus (1), £. lantlw- nifes (2), E. variabilis (1), E. sp. (1), Leptodactyhis mystaceus (1), dendro- batid (1), Hyla fasciata (1), H. granosa (2), //. sp. (1), Osteocephahis Jeprieurii (1), and unidentified frog (2). Inverte- brate prey included one each: leech, annelid worm, homopteran, and mole cricket (Gryllotalpidae). Some of the prey items were nearly as long as the frogs that had eaten them. Remarks. — Some individuals, when disturbed, open their cavernous mouths exposing the bright yellow tongue. Sub- sequent prodding by a careless observer can result in a quick snap of the jaws in a vice-like grip. In 1967 and 1968 Hcmiphractus was reasonably abundant at Santa Cecilia. In- creasing disturbance of the forest seemed to result in the disappearance of these frogs except in deep primary forest. Conceivably Hcmiphractus johnsoni Noble and //. scutatus (Spix) occur in the environs of the Rio Aguarico. The latter species has a broader head with shorter posterolateral horns and no no- ticeal)ly expanded discs on the digits; //. johnsoni also is more robust than H. prohoscideus and has a much smaller proboscis ( see Trueb, 1974, for diagnoses of the species ) . Hyla alboguttata Boulenger Htila alho^iiltata Boulenffer, 1882, Cat. Batr. ' Sal. Brit. Miis.: 385 [Type Ideality.— Sara- yacu, Pio\inda Pastaza, Ecuador]. Material. — Santa Cecilia, 153. Identification. — Although this dull brown Hyht could be confused with sev- eral other brown frogs in the area, it is distinguishable by several characters. EQUATORIAL HERPETOFAUNA 129 The absence of webbing between the fingers and/or the absence of flesliy cal- cars on the heels distingnish H. aJlyogut- tata from H. hoans, colcarofa, fasciata, and g^cogniphica, all of which have dark markings on the flanks and posterior sur- faces of the thighs ( absent in H. alhogiit- tata). Hijla alhoguttoia shares the ab- sence of webbing on the hand and re- duced webbing between the first and second toes with H. cruentomma, fii- nerea, g,arhei, and rubra. The latter has yellow spots on the posterior surfaces of the thighs; H. garhei has a pointed snout, tubercles on the heels, and vertical black and yellow bars on the posterior surfaces of the thighs. Hyla cruentomma and ju- nerea have dark brown longitudinal marks on a tan or olive-tan dorsum. Ju- veniles of H. alhoguffata and Osteoceph- alus Jepreurii have broad, creamy white dorsolateral stripes. The juveniles of O. leprieurii have dark transverse bars on the limbs; these are absent in //. albogut- tata. The head is broad and flat. Webbing is absent on the hand, and the toes are only about one-half webbed. The skin on the dorsum is smooth; that on the venter is granular. The tympanum is large. The dorsum is tan, usually marked with small, irregular dark brown or black spots; the spots are most numerous mid- dorsally on the posterior half of the body. Dark brown spots are present on the dorsal surfaces of the shanks. The flanks and anterior and posterior surfaces of the thighs are creamy tan. The throat and belly are white with minute, reddish brown flecks, and the ventral surfaces of the limbs are dull brown. The edge of the upper lip is cream. Juveniles ( < 25 mm snout- vent length) have broad, creamy white dorsolateral stripes that are narrowest on the head and widest in the scapular region. On the head the stripes are bordered below by dark brown canthal and supratympanic stripes. In specimens 25-30 mm in snout- vent length, the dorsolateral stripes and the dark stripes became diffuse, and in most specimens having snout-vent lengths of more than 30 mm, there is no evidence of stripes. The iris is greenish gold with bold black reticulations. Breeding males have brown horny nup- tial excrescences on the thumbs. S 34 mm; 9 46 mm. Occurrence. — Although Hyla alho- guttafa was found in a wide variety of habitats including clearings, banana groves, swamps, and primary forest, 75 percent of the individuals were in second growth forest, where they were found on low vegetation (< 1.5 m) at night (2% were at heights of more than L5 m) throughout the year. Juveniles and sub- adults were found during the day in un- rolling Heliconia leaves. Life History. — Despite the abundance of this species, no l^rceding or calling were observed. Females with mature ovarian eggs were found in every month, except December. Twenty-one gravid females with snout-vent lengths of 35.0- 46.0 (X=:41.4) contained 218-870 (X=: 440.1) eggs. The eggs are small (1.4 mm) and pigmented. The smallest ju- venile, taken in July, has a snout-vent length of 10 mm. Tadpoles. — The tadpoles are im- known. On the basis of the number and size of ovarian eggs, it is presumed that Hyla alhoguttata deposits eggs in stand- ing water. Mating Call. — The call is unknown. Food. — Nine arthropod orders were represented in 25 stomachs. Orthopter- ans were the most common prey (48% of stomachs), followed by dermapterans (24%), beetles (16%), spiders (16%), and lepidopteran larvae (12%). Other prey items were found in less than 10 percent of the stomachs. Remarks. — Field investigations on this abundant species have been frus- trated by the apparent absence of activ- ity by the frogs. Despite the collection of 153 specimens and observations on at least twice that many individuals in the field, no frog was observed to do more than sit on a leaf or branch. 130 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Hyla bifurca Andersson (Plate 2) Hi/la (Hi/leUa) hi f urea Andersson, 1945, Ark. ' Zool. " Stockholm, 37A(2):79 [Type local- ity.— Rio Pastaza, Provincia Pastaza, Ecua- dor]. Hyla hifuica — Duellman, 1974, Occas. Papers Mus. Nat. Hist. Univ. Kansas, 27:15. Material. — Puerto Ore, 31; Santa Ce- cilia, 96+14. Identification. — This .small brown frog with cream dorsal markings can be confused only with other members of the Hyla leucopJu/IJata group, all of which have axillary membranes and external tympana. Hyla J)ifurca is dark brown with the top of the head anterior to the eyes cream; beginning on the eyelids is a pair of parallel, straight-edged, nar- row, dorsolateral cream stripes extending posteriorly at least to the sacrum. A small cream spot is present on the rump and on each heel. The dorsal surfaces of the shanks are uniform brown, the flanks, hidden surfaces of the thighs, and web- bing on the feet are orange brown. Hyla bifurca is most likely to be confused with H. leucophyllata and H. sarayaciiensis, both of which have orange webs and hidden surfaces of the thighs. In Hyla leucophyllata the dorsolateral stripes are broad, constricting the middorsal brown area in the scapular region, thereby forming an hour-glass shaped, middorsal dark mark; the dorsal surfaces of the thighs are predominantly tan or cream. The dorsal markings in H. sarayacuensis are similar to those in H. bifurca, but in the former the dorsolateral stripes are broad, have irregular edges, and extend posterolaterally; furthermore, irregular transverse cream markings are present on the shanks. Hyla favosa also has orange Hash colors, but the dorsum is dark brown with a network of narrow cream lines. The flash colors are pink or red in H. triangulum, and the dorsum is cream to tan with or without one or more dark brown spots. The snout is broadly rounded in dor- sal view and truncate in profile. An ex- tensive axillary membrane extends nearly to the elbow. The fingers are webbed basally, and the toes are about two-thirds webbed. The skin on the dorsum is smooth, and that on the belly is granular. At night the dorsum is pale brown, with or without minute black flecks; the snout, dorsolateral lines, and spots on the rump and heels are yellowish tan. The hands, feet, axillae, and ventral surfaces of the limbs are orange. Of 111 specimens the rump spot is discrete in 83, connected with one of the dorsolateral lines in 15, and connected with both lines in 13. The vocal sac is yellow; males lack horny nuptial excrescences. The iris is grayish bronze. S 28 mm; 9 35 mm. Occurrence. — No individuals were found in primary forest, but 9 (12%) were found in secondary forest and 11 (14%) in forest-edge habitats. These individuals were perched on leaves or branches of herbs or low bushes; five frogs were on vegetation higher than 1.5 m above the ground. Most of the frogs were found at breeding sites in non-forested areas - temporary ponds or water-filled ditches in clearings or at the edge of a lake. None was found by day. Life History. — Hyla bifurca seems to be an opportunistic breeder. Males call after rains throughout the year, and usu- ally perch on emergent vegetation within 30 cm of the water. Gravid females were found in all months, except June and December through February. Twenty gravid females having snout- vent lengths of 29.0-35.0 (X=:32.9) mm contained 145-255 (X= 190.0) eggs. Fifteen clutches contained 153-255 (X=191.0) eggs, 2.0 mm in diameter. The heavily pigmented eggs are deposited on the upper surfaces of leaves of emergent plants, usually within 30 cm of the water. Tadpoles. — The following description is based on tadpoles (KU 146785) in developmental stage 32 having body lengths of 9.5 mm and total lengths of EQUATORIAL HERPETOFAUNA 131 31.0 mm. In dorsal view, the boch^ is violin shaped with a truncate snout; in lateral view, it is ovoid and deepest pos- teriorly with an acutely rounded snout. The e)'es are large and situated laterally. The nostrils are about midway between the tip of the snout and the orbits. The opening of the sinistral spiracles is di- rected posterodorsally on the midline at about midlength of the body. The mouth is anterior and lacks lateral folds. The median part of the upper lip is bare; elsewhere there is one row of large, blunt labial papillae. The beaks are mod- eratelv robust and finelv serrated. Den- tides are absent. The tail is about 70 percent of the total length. The fins are moderately deep, the depth of the cau- dal musculature at midlength of the tail being about 75 percent of the depth of the dorsal fin, which does not extend onto the body. The tail tenninates in a long filament. The dorsum of the body is tan with broad black lateral stripes bordered below by a metallic cream stripe. The belly is silvery white with three longitudinal black stripes. The caudal musculature and distal part of the dorsal fin are black; a cream stripe is present on the proximal part of the cau- dal musculature. The tip of the tail and other fins are cream, with black flecks on the dorsal fin. The iris is reddish bronze. Tadpoles have been found in shallow, vegetation-choked, temporary pools and ditches. Although individual tadpoles have been observed near the surface in open water, upon being disturbed they seek refuge amidst the vegetation. Mating Call — The call consists of a harsh primary note usually followed by two shorter secondary notes. Analysis of nine recordings made at Santa Cecilia revealed that the note repetition rate of primaiy notes is 15.3-73.2 (X=33.9) notes per minute. The duration of the primary notes is 0.06-0.15 (X=0.09). The pulse rate is 100-120 (X=105) pulses per second. There usually are three harmonics, of which the funda- FiG. 81.— Hijla boons, 9 , 90 mm SVL (KU 109424). mental at 2625-3339 (X=2984) Hz is emphasized. Food. — Twenty-five adults had eaten a variety of small arthropods. Dipterans (36% of stomachs) were the most com- mon prey, followed by spiders (20%), lepidopterans (12%), and beetles (8%). The largest item of prey was a moth 16 mm long eaten by a frog having a snout- vent length of 25 mm. Hyla beans (Linnaeus) ( Figure 81 ) Rana hoans Linnaeus, Syst. Nat. 10, Stockholm, p. 2L3 [Type locality. — "America"]. Ht/la hoans — Duellman, 1971, Herpetologica, " 27:398. A/afer/a/.— Santa Cecilia, 123+1. Identification. — This large brown tree frog has small triangular calcars, exten- sive webbing on the hands and feet, and pigmented reticulations on the lower eyelid. FurtheiTnore, males have a pro- jecting propollex. Uijla geographica is the only other brown tree frog in the area having calcars and reticulations on the eyelid; it diff^ers from the larger H. hoans in coloration and in the amount of webbing. Webbing extends to about the middle of the outer finger in H. geo- graphica but to the base of the disc in H. hoans. In H. geographica the flanks and anterior and posterior surfaces of the thighs are marked by distinct, narrow, dark vertical bars, whereas in H. hoans vertical bars are present, but they are diffuse on the flanks and barely distin- 132 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY guishable on the thiglis. All other brown tree frogs, with which //. hoans might be confused, lack reticulations on the lower eyelid. The head is broad; a tympanum is present. The toes and all fingers, except the first, are webbed to the bases of the discs. In males there is a curved prepol- lical spine. A small triangular calcar is present on the heel. The skin on the dor- sum is smooth; that on the belly is granu- lar. The dorsal color varies from tan to reddish-brown, grayish brown, or dark brown, usually with slightly darker mot- tling on the back and transverse bars on the limbs. By day, the dorsal color usually is paler than at night. The flanks are grayish tan with dark brown vertical marks, and the posterior surfaces of the thighs are grayish brown with faint brown vertical marks. The webbing is brown, and the venter is creamy white. The iris is bronze, and the lower evelid is reticulated with silvery gold, i 118 mm; 9 110 mm. Occurrence. — Although males have been heard calling from trees in primary and secondary forest, all individuals have been found along the Rio Aguarico, where they were sitting on the river bank or in bushes or on large cane-like grasses. These large frogs are most active in the drier times of the year. Their calls sel- dom were heard during rainy periods. Throughout March 1967 one male (pre- sumablv the same individual) began call- ing at dusk (1S00-1S30 hrs) daily. Life History. — Normally, the males of this large species construct shallow, ba- sin-like nests (up to 40 cm in diameter) in muddy river banks; water seeps into the basins, from which the males call and in which females deposit eggs on the surface of the water (Duellman, 1973a). Despite some exposed mud flats on the north side of the large island in the Rio Aguarico, no nests were found there. However, males were found calling from, and tadpoles living in, natural depres- sions in the pebble bank of the Rio Aguarico. Many small basins remain filled with water when the level of the river drops. If the presence of calling males along the banks of the river is a true reflection of breeding activity, then breeding in Iltjla hoans is sporadic and associated with drier times of the year - normally August-September and Decem- ber-February. Gravid females were found in February (1), March (1), July (IS), August (lb), October (1), and December ( 1 ) . Twenty-three of these frogs had snout- vents lengths of SO- 110 (X:=99.0) mm and contained 1300-4800 (X=r3l54) mature ovarian eggs. The pigmented eggs had maximum diameters of 2.1 mm. Tadpoles. — The following description is based on tadpoles (KU 10942S) in stage 37 having body lengths of 15 mm and total lengths of 43 mm. The body is broadly ovoid, widest and deepest pos- teriorly, and ovoid in lateral view. The snout is round. The eyes are large, mod- erately separated, and directed dorso- laterally. The nostrils are about three- fifths of the distance from the tip of the snout to the orbits. The opening of the sinistral spiracle is directly posterodor- sally on the midline at about midlength of the body. The mouth is ventral. The median part of the upper lip is bare, and the lips are folded laterally; a single row of small labial papillae border the mouth laterally and ventrally. The beaks are moderately robust and finely serrate. There are two upper and four lower rows of denticles, of which the second upper row is interrupted medially and the fourth lower row is much shorter than the others. The tail is about 65 percent of the total length and is acutely round- ed terminally. The dorsal fin does not extend onto the body, and at its deepest point at alwut midlength of the tail the depth of the dorsal fin equals the depth of the caudal musculature and twice the depth of the caudal fin. The dorsum is reddish tan with dark olive-brown mark- ings; the belly is white with a golden hue. The tail is cream with reddish brown reticulations. The iris is bronze. EQUATORIAL HERPETOFAUNA 133 Tadpoles were found in a rocky pool in the riverbed, where they sought ref- uge under and between stones. The pool \\'as about 1 m in diameter, and the water was no more than 15 cm deep. Mating Call — The call consists of a series of short, loud, low-pitched notes, usually produced in groups of three to ten. Analysis of the calls of three indi- \ iduals from Santa Cecilia revealed that notes were repeated at rates of 21.4-66.7 (X— 38.2) notes per minute. The dura- tion of the notes is 0.18-0.27 (X=0.23), and pulses are produced at a rate of 100-140 (X=117) per second. There are at least two hamionics; the fundamental frequency at 435-565 (X=512) Hz is dominant in three of the notes, whereas tlie second harmonic is dominant in six others. Food. — Stomachs of 13 adults re- vealed the presence of orthopterans ( 85% of stomachs), lepidopteran larvae (15%), and moths (8^). One frog having a snout-vent length of 76 mm had eaten an orthopteran 25 mm long. Remarks. — This is the only hylid frog in the area in which males attain a larger size than females. In Panama //. ])oans is even larger - males to 132 mm and females to 117.5 mm (Duellman, 1970b: 258). Breder (1946:409) reported nest building by males of H. roscnhcrgi, and Lutz (1960:61) reported the habit in H. fahcr and also observed territoriality and aggressive behavior in H. fahcr. All three species are closely related ( Duell- man, 1970b: 246). Thus, it can be as- sumed that territoriality is common to all three nest-building species and that perhaps selection favors larger size in males. Larger individuals would tend to be more successful in combat than small- er individuals. Hyla bokermanni Coin ( Figure 82 ) HyJa bokermanni Goin, 1960, Ann. Mag. Nat. Hist. (13)2:721 [Type locality.— Tarauaca, Tenitorio do Acre, Brasil]. Duellman and Fig. 82. — Hi/la hokcrmani. Upper: $, 23 mm SVL (KU 123089). Lower: ?, 25 mm SVL (KU 126552). Crump, 1974, Occas. Papers Mus. Nat. Hist. Univ. Kansas 23:14. Afafe7-/a/.— Santa Cecilia, 99+12. Identification. — This small tan frog ^^ith a short, truncate snout has dark thighs with yellow spots. Two other small frogs in the area have a pattern of pale spots on dark thighs. Hyla hrevi- frons is like H. bokermanni in having one to three small yellow spots on the dorsal surface of the thigh; however, in H. hrcv- ifrons the anterior surface of the thigh is uniform dark brown, whereas in most specimens of U. ])okermanni there is a large black-bordered yellow spot on the anterior surface of the thigh. Hyla parvi- ceps has dark brown thighs with two small cream spots on the anterior sur- faces. Furthermore, H. parviccps has a bright orange spot on the proximal ven- tral surface of the shank (unmarked in H. hokerm^aivii) and. one vertical cream bar on the upper lip (two bars in H. 134 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY hokermanni) . Other small yellowish tan or brown frogs in the area do not have one to three yellow spots on dark brown thighs. The snout is short and truncate in dorsal view and profile. The tympanic ring is visible ventrally. An axillary mem- brane extends about one-half the length of the upper arm. The fingers are about one-third, and the toes about one-half webbed. Breeding males lack nuptial excrescences. The skin on the dorsum is smooth, and that on the belly is granu- lar. At night the dorsum is pale yellow- ish tan; dorsal markings and dorsolateral stripes are barely discernable. By day the dorsum is tan usually with dark markings consisting of many small flecks, three transverse bars, or a large blotch anteriorly and a transverse bar posterior- ly. The shanks and forearms are tan with distinct brown crossbars. The thighs are dark brown with two creamy yellow spots on the dorsal surfaces; in 80 per- cent of the specimens there is a large, oval bright yellow spot, usually with a narrow black border on the anterior sur- face of the thigh. There are two pale creamy yellow vertical or diagonal bars below the eye. In about half of the specimens a narrow creamy yellow, verti- cal rostral stripe is present; usually in these specimens the rostral stripe is con- fluent with a narrow canthal stripe. In males a narrow creamy yellow dorsolat- eral stripe extends from the outer edge of the eyelid to the mid-upper flank; a short stripe is present in the groin. In females a broad, creamy white stripe and inguinal spot are present. The venter is creamy white, except for a yellow vocal sac in males. The iris is silvery gray with a narrow red ring around the pupil. c5 24 mm; $ 28 mm. Occurrence. — About 75 percent of the HijJa hokermanni were found in secon- dary forest; the others were in primary forest. All were found at night on vege- tation; only four of 101 individuals were more than 1.5 m above the ground. Males usually call from emergent vegetation in swamps in, or, less frequently, at the edge of, forest. Most calling males are 1-1.5 m above the water. The frogs were found throughout the year, except De- cember. Life History. — Gravid females were found in January (1), February (17), March (29), April (29), May (9), June (2), October (1), and November (5). Hijla hokermanni seems to be an oppor- tunistic breeder, reaching peaks of re- productive activity at times of heavy rainfall. Duellman and Crump (1974: 23) noted that a He//con/a-choked swamp in secondary forest there were peaks of breeding activity in October- November 1971 and in February-April 1972. These were times of greatest water depth in the swamp. During breeding peaks 300-500 males and 10-50 females were estimated to be in the swamp on any given night. Twenty-nine females having snout-vent lengths of 22.5-25.5 (X=:23.9) mm contained 65-135 (X=: 98.3) mature ovarian eggs. HijJa hoker- manni deposited eggs on vegetation over the water. The small, lightly pigmented eggs are 1.5-1.8 (X=1.6, N=53) mm in diameter. Clutches contained 60-196 (X= 116.8, N=53) eggs. The eggs re- quire 4.0-5.3 (X=4.4, ]V=20) days to hatch, at which time the hatchlings drop into the water. Hatchling tadpoles have total lengths of 4.0-4.5, (^=4.2, N=20) mm. Two recently metamorphosed young have snout- vent lengths of 11.3 and 11.8 mm. Tadpoles. — The following description is based on tadpoles (KU 124193) in stage 35 having a body length of 6.8 mm and a total length of 18.8 mm (see Duell- man and Crump, 1974:13 for illustra- tion). The body is ovoid, widest me- dially and deepest posteriorly, with a round snout. The eyes are large, widely separated, and directed laterally. The nostrils are about midway between the tip of the snout and the orbits. The opening of the sinistral spiracle is just below the midline at about three-fifths of the length of the body. The mouth is EQUATORIAL HERPETOFAUNA 135 anterior and lacks lateral folds. The up- per lip is bare; one row of labial papillae border the mouth laterally and ventrally. The beaks are moderately robust and finely serrated. Denticles are absent. The tail is about 64 percent of the total length and terminates in a short filament. The tail is deepest at midlength, where the caudal musculature and dorsal fin are of equal depth, but the ventral fin is about 60 percent as deep as the dorsal fin, which does not extend onto the body. The body is dark brown with two short, broad, longitudinal yellowish tan bars on the snout. Proximally, the tail is pale orange; distally, the tail is unpigmented, except for broad gray vertical marks. The iris is silver with a red tint medially. The tadpoles are pelagic in swamps; they seek cover amidst vegetation or by diving towards the bottom. Mating Coll. — The call consists of a high-pitched primary note followed by two to five shorter secondary notes. Anal- ysis of recordings of four individuals at Santa Cecilia revealed a note repetition rate (primaries) of 5-19 (X=12.2) notes per minute having durations of 0.23-0.28 (X=0.259) sec and 100-190 (X=rz 132.5) pulses per second. The dominant fre- quency is at about 4500 Hz. Food. — Sixteen stomachs contained a variety of small arthropods. Ants and orthopterans were each represented in 50 percent of the stomachs; dipterans and lepidopteran larvae were each pres- ent in 19 percent of the stomachs. Single prey items were a beetle, beetle larva, homopteran, and spider. These small frogs are capable of taking large prey relative to their body size. One frog hav- ing a snout-vent length of 22 mm con- tained an orthopteran 15 mm in length, and three individuals 18, 23, and 25 mm in snout-vent length contained caterpil- lars 17, 23, and 15 mm in length. Remarks. — As noted by Duellman and Cmmp (1974:22), the three species in the HyJa parviceps group {H. hoker- manni, hrevifrons, and porviceps) at Santa Cecilia are segregated ecologically. Hijla parviceps is a habitat generalist and breeds sympatrically and synchronously with the other two species, which only rarely were found together. Further- more, H. parviceps deposits eggs in wa- ter, whereas the others deposit eggs on vegetation over water. Hyla brevifrons Duellman and Crump ( Figure 83 ) Hijla brevifrons Duellman and Crump, 1974, Occas. Papers Mus. Nat. Hist. Univ. Kansas 23:15 [Type locality. — Santa Cecilia, Pro- vincia Napo, Ecuador]. Material— Lugo Agrio, 3; Puerto Libre, 3; Santa Cecilia, 153+5. Identification. — This small tan frog with a short, truncate snout has one to three small yellow spots on the otherwise dark brown thighs. Two other frogs in the area have pale spots on dark thighs. Hyla hokermanni also has yellow spots on the dorsal surfaces, but most speci- mens also have a large black-bordered yellow spot on the anterior surface of the thigh. Hyla parviceps has two cream spots on the anterior surface of the thigh, a bright orange spot on the proximal ventral surface of each shank ( unmarked in H. hrevifrons), and one vertical cream bar on the upper lip (two in H. hrevi- frons). Other small yellowish tan or brown frogs in the area lack pale spots on the thighs. The snout is short and truncate in dorsal view and profile. A tympanic ring is visible ventrally. An axillary mem- brane extends one-half the length of the upper ami. The fingers are about one- half, and the feet about two-thirds, webbed. Breeding males do not have nuptial excrescences. The skin is smooth on the dorsum and granular on the belly. At night the dorsum is pale tan with faintly darker dorsal markings. By day the dorsum is olive-tan with three or four broad transverse brown marks on the back and transverse bars on the fore- arms and shanks. The thighs are dark brown with 1-3 creamy yellow spots on the dorsal surfaces. Two creamy yellow 136 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 83. — Hyla brevifrons. Upper: S, 19 mm SVL (KU 126370). Lower: ?, 23 mm SVL (KU 126371). vertical or diagonal suborbital bars are invariably present; a narrow vertical ros- tral stripe is present in about half of the specimens, and a narrow cream canthal line is present in about two-thirds of the specimens. In most males and in all females a creamy yellow dorsolateral stripe extends from the outer edge of the eyelid to the midHank region and then ventrally on the flank. The stripe is a very narrow line in males and a distinct broad stripe in females. The chin and chest are white in females, and the vocal sac is yellow in males; other ventral sur- faces are unpigmented. The iris is silvery gray with a narrow red ring around the pupil. 6 22 mm; 9 25 mm. Occurrence. — All individuals were in secondary forest. Thirteen were perched on low ( < 1.5 m) vegetation in the for- est, and 121 were on vegetation in and around swamps at the forest edge or, less frequently, in forest. All were found at night. Individuals were found through- out the year, except in February. Life History. — Gravid females were found in January (1), March (4), April (1), June (1), July (3), October (2), November (1), and December (2). Hyla brevifrons seems to be an opportunistic breeder with peaks of breeding activity correlated with heavy rains. Fifteen fe- males having snout-vent lengths of 18.0- 23.5 (X=2d.9)mm containe^d 48-114(X =79.1) mature ovarian eggs. The small (1.2-1.6, X=1.3 mm; N=e) lightly pig- mented eggs are deposited on vegetation over water. One clutch required 6.3 days to hatch, at which time the tadpoles dropped into the water. A hatchling tad- pole had a total length of 6.0 mm. Two recently metamorphosed young had snout- vent lengths of 7.6 and 8.4 mm. Tadpoles. — The following description is based on a tadpole (KU 125899) in stage 37 having a body length of 7.2 mm and a total length of 23.0 mm ( see Duell- man and Crump, 1974:13, for illustra- tion). The body is ovoid, widest ante- riorly and deepest posteriorly, with a rounded snout. The eyes are large, wide- ly separated, and directed dorsolaterally. The nostrils are about midway between the tip of the snout and the eyes. The opening of the sinistral spiracle is direct- ed posterodorsally just below the midline at about three-fifths of the length of the body. The mouth is anterior and lacks lateral folds; the upper lip is bare, but a row of small labial papillae arc present laterally and ventrally. The beaks are moderately robust and finely serrated; denticles are absent. The tail is about 68 percent of the total length and termi- nates in a short filament. The tail is deepest at about two-thirds of its length, where the ventral fin is nearly as deep as the dorsal fin, and the caudal muscula- ture is about two-thirds of the depth of the dorsal fin, which does not extend on- to the body. The body is dark brown with an indistinct, diffuse, tan dorsolat- eral stripe and cream flecks laterally. The tail is tan with dark brown mottling and EQUATORIAL HERPETOFAUNA 137 a briglit red streak on the dorsal fin. The iris is grayish bronze with a red ring around the pupil. Tadpoles were found in deep water ( ± 1 m ) in forest ponds, where they took refuge amidst detritus on the bottom. Mating Call. — The call consists of a series of short insect-like chirps repeated at a rate of 26-46 (X=36.8, N=9) notes per minute. Further analysis of record- igns of nine individuals at Santa Cecilia reveals that notes have a duration of 0.43-0.49 (X=0.467) sec and 30-40 (X =38.9 ) pulses per second. The dominant frequency is at about 4600 Hz. Food. — In 25 stomachs, small beetles (44% of stomachs) were the most com- mon prey, followed by lepidopteran lar- vae (20%), homopterans and spiders (16% each), orthopterans (S%), and one fly and one ant (4% each). The largest prey item was a caterpillar 17 mm long in a frog 18 mm in length. Hyla calcarata Troschel ( Figure 84 ) Htjia calcarata Troschel, 1848 Amphihien, in M. R. Schomburgk, Reisen Biitisch-Giiiana, 3:60 [Tvpe locality. — Guyana]. Duellman, 1973, Copeia, 1973(3):552. Material. — Puerto Ore, 12; Santa Ce- cilia, 96 -f 5. Identification. — This brown tree frog has large elongately triangular calcars, pale flanks and hidden surfaces of the thighs with bold black markings, no re- ticulations on the lower eyelids, and no weighing between the fingers. The ab- sence of reticulations on the lower eyelid and webbing between the fingers distin- guish H. calcarata from H. hoans and H. geop,raphica; the latter has the fingers half webbed, and H. hoans has nearly fully webbed fingers. Both species have golden reticulations on the lower eyelid. Hijla calcarata is most easily confused with H. fasciata. In the latter the calcars are smaller and proportionately broader, the black markings on the flanks and Fig. 84.— Hyla calcarata, $ , 37 mm SVL ( KU 111677). thighs usually are irregular spots, and brown flecks are present on the throat and chest. Hyla calcarata has large, elon- gate calcars, the black markings on the flanks and thighs usually in the fomi of vertical bars, and an immaculate venter. Two other hylids in the area have calcars and vertical dark marks on the thighs; Hyla garhei has broad black and orange bars on the thighs, small conical calcars, a pointed tuberculate snout, and a mot- tled green, tan, and black dorsum. Phyl- lomedusa tomopterna has bright orange thighs with brown bars, large triangular calcars, no webbing between the toes, and a uniformly green dorsum. The head is broad; a tympanum is present. Webbing is absent or vestigial between the fingers; the toes are one-half to two-thirds webbed. Breeding males have horny nuptial excrescenses on the thumbs. Large, elongate, triangular cal- cars are present on the heels. The skin on the dorsum is smooth; that on the venter is granular. At night the dorsum is pale yellowish tan with faint darker markings; by day the dorsum is pale red- dish brown with brown markings. The dorsal markings consist of: 1) five to seven broad transverse marks, intercon- nected or not; 2 ) narrow transverse lines in the interspaces between the broad marks in some individuals; 3) narrow dark brown middorsal line from the tip 138 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY of the snout to a point between the occi- put and rump; 4) transverse bars on the hmbs (one or two each on upper ami and forearm and three to five each on thigh, shank, and foot). The webbing on the foot is brown. The posterior half of the flanks and the anterior and pos- terior surfaces of the thighs are bluish white in small adults and blue in large adults, usually with vertical black bars about equal in width to the interspaces; in a few individuals the dark markings are in the form of irregular spots. As many as eight vertical bars may be pres- ent on the flank or thigh. Narrow creamy white stripes are present above the anus and on the outer edge of the foot. The venter is white; in large females there is a tan suffusion along the margin of the chin. The coloration of juveniles is no- ticeably different (see Remarks). The iris is creamy silver. S 41 mm; 9 61 mm. Occurrence. — Hyhi calcarata inhabits forests. All individuals were found on vegetation; most were on branches of bushes less than 1.5 m above the ground. Of 92 individuals with adequate ecologi- cal data, 23 were in a forest swamp, 32 were at the edge of a lake, and 5 were along streams in the forest; the others were scattered in the forest away from water. The frogs were observed every month of the year, except December. Life History. — Although males were heard calling throughout most of the year, only ten gravid females were found - March (3), April (2), August (1), September (1), October (2), November ( 1 ) . Breeding seems to be associated sporadically with rains throughout the year. The ten females had snout-vent lengths of 47-61 (X=53.9) mm and con- tained 835-1600 (X=1143) mature ovar- ian eggs. Three females having snout- vent lengths of 52.5-57.0 (X=55.0) mm deposited clutches of 1060-1250 {X= 1153) eggs in the laboratory. The pig- mented eggs (2.0 mm in diameter) were deposited as a film on the surface of the water. The eggs hatched in 79-86 hrs; hatchling tadpoles had total lengths of 5.5-7.0 (X=6.0) mm. Tadpoles.— Tadpo\es (KU 152292) in developmental stage 33 have body lengths of 9.5 and total lengths of 32.5 mm. The body is ovoid, widest and deep- est at midlength, with an acutely round- ed snout. The eyes are large, closely set, and directed dorsolaterally. The nostrils are about three-fifths of the distance from the tip of the snout to the orbits. The spiracular tube is long, and the opening of the spiracle is directed pos- teriorly on the midline at about two- thirds the length of the body. The mouth is anteroventral and has lateral folds. The median part of the upper lip is bare; elsewhere the mouth is bordered by one row of small labial papillae. The beaks are slender with moderate-sized serra- tions. There are two upper (second in- terrupted medially) and three lower rows of denticles. The tail is about 72 percent of the total length, and it temii- nates in a pointed tip. The dorsal fin does not extend onto the body and is deepest at the midlength of the tail, where it is equal in depth to the caudal musculature and half again as deep as the ventral fin. The dorsum of the body is dark brown with faint tan mottling and tan interorbital bar. The tail is yellowish tan with vertical brown bars. Tadpoles were found in swamps and small sluggish streams in forest. Matinp, Call. — The call consists of one or more soft, low-pitched, poorly modulated, monophasic notes. Analysis of one recording from Santa Cecilia re- vealed a repetition rate of 8 notes per minute. The notes were 0.07 sec in dura- tion and were produced with a pulse rate of ISO pulses per second. The dominant frequency was at about 1400 Hz. Food. — Twenty-five individuals con- tained the remains of a diversity of small arthropods and one small frog. Orthop- terans were the most common prey (40% of the stomachs), followed l:)y spiders (24%), beetles (8%), homopterans (8%), EQUATORIAL HERPETOFAUNA 139 lepidopteran larvae (8%), and five other groups eacli in fewer than 5 percent of the stomachs. Remarks. — Juveniles having snout- vent lengths of less than 25 mm lack transverse markings on the dorsum and vertical black bars on the flanks and thighs. A faintly darker brown middor- sal stripe has five or six lateral expan- sions, which may be incipient transverse marks. Hyla cruentomma Duellman (Figure 85) Ilyla cruentomma Duellman, 1972, Copeia 1972 (2):266 [Type locality. — Santa Cecilia, Provincia Napo, Ecuador]. Material. — Lago Agrio, 2; Santa Ce- cilia, 118+13. Identification. — This is one of three species of moderate-sized tree frogs in the area having a yellowish tan to green- ish tan dorsum with darker longitudinal markings. Among these species, //. crti- entonima is distinctive in having a silvery bronze iris with a median horizontal red streak. UijJa cruentomma is like H. ru- bra in having smooth skin on the dorsum, but //. rubra differs by having bold black or dark lirown reticulations enclosing yellow spots in the groin and on the pos- terior surfaces of the thighs. These sur- faces are uniform pale green in H. cruen- tomma and yellow or pale green with brown spots or streaks in H. funerea, a species having small tubercles on the dorsum and a greenish bronze iris with black flecks. The snout is round in dorsal view and in profile; an external tympanum is pres- ent. Webbing is absent between the fingers; the toes are about one-half webbed, but the webbing is vestigial be- tween the first and second toes. Nuptial excrescences are absent in breeding males. The skin on the dorsum is smooth and that on the belly is granular. The dorsum is yellow, tan, or pale green usu- ally with dark brown markings, most Fig. 85. — Htjia cruentomma, S , 27 mm SVL (KU 111924). often consisting of a canthal stripe, inter- orbital bar, transverse bars on the fore- arms and shanks, and irregular longi- tudinal marks on the body. The groin and anterior and posterior surfaces of the thighs are uniform pale green or yellow- ish green. The ventral surfaces are white in females and pale yellow in males, which have bright yellow vocal sacs. The iris is silvery bronze with a median hori- zontal red streak. The bones are green. c^ 28 mm; 9 32 mm. Occurrence. — Most individuals were found in low (< 1.5 m) vegetation in and around temporary ponds; about equal numbers were found in ponds in clearings and in a f/ total length and is pointed terminally. The greatest deptli of the tail is at two-fifths of its IcMigth, where the deptli of the caudal fin is slightly less than, and the depth of the caudal musculature is one- half, the depth of the dorsal fin, which extends onto the body. The body is pale green dorsally and laterally and silver ventrally. The tail is pale orange with brown flecks, and the iris is silvery bronze. Larger tadpoles (stage 36) are more yellowish green on the body and tail with brown flecks on the tail, and the iris is bronze. Tadpoles were found in a pond in primary forest by day and in a pond at the edge of the forest at night. Mating Call — The call consists of a moderately long, rather high-pitched, single note. No recordings were ob- tained. Food. — Orthopterans, which were found in 64 percent of 25 stomachs con- taining food, were the most common prey items, followed by lepidopteran larvae (16%), beetles '(16%), spiders (12%), and homopterans, dermapterans, dipterans, and dipteran larvae (4% each). Caterpillars 35 and 27 mm long had been eaten by frogs having snout-vent lengths of 41 and 3S mm. The latter also contained a roach 14 mm long, and another individual having a snont-vent length of 31 mm had eaten an orthop- teran 17 mm long. Remarks. — See the account of Hyla cruentomma for a discussion of the eco- logical distribution of H. funerea with respect to other members of the Hijla ruhra group. The variation in, and dis- tribution of, Hijla funerea were discussed by Duellman "(1971). Hyla garbei (Miranda-Ribeiro) ( Figure 88 ) Gmheana ^arhci Miranda-Riheiro, 1926, Arch. Mus. Nac. Rio dc Janeiro, 27:96 [Type lo- cality.— Eirnnepe, Rio Junia, Territorio do Ama/oiias, Brasil], Ibjla f^aihci — Duellman, 1970, Copeia, no. 3:536. Material — Puerto Libre, 2; Puerto Ore, 1; Santa Cecilia, (S2-f-16. Identification. — This distinctive small EQUATORIAL HERPETOFAUNA 145 Fig. 88.—HuIa garhei, $, 37 mm SVL (KU 111680). hylid is readily distinguished from all other frogs in the area by the following eombination of characters : 1) long, acu- minate, projecting snout; 2) long slender fingers lacking webbing and bearing truncate discs; 3) elongate tubercle on heel; 4) row of tubercles along edge of lower jaw; and 5) groin and hidden sur- faces of thighs yellow or orange with broad black vertical bars. The general appearance of Hijla p^arhci is suggestive of Eleuflierodacfyhis more than Hijla: however, the combination of characters given abo\'e distinguishes H. p^arhci from Eleiitlierodactyhis, as well as other IlijJa. The snout is long, acuiuinate, project- ing, and usually bearing at least one temiinal tubercle. A tympanum is pres- ent. The fingers are long and slender, bearing broad, truncate discs; webbing is absent between the fingers, and breed- ing males lack nuptial excrescences. The toes are about one-half webbed, except that webbing is vestigial between the first and second toes. The skin is smooth above the tubercles on the snout, eyelids, supratympanic fold, heels, and along the outer edges of the forearms and feet. The skin on the throat is smooth and that on the belly granular. The dorsum is pale dull green or reddish brown with dark brown or black markings always consist- ing of a triangle on the head with the corners on the eyelids and the apex pos- teriorly. Other dorsal markings usually consist of: 1) supratympanic stripe; 2) indistinci bars on lips, weakly defined transverse bars on the limbs; 3) larGie, ovoid marks posterodorsal to the axillae; and 4) small dashes or transverse marks posteriorly on the dorsum. The anterior and posterior surfaces of the thighs are yellow or pale orange with broad black or dark brown vertical bars, usually three on each surface. The venter is creamy white with gray flecks on the chin and, in some individuals, laterall)' on the bell)'. The iris is pale creamy bronze with a median horizontal reddish brown streak. <5 37 mm; 9 48 mm. Occurrence. — IhjJa ^^arhei is wide- spread and abundant at Santa Cecilia, but only one individual was found in pri- mary forest. Others were about equally distributed between clearings and secon- dary forest, but most of those in secon- dar\- forest were at breeding ponds. One individual was on the ground in a clear- ing; all others were on low ( < 1.5) vege- tation. All of the frogs were found at night. Individuals were found through- out the year, except in November and Jaiuiary; this probably is an artifact of sampling. Life History. — Hyhi ' having acuminate, pro- truding snouts. The one bright green Eleutlwrodactylus (E. acuminatus) lacks webbing on the hands and feet, lacks an external tympanum, and has a pointed snout and truncate discs, whereas H. gra- nosa has a tympanum and round discs. The species of CentroJcncUa all have truncate discs and gold, )'ellow, or white flecks on a green dorsum, and PlujIIo- nieclusa lack webbing on the feet. The snout is broadly rounded in dor- sal view and round in profile. An exter- nal tympanum is present. The outer finger is about one-third webbed, and the second and third fingers are basally webbed; the toes are about two-thirds webbed. Breeding males lack horny nup- tial excrescences, but adult males have projecting prepollical spines. The skin on the dorsum is finely granular; that on the belly is coarsely granular. In most in- dividuals the dorsum is uniform pale green. In some individuals minute red flecks are visible on the dorsum. In about five percent of the specimens, there are reddish brown or black dorsal markings that maximally consist of a canthal stripe, interorbital bar, two to four spots on the sacral region, or transverse bars on the shanks. The groin and hidden surfaces of the limbs are green, with a bluish tint in some individuals. The venter is nearly transparent but has a faint greenish tint. The fingers and toes are greenish yellow, and the edge of the upper eyelid is cream. The vocal sac is green, and the iris is pale cream. $ 54 mm; 9 44 mm. Occurrence. — Only two individuals were found in clearings. All others were in primary or secondary forest. All frogs were on vegetation at night. Males com- monlv were heard after rains throughout the year along the forested shores of the lake and in two forest-edge swamps. No individuals were found in November through January. Life History. — IlyJa granosa breeds sporadically at times of heavy rainfall. Gravid females were found in February (1), March (2), April (3), June (2), July (2), and August (1). Eleven fe- males having snout-vent lengths of 40.0- 44.0 (X=41.2) mm contained 350-520 (X=426) pigmented eggs. Presumably, these are deposited as a surface film on the water. The smallest, recently meta- morphosed young has a snout-vent length of 14.2 mm. Tadpoles. — The following description is based on tadpoles (KU 125902) in stage 25 having body lengths of 7.5 mm and total lengths of 22.5 mm. The body is ovoid, wider than deep, and widest anteriorly. The snout is broadly rounded in dorsal view and round in profile. The 150 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY eyes are large, narrowly separated, and directed dorsolaterally; the nostrils are midway between the tip of the snout and the orbits. The opening of the sinis- tral spiracle is directed posterodorsally below the midline at about the mid- length of the body. The mouth is antero- ventral and has lateral folds. The median part of the upper lip is bare; elsewhere one row of small lal:)ial papillae border the mouth. The beaks are slender with fine serrations. The single upper row of denticles is narrowly interrupted medi- ally; the two lower rows are complete. The tail comprises about 67 percent of the total length and terminates in a point. The fins are shallow; the tail is deepest at midlength, where the depth of the caudal musculature is equal to the depth of the dorsal fin, and where the depth of the ventral fin is about two- thirds of the depth of the dorsal fin, which does not extend onto the body. The body and caudal musculatiue are dull, pale green, and the fins are trans- parent. Tadpoles were obtained from amidst aquatic vegetation in swamps at the edge of the forest. Mating Call. — The call consists of a series of two to four notes, "boop-boop- boop." Analysis of ten recordings made at Santa Cecilia reveals that the call rate is 14-40 (X=:24.7) notes per minute and that the notes are produced at a rate of 40-80 (X=57.S) per minute. Notes are 0.03-0.18 (X= 0.078) sec in duration and have 240-400 (X=299) pulses per sec- ond. The dominant frequency is at 1217- 1587 (X=1390) Hz. Food. — Eight stomachs contained four beetles, two moths, three orthop- terans, and two spiders. Hyla lanciformis (Cope) ( FiGUHE 91 ) Hijpsihou-i lanciformis Cope, 1870, Pioc. Amer. ' Philos. Soc, 11:556 [Type locality.— Pelxis, Departamcnto Lorcto, Peru]. llijla lanciformis — Boulenfrcr, 1882, Cat. Batr. Sal. Brit. Mus., p. 361. Fig. 91. — Htjla lanciformis. Upper: $ , 80 mm SVL (KU 122778). Lower: juvenile, 19 mm SVL (KU 122739). Material. — Puerto Libre, 1; Puerto Ore, 2; Santa Cecilia, 176+19. Identification. — Adults of this large tree frog are easily distinguished from all other frogs in the area by the following combination of characters: 1) snout long and pointed; 2) creamy yellow labial and supra-anal stripes; 3) throat and chest brown with large cream spots; 4) fingers basally webbed; 5) males having pro- jecting prepollical spine. Leptodactyhis mystaceus also has a cream labial stripe, but it has a unifonn white or yellow venter and no prepollical spines. Juve- niles (< 25 mm snout-vent length) are yellowish tan with longitudinal brown lines on the dorsum. The presence of a white supra-anal stripe and a brown me- dian stripe and brown dashes on the tliroat distinguish them from other juve- nile hylids having similar dorsal patterns (Hyla alboguttata, H. calcarata, H. fas- ciata, and Osteocephalus leprieiirii). The snout is long and pointed in dor- sal view and round in profile. The promi- EQUATORIAL HERPETOFAUNA 151 nent tympanum is nearly as lar^e as the eye. The outer fingers are webbed ba- sally, and tlie inner fingers lack webliing; the toes are about two-thirds webbed. Males lack horny nuptial excrescences l)ut have a projecting prepollex. The skin on the dorsum is smooth; that on the ])elly is granular. At night the dorsum is tan; by da\' it changes to brown or olive- tan. Dorsal markings consist of six to eight transverse brown or olive-brown bars on the dorsum of the body. Usually the anteriormost bar is between the eyes. A distinct creamy yellow to silvery \\'hite labial stripe extends from the tip of the snout to the angle of the jaw. The side of the head is olive-green to dark brown. The anterior and posterior surfaces of the thighs are tan to pale olive-green. A transverse creamy \ellow to silvery white stripe is present above the anus, and similarly colored narrow stripes are pres- ent on the heels. The throat, chest, and ventral surfaces of the limbs are brown; large cream spots are present on the chest. The belly is cream, with a snfi^u- sion of gray in some individuals. The dorsal surfaces of the discs on the first three fingers are creamy white. The iris is bronze. 6 80 mm; 9 94 mm. Occurrence. — Ilijhi lanciformis is an abundant frog, acti\e throughout the year. Although the frogs are most abun- dant in clearings, where they are active on the ground, they are most readily ob- served in second growth forest, where many individuals perch at night on branches or stems of low (< 1.5 m) bushes and trees. Observations at Muiiozlandia suggest that these frogs seek diurnal refuges beneath fallen palm fronds and other debris and in clumps of grass. Life History. — This species apparent- ly breeds continuously throughout the year. Males call every night of the year; calling sites usually are branches or trunks of low trees and bushes in, or adjacent to, shallow, temporary ponds in clearings or at the edge of the forest. Gravid females were obtained in every month, except January, June, and De- cember. Twenty-three females having snout- vent lengths of 79.0-94.0 (X= 87.3) mm contained 740-2400 (X=1617) eggs having diameters of 2.0 mm. Three clutches of 2100-2400 (X=:2250) pig- mented eggs, 2.0 mm in diameter, were deposited as a surface film on the water and hatched in about 80 hrs into tadpoles 6.0 mm long. The smallest metamor- phosed young had a snout-vent length of 13.9 mni. Tadpoles. — The following description is based on tadpoles (KU 112332) in stage 35 having body lengths of 14 mm and total lengths of 45 mm. The body is ovoid, wider than deep, widest and deep- est medially. The snout is round in dorsal view and in profile. The eyes are large, widely separated, and directed dorsolaterally. The nostrils are about midway between the tip of the snout and the orbits. The opening of the sinistral spiracle is directed posteriorly just below the midline at a point about three-fourths of the length of the body. The mouth is anteroventral and has lateral folds. The upper lip is bare; elsewhere the mouth is bordered by one row of small labial pa- pillae. The beaks are slender and bear moderate-sized serrations. There are two upper and three lower rows of denticles. The second upper row is interrupted medially, and the third lower row is much shorter than the others. The tail is about 70 percent of the total length and terminates in a point. The fins are shallow. The tail is deepest at midlength, where the depth of the ventral fin equals about 60 percent, and the depth of the caudal musculature equals the depth of the dorsal fin, which does not extend on- to the body. The body is dark brown above and laterally, and gray ventrally, with green lichenous markings. The cau- dal musculature is tan, and the fins are transparent, both marked with vertical dark brown bars. The iris is dull reddish brown. Tadpoles were found in nearly every kind of temporary aquatic environment 152 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY in clearings and at the edge of the forest. Mating Call — Tlie call consists of a loud chuckle-like note. Analysis of four recordings made at Santa Cecilia reveals that notes are produced at a rate of 9.1- 18.S (X=13.0) per minute. The notes are 0.22-0.39 (X=0.275) sec in duration and have 160-220 (X=200) pulses per second. The dominant frequency is the second harmonic at 660-728 (.^=697) Hz. Food. — Examination of 20 stomachs revealed orthopterans (60^ of the stom- achs) to be the most common prey, fol- lowed by beetles (10%), homopterans (10%), and dermapterans, lepidoptcrans, dipteran larvae, pedipalpids, and spiders (5% each). For its size, Ilyhi lanciformis does not eat large prey. Except for one grasshopper 45 mm long eaten by a frog having a snout-vent length of 78 mm, the largest prey items were 17 and 19 mm. Remarks. — On several occasions, Hyh lanciformis were observed to be captured in Muiiozlandia by day by foraging do- mestic chickens, and two were captured by diurnal snakes .(Chironius). When grasped b\' a snake or chicken, the frogs emit a loud distress call. When captured, females frequently emit a distress call, but males seldom do. Hyla leucophyllata (Bereis) ( Plate 2 ) Rana Icucopht/Uata Bereis, 1783, Slier. Gesell. Naturf. Fr. Berlin, 1783:182 [Type local- ity.— Surinam]. Htjla leucophyllata — Gravenhorst, 1829, Delic. ' Mus. Zool. Vratislav., Batr., p. 31. Duell- man, 1974, Occas. Pap. Mius. Nat. Hist. Uni\'. Kansas, 27:18. Material. — Santa Cecilia, 81+7. Identification. — This distinctive spe- cies can be confused only with //. hifiirca and //. sarayacuensis, all of which ha\'c orange webbing and hidden surfaces of the thighs, although the orange is heavily suffused with brown in some H. hifurca. In all three species the top of the head anterior to the eyes is cream to tan; a pair of pale marks extends posteriorly from the eyelids. These marks have ir- regular edges and extend posterolaterally in H. sarayacuensis, whereas they have smooth edges and are parallel, extending posteriorly in H. hifurca and H. leuco- phyllata. The marks are narrow in H. hifurca, in which the dorsal surfaces of the shanks are uniformly dark brown, and a small cream or tan spot is present on the rump and on each heel. The marks are broad in H. leucophyllata and delineate a dark brown, middorsal hour- glass-shaped mark; the dorsal surfaces of the shanks are predominantly pale. Hyla favosa, which also has orange flash col- ors, has a dark brown dorsum with a network of creamy white lines. The morphologically similar Hyla triangulum has pink or red flash colors and a uni- formly pale dorsum or one or more small dark spots. The snout is broadly rounded in dor- sal view and truncate in profile. An axillary membrane extends to about the midlength of the upper arm. The fingers are about half webbed and the toes about two-thirds webbed. The skin on the dorsum is smooth, and that on the belly is granular. At night the dorsum is yellowish tan with brown markings consisting of a large hour-glass-shaped middorsal mark extending from the eye- lids to the sacrum and a border around the head and body. Thus, the tip of the snout, sides of the head, and flanks are brown. The supra-anal region, dorsal surfaces of the thighs and arms, and outer sinfaces of the shanks are brown. In some individuals the posterolateral corners of the middorsal mark are con- nected with the dark flanks. In other specimens one or two narrow brown transverse bars are present on the dorsal surfaces of the shanks, which otherwise are yellowish tan. By day, the dorsum is creamy tan to white with dark brown markings. The venter is cream. At night the hands, feet, axillae, anterior and pos- terior surfaces of the thighs, and ventral surfaces of the limbs are yellowish EQUATORIAL HERPETOFAUNA 153 orange; by day these surfaces are orange. The vocal sac is yellow; males lack nup- tial excrescences. The iris is dull bronze. S 36 mm; 9 44 mm. Occurrence. — Only three individuals were found on vegetation at night in secondary forest; none was found in pri- mary forest. Most individuals were found on bushes and emergent grasses in a permanent forest-edge swamp, and others were in a Heliconio swamp and a temporary forest-edge swamp. Life Histonj. — Hi/la Jeticophyllata breeds sporadically after heavy rains, at which time males call from bushes around, and herbs or grass emerging from, forest-edge ponds. With the ex- ception of September, gravid females have been found in April through Oc- tober. Twelve females having snout-vent lengths of 40.0-44.0 (X=42.1) mm con- tained 220-928 (X=572.3) eggs. Three clutches contained 570-769 (X=669.7) eggs, 1.5 mm in diameter. The lightly pigmented eggs are deposited on the upper surfaces of leaves of emergent vegetation, usually less than 50 cm above the water. Tadpoles. — The following description is based on tadpoles (KU 112337) in stage 36 having body lengths of about 8.0 mm and total lengths of 28.0 mm. In dorsal view, the body is violin-shaped with a round snout; in lateral view, shal- lowly ovoid, deepest posteriorly, with a round snout. The eyes are large and lat- eral in position. The nostrils are about midway between the snout and the or- bits. The opening of the sinistral spiracle is directed posterodorsally at a point be- low the midline at about three-fifths of the length of the body. The mouth is directed anteriorly and lacks lateral folds; one row of large labial papillae is present laterally. The beaks are moder- ately ovoid and finely serrate. Denticles are absent. The tail is about 70 percent of the total length. The fins are moder- ately deep; the ventral and dorsal fins are equal in depth, and the latter does not extend onto the body. The dorsal fin is deepest at about three-fifths of the length of the tail. The tail terminates in a long filament. The dorsum of the body is chocolate brown, and the flanks are black. A white lateral stripe below the eye extends onto the proximal ventro- lateral caudal musculature. The belly is black with cream longitudinal dashes. The tail is black with fins clear proxi- mally and black distally. Tadpoles were found in deep forest pools and in forest-edge pools, where they apparently swim about amidst sparse vegetation. Mating Call — The call consists of a harsh primary note, usually followed by 2-7 (X=3.4) shorter, secondary notes. Analysis of nine recordings made at Santa Cecilia indicates that the repeti- tion rate of primary notes is 17.1-120.0 (X=77.9) notes per minute. The dura- tion of primary notes is 0.11-0.35 (0.19) sec and of secondary notes, 0.03-0.06 (X=0.045) sec. The pulse rate is 100- 180 (X=zll8.3) pulses per second. There are three harmonics, of which the funda- mental at 1913-2522 (X=2304) Hz is emphasized. Food. — Only three individuals con- tained identifiable food items. One frog contained a beetle ( Coleoptera ) , another a roach (Blattidae), and another two moths ( Lcpidoptera ) . Remarks. — Breeding congregations of H. leucopht/Uata and H. sarayocuensis arc mutually exclusive in forest ponds, although occasionally, calling males of one species are present at ponds where the other species is calling. Hyla leuco- phyllafa has not been found at ponds in primary forest, where H. sarayacuensis is abundant. Hyla marmorata (Laurenti) ( Plate 2 ) Bufo marmorata Laurenti, 1768, Synop. Rept., p. 29 [Type locality. — Surinam]. Hyla marmorata — Daudin, 1802, Hist. Nat. Rain. Gren. Crap., p. 34. Bokermann, 1964, Senck. Biol., 45:250. 154 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Material — Lago Agrio, 26; Santa Ce- cilia, 54+7. Identification. — This distinctive tree frog diflFers from all other frogs in the area by having a white or yellow belly with black spots or mottling and scal- loped dermal fringes on the outer edges of the hands and feet. The snout is round in dorsal view and in profile. A tympanum is present. The outer finger is webbed to the base of the disc; the other fingers are about two- thirds webbed. The toes arc webbed to the bases of the discs. Breeding males lack horny nuptial excrescences. Scal- loped dermal folds are present on the outer edges of the feet, foreanns, and hands. The skin on the dorsum is weakly tuberculate and that on the venter is granular. The dorsum varies from gray- ish tan to greenish tan with black, dark brown, or reddish brown mottling that usually encloses olive-green areas dorso- laterally. In most indi\'iduals, there is a pair of large brown or reddish brown scapular marks. The axilla, groin, and posterior surfaces of the thighs are orange-yellow with black spots or mot- tling on the thighs. The anterior surfaces of the thighs are pale greenish yellow with black spots. The chin and belly are white or pale yellow with black spots; the ventral surfaces of the limbs are dark gray to black. The webbing is orange distally and black proximally. The iris is pale gray with fine black reticulations and a median horizontal reddish brown streak, i 43 mm; 9 55 mm. Occurrence. — About two-thirds of the individuals from Santa Cecilia were around temporary ponds and water-filled ditches in clearings near forest. Six were on limbs of trees in secondary forest, and two were in primary forest. At Lago Agrio, the frogs jumped out of the branches of trees felled in primary forest. Many of the frogs had been at heights of more than 20 m. Hyla marmorata is an arboreal forest inhabitant, but individ- uals migrate to breeding sites in clear- ings. The frogs were found in February through Jul)' and October through De- cember. Life History. — HyJa marmorata breeds sporadically after heavy rains. After some rains, numerous adults con- gregate at breeding sites, but after other equally heavy rains no congregations oc- cur. Males call from the ground, grasses, herbs, or low trees adjacent to temporary pools. Gravid females were found in February (2), April (1), May (6), July (1), October (4), and December (1). Ten females having snout-vent lengths of 46.0-53.0 (X=49.3) mm contained 630- 15(S0 (X=1146) mature ovarian eggs. Six females having snout-vent lengths of 48.0-53.0 (X=49.8) mm deposited 740- 1580 (X=1170) eggs in the laboratory. The pigmented eggs, 1.5-2.0 (X=1.7) nun in diameter, were deposited as a sur- face film on the water. They required 32-48 hrs to hatch; hatchlings had total lengths of 5.0-5.5 (X=5.1) mm. The smallest metamorphosed young had a snout- vent length of 14.5 mm. Tadpoles. — The following description is based on tadpoles (Ku" 109423) in stage 27 having bod)' lengths of 9.0 mm and total lengths of 25.0 mm. The body is ovoid, widest at midlength, deepest posteriorly, and with a round snout. The eves are large and lateral; the nostrils are about one-third of the distance from the tip of the snout to the orbits. The opening of the sinistral spiracle is di- rected posterodorsally just below the midline at about two-thirds of the length of the body. The mouth is anterior and lacks lateral folds. The mouth, except for a bare upper lip, is bordered by one row of small labial papillae. The beaks are robust and bear fine serrations; rows of denticles are absent. The tail comprises about 64 percent of the total length and terminates in a slender point. The tail is deepest at midlength, where the dorsal and ventral fins and the caudal muscula- ture are ecjual in depth. The dorsal fin does not extend onto the body. The l)ody is olive-tan with brown transverse marks. The throat is gray with brown EQUATORIAL HERPETOFAUNA 155 flecks, and the belly is white. The caudal musculature is creamy tan with brown spots. The fins are transparent with brown spots anteriorly and black spots posteriorly. The iris is silvery white. Larger tadpoles (stages 38-40) are olive green with brown blotches dorsally. The venter is white with black flecks on the chin. There is a rectangular reddish tan blotch dorsally on the body at the base of the tail, which is heavily spotted with black. Tadpoles were found in shallow grassy ponds and ditches. Mating Call. — The call consists of one to three moderately long, pulsed, low notes. Analysis of ten recordings made at Santa Cecilia reveals that notes are produced at a rate of 11.5-35.3 (X=20.2) per minute. The notes have a duration of 0.11-0.27 (X=:0.209) sec and 120-140 (X=130) pulses per second. The funda- mental frequency is at 330-423 (X=373) Hz, and the fourth or fifth harmonic at 1320-1668 (X=1514) Hz is emphasized. Food. — In 13 stomachs, beetles and orthopterans were the most abundant prey items ( each in 38% of the stomachs) . Other food items included one each - spider, mite, caterpillar, and dermap- teran. The largest prey was a caterpillar 35 mm long eaten by a frog having a snout-vent length of 38 mm. Remarks. — Recently metamorphosed young differ from adults in coloration. The dorsum is mottled dull olive-gray and black. The rump, upper arms, heels, and outer edges of the feet are creamy gray. The hands, forearms, thighs, and shanks are black with narrow grayish tan transverse bars. The posterior surfaces of the thighs are brownish black. The belly is cream; the throat, chest, and ventral surfaces of the thighs are gray, and the other ventral surfaces are black. The iris is grayish white. Hyla minuta Peters ( Plate 2 ) Htjla wiuuta Peters, 1872, Monatsb. Akad. Wiss. Berlin, 1872:680 [Type locality.— Rio de Janeiro, Brasil]. Cochran and Coin, 1970, Bull. U.S. Natl. Mns., 288:277. Material. — Santa Cecilia, 17. Identification. — This small brown frog characteristically has a dark brown in- terorbital bar and two, broad chevrons on the back. These marks and the broad transverse bars oh the shanks are bor- dered by narrow cream lines. A trans- verse cream stripe is present above the anus and on each heel. These markings serve to distinguish Hyla minuta from other small yellowish tan or brown Hyla in the area, none of which has cream anal and heel stripes. The snout is broadly rounded in dor- sal view and truncate in profile. A tym- panum is present; an axillary membrane is absent. The fingers are about half webbed and the toes are about three- fourths webbed. Breeding males do not have nuptial excrescences. The skin is smooth on the dorsum and granular on the belly. The dorsum is tan to pale reddish brown with dark brown to dark reddish brown markings (each narrowly bordered by cream), consisting of a broad interorbital mark, two broad chev- rons on the back, and transverse bars on the forelimbs and shanks. The thighs are orange-tan. A narrow cream supra-anal stripe is confluent with transverse stripes on the heels when the frog is in a resting position. The venter is creamy white, and males have a yellow vocal sac. The iris is reddish tan. i 22 mm; 9 24 mm. Occurrence. — All but two individuals were found in a restricted area of dis- turbed forest, where they were sitting on low vegetation (<1.5m) at night. They were found in May (3), July (8), and August (4). Two were on low vegetation at night at the edge of a pond in primary forest in June. Life History. — Five females - May (1), July (3), August (1) - having snout-vent lengths of 23.0-24.0 (X=23.8) mm contained 160-265 (211.6) lightly pigmented eggs 1.0 mm in diameter. No 156 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY observations of breeding activity were made at Santa Cecilia, but at Belem, Brasil I have observed males of Hijla minuta calling from emergent \egetation in temporary ponds. Kenny (1969:47) noted in Trinidad that spawning takes place throughout the rainy season and that eggs are laid in adhesive strings or clumps on submerged leaves or debris. Tadpoles. — The following description is based on one tadpole (KU 155522) in stage 41 having a body length of 14.2 mm and a total length of 41.7 mm. The body is ovoid, widest at midlength and deep- est posteriorly. The snout is broadly rounded in dorsal view and round in pro- file. The eyes are large and lateral. The nostrils are about 40 percent of the dis- tance from the tip of the snout to the orbits. The opening of the sinistral spir- acle is directed posterodorsally on the midline at about two-thirds of the length of the body. The mouth is anterior and has no lateral folds. The median part of the upper lip is bare; elsewhere two rows of small labial papillae are present. The beaks are moderately robust, broad, and finely serrate. There arc two lower, but no upper, rows of denticles. The tail is 68 percent of the total length and termi- nates in a long, slender filament. The dorsal fin extends onto the body. The tail is deepest at about midlength where the dorsal and ventral fins are equal in depth and twice as deep as the caudal musculature. The body is brown dor- sally and laterally and white vcntrally with brown ventrolateral flecks. The tail is cream with small brown blotches and fine brown reticulations. The tadpoles were obtained from a forest pond in October. Mating Call. — No calling individuals were found at Santa Cecilia, but record- ings of six individuals from Belem, Bra- sil were analyzed. The call consists of a harsh primary note, followed by 1-3 shorter secondary notes, repeated at a rate of 8-13 (X=:11.0) notes per minute and having a duration of 0.07-0.23 (X= 0.164) sec and 250-375 (X=290) pulses per second. The dominant frequency is at about 3200 Hz. Food. — Each of three individuals hax'ing snout-\ent lengths of 22 mm con- tained caterpillars of 17, 19, and 23 mm. Remarks. — Elsewhere Hyla ]ninufa usually is abundant; however, it is not only uncommon, l)ut also extremely re- stricted in its occurrence at Santa Cecilia, the only localit)' in Ecuador from which it is known. Hyla parviceps Boulenger ( Figure 92 ) Hyla parviceps Boulenger, 1882, Cat. Batr. Sal. Brit. Mus., London, p. 393 [Type loeality.— Sarayacu, Pro\incia Pastaza, Ecuador]. Duellman and Crump, 1974, Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 23:19. Material — Dureno, 2; Lago Agrio, 1; Santa Cecilia, 220+11. Identification. — This small frog with a short, blunt snout has a reddish brown dorsum with dark brown markings and a gray and white venter with a bright orange spot on the proximal ventral sur- face of the shank. The only other frogs in the area having bright orange or yel- low spots on the shanks are dendrob- atids, all of which have truncate, ex- panded digits with pairs of scutes on the dorsal surface of the distal digit; further- more, none of the dendrobatids has large dark brown marks on the dorsum. Hyla hrcvifrons and //. hokermanni are like //. parviceps in having small cream or yellow spots on dark brown thighs, but neither of those species has a spot on the ventral surface of the thigh. The snout is short and truncate in dorsal view and in profile. The tympanic ring is visible. The axillary membrane is abbreviated or absent. The fingers are about half webbed, and the toes are about three-fourths webbed. Breeding males lack nuptial excrescences. The skin on the dorsum is smooth with scattered small tubercles; the skin on the belly is granular. At night, the dorsum is tan with faintly darker markings. By day, the dorsum is orange-tan, reddish brown or EQUATORIAL HERPETOFAUNA 157 Fig. 92. — Htjia pawiceps. Upper: S , 18 mm SVL (KU 126475). Lower: 9, 25 mm SVL (KU 126476). brown with dark brown markings con- sisting of a quadrangidar or A-shaped mark anteriorly, a transverse bar in the sacral region extending onto the flanks and broad transverse bars on the fore- limbs and shanks. The flanks are paler than the dorsum. The thighs are black with one or two cream spots on the an- terior surfaces. A single creamy white suborbital bar is present. Narrow creamy white canthal stripes usually are evident. The venter is gray or white with a gray or black suffusion most prominent later- ally on the throat and belly. The ventral surfaces of the limbs are gray with a bright orange spot proximally on the shank. Some individuals also have an orange spot in the axilla. The iris is sil- very gray with a red ring around the pupil. 6 IS mm; 5 27 mm. Occurrence. — This small Hyla is ubiq- uitous throughout the year in primaiy and secondary forest, where individuals perch on leaves of herbs or leaves and branches of low (< 1.5 m) bushes at night. Males call from leaves of plants in, and peripheral to, swamps in or at the edge of forest. Life History. — Hylo parviccps breeds throughout the year. Gravid females were found in every month; of 175 gravid females, 2S were found in March, 30 in October and 49 in July. Duellman and Crump (1974:23) noted that at one HeJiconia-choked pond breeding activity reached peaks in October-November and in June-July and that males ceased call- ing: when the water level was low in the swamp. One hundred and one gravid females had snout-vent lengths of 20.0- 25.5 (X=23.3) mm and contained 120- .395 (X=234.1) matiu-e ovarian eggs. The eggs are heavily pigmented and small - 1.0-1.2 (Xml.l, N=15) mm in diameter. The eggs are deposited as clumps in the water; fifteen clutches con- tained 165-385 (X=302.8_) eggs. Hatch- ing occurs in 1.3-2.3 (X=1.9, N=13) days, and hatchling tadpoles are 4.0-4.5 (X=r4.1, N=13) mm in total length. Tadpoles. — The following description is based on tadpoles (KU 146790) in stage 27 having body lengths of 6.2 mm and total lengths of 18.9 mm. The body is ovoid, deepest posteriorly, with a bluntly rounded snout. The eyes are large and lateral; the nostrils are about 60 percent of the distance from the snout to the eyes. The opening of the sinistral spir- acle is directed posterodorsally at a point below the midline at about three-fifths of the length of the body. The mouth is anterior and lacks lateral folds. The up- per lip is bare; laterally and vcntrally the mouth is bordered by one row of large labial papillae. The beaks are robust and finely serrated. Denticles are absent. The tail is about 68 percent of the total length and temiinates in a long filament. At niidlength of the tail, the dorsal and ven- tral fins are equal in depth; the dorsal fin 158 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY is about three-fourths of the depth of the caudal musculature and does not extend onto the body. The body is dark brown with a broad, transverse, cream bar dor- sally on the snout and another posterior to the spiracle. The tail is pinkish orange to orange-tan with dark brown mottling. Tadpoles were found in shallow vege- tation-choked temporary and semipenna- nent ponds. Mating Coll — On the basis of analy- sis of one recording from Santa Cecilia, the call is a series of short, high-pitched notes produced at a rate of 54.5 notes per minute and having a duration of 0.12- 0.14 sec and 140 pulses per second. The dominant frequency is at about 6200 Hz. The call is soft and seldom audible in mixed choruses. Food. — In 25 stomachs, beetles (36% of stomachs) and lepidopteran larvae (32%) were the most common items; orthopterans, homoptcrans, diptcrans, and ants each occurred in 12 percent of the stomachs. One fly and one beetle, each 8 mm in length were contained in frogs having snout-vent lengths of 22 and 26 mm. The fonner also contained a caterpillar 28 mm long. Remarks. — Despite its abundance throughout the \'ear, no individuals were found in stomachs of the many species of snakes that feed on small frogs. Hyla punctata (Schneider) ( Figure 93 ) Calamita punctata Schneider, 1799, Hist. Amph., 1:170 [Type locality. — Surinam]. Hijla punctata — Daudin, 1802, Hist. Nat. Rain. Gren. Crap., p. 41. Duellman, 1974, Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 27:10. Material. — Santa Cecilia, 48-(-8. Identification. — This moderate-sized, primarily green Hijla is most easily con- fused with Uijla granosa, a green frog lacking dorsolateral stripes. Hyla punc- tata has red flecks on the dorsum and a red dorsolateral stripe bordered above by a yellowish white stripe. Furthermore, in H. punctata the outer finger is only Fig. 93.— Hyla punctata, 9 , 39 mm SVL ( KU 111785). basally webbed, the dorsal skin is smooth, and males lack projecting pre- pollical spines. In H. granosa the outer finger is about one-third webbed, the skin on the dorsum is finely granular, and males have projecting prepollical spines. The species of Spliaenorhynclius difl^er from H. punctata by having much more webbing on the feet and has a pointed snout and truncate discs, where- as H. punctata has a txinpanum and round discs. The species of CentrolcneUa all luue truncate discs and gold, yellow, or white Hecks on a green dorsum, and FhyUomcdusa lack webbing on the feet. The snout is round in dorsal view and in profile. A tympanum is present. The fingers are webbed basally, and the toes are about half webbed. Breeding males lack horny nuptial excrescences. The skin on the dorsum is smooth, and that on the venter is granular. At night the dorsum in some individuals is pale me- tallic red, and the flanks and limbs are pale green; others have the coloration of individuals foimd by day, in which the dorsum is pale green with dark red Hecks and a yellowish white dorsolateral stripe bordered below by a narrow dark red stripe. The throat is green, and the belly is white; other \'entral surfaces lack pigment but have a pale greenish gra\' appearance. The iris is white with fine black reticulations, ci 40 mm; 9 41 mm. Occurrence. — Ihjhi punctata was found throughout the year except in De- cember and January. Most individuals were on low (< 1.5 m) vegetation in EQUATORIAL HERPETOFAUNA 159 secondary forest, but three were in pri- mary forest. Breeding congregations were found in clearings, forest-edge swamps, and swamps in primary and secondary forest. Life History. — Breeding occurs in temporary and permanent ponds sporadi- cally after heavy rains, most frequently in April-June and October-November. Gravid females were found in April (5), May (1), June (4), July (2), October (3), and November (1). Twenty fe- males having snout-vent lengths of 35.0- 41.0 (X=:37.2) mm contained 230-430 (X=325) mature ovarian eggs. One fe- male 36.0 mm in snout-vent length de- posited 310 pigmented eggs 1.5 mm in diameter. The eggs hatched in 86 hrs; the hatchlings had total lengths of 4.0 mm. Two recently metamorphosed young had snout- vent lengths of 11.3 and 12.2 mm. Tadpoles. — The following description is based on tadpoles (KU 152298) in stage 36 having body lengths of 11.5 mm and total lengths of 37.5 mm. The body is shallowly ovoid, wider than deep and widest at midlength. The snout is broad- ly rounded in dorsal view and round in profile. The eyes are large and lateral; the nostrils are midway between the tip of the snout and the orbits. The open- ing of the sinistral spiracle is directed posterodorsally on the midline at about two-thirds of the length of the body. The mouth is anteroventral and has lateral folds. The median part of the upper lip is bare; elsewhere the lips are bordered by a single row of small papillae. The beaks are slender and bear fine serra- tions. There are two upper and three lower rows of denticles; the second up- per row is narrowly interrupted medially. The tail comprises about 70 percent of the total length and teiminates in a point. The tail is deepest at three-fifths of its length, where the caudal musculature is slightly shallower than the dorsal fin, and the ventral fin is two-thirds as deep as the dorsal fin, which does not extend onto the body. The body is dark green with black flecks, and the belly is dark gray. The tail is olive green, and the iris is pale bronze. Tadpoles were found in open water in ponds at the forest edge and in clear- ings. Mating, Call. — The call consists of a series of short, low-pitched notes. Anal- ysis of four recordings made at Santa Cecilia reveals that there are 3-17 (X== 8.4) notes per call group. The notes are 0.15-0.67 (X=0.377) sec in duration and have 60-80 (X=66) pulses per second. The fundamental frequency is at 783- 1130 (X=964) Hz. Food. — The stomachs of nine individ- uals contained four homopterans, three orthopterans, one hymenopteran, and one dipteran. Hyla rhodopepla Giinther Hijla rhodopepla Giinther, 1859, Cat. Batr. Sal. Brit. Mus., p. 112 [Type locality. — Andes of Ecnador]. Duellman, 1972, Herpeto- logica, 28:369. Material. — Santa Cecilia, 118+14. Identification. — The pale, yellowish tan dorsum with red flecks, broad red- dish brown lateral stripe, and white labial stripe immediateh' distinguish this small frog from all others in the area. Hyla riveroi is similar to H. rhodopepla but lacks the distinct lateral stripe and has brown instead of red markings. In Hyla rhodopepla the thighs are unpig- mented; other small yellowish tan or black frogs in the area have at least the dorsal surfaces of the thighs pigmented. The snout is round in dorsal view and in profile. An external tjmpanum is present; an axillary membrane is absent. The fingers are about one-third webbed, and the toes are about half webbed. Nup- tial excrescences are absent in breeding males. The skin on the dorsum is smooth; that on the belly is granular. At night the dorsum is pale yellowish tan with red flecks and a broad red band from the 160 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY snout to the rump. By day the dorsum is silvery white, and the flecks and band are reddisli brown. A broad, creamy white labial stripe extends posteriorly on the flanks, forming the ventral border to the brown band. Narrow, irregular, diagonal brown marks are present on the shanks. The venter is pale yellow, and calling males have bright yellow vocal sacs. The iris is pinkish gray with gray predominating peripherally. 6 23 mm; 5 29 mm. Occurrence. — Of 105 individuals bearing ecological data, all ^\•ere found at night, and two were on low vegeta- tion at the edge of a pond in a clearing, one was in an open marsh, four were on vegetation at the edge of the forest, seven were on vegetation in secondary forest, and 91 were on vegetation in and around swamps in secondary or primary forest. Males usually call from stems of emergent plants, and most frequently, they perch transversely on the stem within 20 cm of the water. The frogs are active throughout the year. Life Uistonj. — Although gravid fe- males were found throughout the year, Hijla rhodopepJa must be considered to be an opportunistic breeder. A few days with no appreciable rainfall will cause a cessation of breeding activit\' until the next heavy rain. Of the 115 gravid fe- males, 102 were found in five months - March (16), June (23), July (28), Oc- tober (20), and November (15). Fifty- two gravid females having snout-vent lengths of 23.0-2-8.0 (X=26.8) mm con- tained 140-422 (X=285.4) heavily pig- mented eggs about 1.1 nmi in diameter. Forty-three clutches contained 253-526 (X=3_.53.4) eggs, which hatched in 2.3- 3.3 (X=2.9) days. Hatchling tadpoles had total lengths of 5.0-5.5 (X=r5.0) mm. One female captured on 19 May 1972 mated and deposited 390 eggs in the laboratory; she was marked and released the next day. On 19 June 1972 she was recaptured; that night she deposited 260 eggs. These are the only absolute data available on frequency of mating and oviposition in any frogs at Santa Cecilia. Tadpoles. — The following description is based on tadpoles (KU 125917) in stage 30 having body lengths of 4.9 mm and total lengths of 16.2 mm. The body is ovoid, wider than long, widest at an- terior border of orbits, and deepest pos- teriorly. The snout in dorsal view is broadly rounded, in lateral view, in- clined anteroventrally. The eyes are large and lateral. The nostrils are about 40 percent of the distance from the snout to the eyes. The opening of the sinistral spiracle is directed posteriorly just below the midline at about three-fifths of the length of the body. The mouth is an- terior and devoid of lateral folds, labial papillae, and denticles. The beaks are robust and weakly serrated. The tail comprises about 70 percent of the total length and terminates in a long filament. The tail is deepest at about midlength, where the caudal musculature and ven- tral fin are equal in depth and each is ecjual to about 65 percent of the depth of the dorsal fin, which extends onto the bod\'. The body and tail are pale orange w ith scattered brown flecks. Tadpoles were found in a variety of temporary and ephemeral ponds, rang- ing from relatively open forest ponds and ponds in clearings with peripheral \egetation to weed-choked shallow ditches. Mating Call. — The call consists of a moderately high-pitched primary note followed by 3-5 shorter secondary notes. Analvsis of recordings of five individuals from Santa Cecilia reveals that primary notes are produced at a rate of 4-10 (X:= 7.9) notes per minute and have a dura- tion of 0.10-0.22 (X=0.15) sec and 200- 220 (X=206) pulses per second. The dominant frequency is at about 3925 Hz, and a second emphasized harmonic is at about 7850 IIz. Food. — Ten stomachs contained a \a- riety of small arthropods including three beetles, two flies, two orthopterans, two spiders, and one each of hemiptera, lepi- EQUATORIAL HERPETOFAUNA 161 doptera, and homoptera. The largest prey item was a roach 8 mm in length contained in a frog having a snout-vent length of 19 mm. Remarks. — Duellman (1972c) re- viewed the status of Hijla rhodopepla and showed that it belonged in the Hyla microcephala group, which is wide- spread and species rich in the lowlands of tropical America. Hijla rliodopcpla is the only species in the group in the upper Amazon Basin, although HyJa mathias- soni occurs in the llanos of central Co- lombia. Hyla riveroi Cochran and Coin ( Figure 94 ) Ht/la riveroi Cochran and Coin, 1970, Bull. ' U. S. Natl. Mus., 288:284 [Type locality.— Leticia, Amazonas, Colombia]. Material. — Santa Cecilia, 10. Identification. — This small, yellowish tan frog has a brown canthal stripe and two or three cream spots on the upper lip. The canthal stripe continues as a dorsolateral stripe to the scapular or sac- ral regions, and tan pigment is present as a narrow stripe on the dorsal surface of the thigh. Other small hylid frogs in the area having pale markings on the upper lip include H. hokernianni, hrevi- frons, and parviceps, all of which have cream or yellow spots on dark thighs. Furthermore, the labial markings consist of one (H. parviceps) or two (H. hoker- manni and //. hrevifrons) distinct sub- orbital, vertical bars. Superficially H. riveroi resembles H. rhodopepla, in which the thighs are unpigmented, the dark stripe is lateral and extends to the groin, and the lips are white. The small yellowish tan H. rossaUeni lacks labial markings. The snout is round in dorsal view and in profile. The tympanic ring is evident ventrally. An axillary membrane extends about three-fourths of the length of the upper arm. The fingers are about one- fourth webbed, and the toes are about two-thirds webbed. Breeding males do Fig. 94.— Hyla riveroi, $, 20 mm SVL (KU 126341). not have nuptial excrescences. The skin is smooth on the dorsum and granular on the belly. At night the dorsum is pale yellow with tan markings. By day, the dorsum is tan with brown markings con- sisting of flecks on the back, transverse bars on the limbs, and indistinct canthal and supratympanic stripes. In most indi- viduals some form of a brown mark is present in the occipital-scapular region. The edge of the lower lip and two or three round labial spots below the eye are pale yellow. The dorsal surfaces of the thighs are tan; other surfaces of the thighs, ventral surfaces of the limbs, and the posterior part of the belly are un- pigmented. The anterior part of the belly is white. Males have yellow vocal seas. The iris is pale red at night, changing by day to pale dull bronze with brown flecks and an orange ring around the pupil. $ 20 mm; 9 23 mm. Occurrence. — One individual was about 3 m above the ground on a branch in secondary forest; eight were on low vegetation in swamps in secondary for- est, and one was on low vegetation in a clearing. All were found at night in March through June. Life History. — Males were calling from emergent vegetation in forest swamps in March, April, and June. The single female contained only small ovar- ian eggs. Presumably this species is a sporadic breeder and deposits its eggs in the water in forest swamps. 162 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Tadpoles. — The tadpoles are un- known. Mating Call. — Analysis of recordings of two individuals from Santa Cecilia shows the call consists of short, high- pitched, insect-like notes repeated at a rate of 37-46 (X=41.5) notes per minute having a duration of 0.02-0.08 (X=0.053) sec. The dominant frequency is at about 4310 Hz. Food. — One individual contained three flies, each 5 mm in length. Remarks. — This small frog has been found only occasionally at Santa Cecilia. Most were taken in a //condary forest, and one was on a hinb of a bush at the edge of the forest. Two were ol^tained in May and one eacli in July and Octo- ber. Life History. — No observations are available. The single female from May contained 1430 ovarian eggs. Presum- ably P. coriacea, like other members of the genus, deposits eggs in a film on the surface of ponds. Tadpoles. — The tadpoles are un- known. Matinfi, Call. — The call is unknown. Food. — The stomachs of two individ- uals contained a roach, orthopteran, ant, beetle, and spider. Phrynohyas venulosa (Laurenti) (Figure 101) Rana venulosa Laurenti, 1768, Specimen medi- cum. . . Reptilium:31 [Type locality. — based on Plate 72, fi^. 4 in Seba, 1734]. Plinjnohyas venulosa — Hemming, 1958, Opin, Declar, Intl. Comm. Zool. Nomencl., 19: 172. Duelinian, 1971, Occas. Papers Mus. Nat. Hist. Univ. Kansas, 4:11. Material. — Lago Agrio, 1. Identification. — This large tree frog has thick, glandular, dorsal skin, usually l)earing many large, low tubercles. The dorsum is gray, tan, or brown, usually with darker markings. The species is most easily confused with Osteocepha- his taiirintis, which differs by having thin skin and a pair of frontoparietal ridges. Fhriinolujas vcntdosa diff^ers from P. co- riacea by lacking red thighs and web- bing. The snout is bluntly rounded, and the head is nearly as wide as long. The skin on the dorsum is thick, glandular, and usually tuberculate. A heavy dermal fold covers the upper part of the tym- panum, which is about three-fourths of the diameter of the eye. The fingers are about one-third, and the toes two-thirds, webbed. The single specimen from Lago Agrio had a greenish gray dorsum with irregular brown marks on the back, brown transverse bars on the limbs, and dark brown spots on the flanks. The venter was greenish cream, and the iris was bronze with radiating black streaks. i 75 mm. Occurrence. — The single specimen EQUATORIAL HERPETOFAUNA 175 Fig. 101. — Phrtjnohyas venulosa, S, 75 mm SVL (KU 126657). was obtained from a felled tree in pri- mary forest. Life History. — No observations were made in Ecuador, but data obtained on this species elsewhere in the Amazon Basin (Leticia, Colombia, and Belem, Brasil ) and in Central America probably pertain to Ecuadorian populations. Breeding usually occurs only after tor- rential rains. Males call from the water or from bushes and trees around ponds. The eggs arc deposited in a film on the surface of temporary ponds. Tadpoles-. — The following description is based on a tadpole (KU 124955) in stage 3S having a body length of 17 mm and a total length of 49 mm from Leticia, Colombia. The body is ovoid, as wide as deep, and deepest posteriorly. The snout is bluntly rounded. The eyes are moderately large and directed laterally. The nostrils are directed anterolaterally at a point midway between the eyes and the tip of the snout. The opening of the sinistral spiracle is directed posterodor- sally at a point below the midline at about the midlength of the body. The anal tube is short and sinistral. The cau- dal musculature is moderately slender and attenuate. The fins are deeper than the musculature and are deepest at about two-fifths of the length of the tail. The dorsal and ventral fins are equal in depth, and the dorsal fin extends onto the body. The tail is about 65 percent of the total length. The mouth is moderately small and directed anteroventrally. The me- dian part of the upper lip is bare; else- where the lips are bordered by two rows of small labial papillae, except that pa- pillae are more numerous in the lateral folds. The beaks are slender and bear fine serrations. The upper beak is in the form of a broad arch, and the lower beak is broadly V-shaped. There are three upper and six lower rows of denticles; the first upper row is broadly interrupted medially, and the last lower row is much shorter than the others. The body is brown above with a few scattered dark flecks; the belly is dull white. The cau- dal musculature is cream with dark brown stripes. The fins are transluscent with a bluish gray tint posteriorly and brown flecks and reticulations on the dorsal fin. The iris is reddish bronze. Tadpoles were in a pond in a clearing in rainforest near Leticia, Colombia. They oriented head-up, with the axis of the bodv at about 60° from the surface. The snouts were just below the surface. When disturbed they took refuge in de- bris on the bottom of the pond. Mating Coll. — The distinctive call of P. venulosa was not heard at Santa Ce- cilia. The call is a loud growl. Duellman (1970b: 168) noted that analysis of re- cordings made in Panama and Costa Rica showed that the averages for various pa- rameters are: 47 notes per minute, 0.30 sec duration, 161 pulses per second, fundamental frequency of 159 Hz, and dominant frequency of 1622 Hz. Food. — The single specimen con- tained no food, but examination of speci- mens from elsewhere indicates that a wide variety of moderately large arthro- pods is eaten. Re/?iflric5'.— Duellman (1971) noted that 12 of 13 specimens from Amazonian Ecuador had uniformly brown dorsal col- oration, whereas the other individual (from Lago Agrio) had a pattern con- sisting of anterior and posterior dorsal 176 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY markings. Proportionately, plain individ- uals are most abundant in the upper Amazon Basin. The largest male from Amazonian Ecuador is 92 mm, and the largest female is 110 mm. Ph\ lloniedusa palliata Peters (Figure 102) PJu/Uomedusa palliata Peters, 1872, Monatsb. Akad. Wiss. Berlin, 1872:773 [Type local- ity.—"Ucayali" (? = Rio Ucayali), De- partamento Loreto, Peru]. Duellman, 1974, Heipetologica, 30:107. Material. — Dureno, 1; Santa Cecilia, 94+5. Identification. — This .small Phyllome- dusa differs from its congeners by having small digital discs, and the sides of the head and flanks cream with brown flecks. All other green frogs in the area have the sides of the head green. Only P. tomop- tcrna has patterned flanks (orange with vertical brown bars); it diff'ers further from P. palliata by having moderately large discs and large calcars (absent in P. palliata). The snout is rounded above and in males truncate below, in females inclined anteroventrally below. The parotoid glands are low, rounded, and extend onlv to the shoulder. The skin on the dorsum is smooth. The discs are small; webbing is absent, and the first toe is much longer than and opposable to the second. Cal- cars are absent. The dorsum and the sides of the head above the nostrils and middle of the orbits are dark green. The lower sides of the head and the flanks are cream with brown flecks. The groin and hidden surfaces of the limbs are orange with irregular brown marks. The venter is cream with brown flecks. The iris is bronze with fine black reticulations. Breeding males have brown nuptial ex- crescences. (^45 mm; 9 50 mm. Occurrence. — Individuals were about equally abundant in primary and secon- dary forest and in forest-edge habitats, where they usually were associated with temporary or permanent ponds. At most breeding sites the frogs were most com- FiG. 102. — Phi/Uomedusa palliata, $ , 44 mm SVL (KU 126661). mon in low ( < 1 m ) herbaceous growth above the water. Individuals were ob- served in March through August and in October and December. Life History. — Breeding occurs spo- radically throughout the year. Males call from low, emergent vegetation in ponds in and at the edge of the forest. Eggs are deposited on the upper edges of leaves, usually in a tear-drop shape ofl^ of the tip of the leaf. Six clutches con- tained 38-71 (X=49.3) unpigmented eggs having diameters of 3.5-4.0 (Xr= 3.6) mm. Fifteen gravid females con- tained 38-106 (X=60.S) mature ovarian eggs. Apparently the entire ovarian com- plement is not necessarily deposited in one clutch. Five metamorphosing young have snout- vent lengths of 20-21 (X= 20.4) mm. Tadpoles. — The following description is based on tadpoles (KU 124214) in stage 37 having a body length of 15 mm and a total length of 48 mm. The body is elongately ovoid, slightly deeper than wide, deepest at midlength of the body. The snout is truncate. The eyes are large, widely separated, and directed laterally. The nostrils are dorsal and di- rected anteriorly at a point about one- third of the distance from the tip of the snout to the eyes. The spiracular open- ing is directed posteriorly just left of the midline at a point at about the midlength of the body. The anal tube is short and dextral. The caudal musculature is ro- EQUATORIAL HERPETOFAUNA 177 bust proximally and gradually diminishes to a filamentous tip. The fins are shal- low; the ventral fin is deepest, and at the midlength of the tail it is equal to two- thirds of the depth of the musculature. The fins do not extend to the tip of the tail, nor does the dorsal fin extend onto the body. The tail is about 70 percent of the total length. The small mouth is terminal; the upper lip is bare, whereas there is one row of small labial papillae laterally and two rows ventrally. There is no lateral fold. The beaks are mod- erately robust and bear small serrations. The upper beak is in the form of a highly curved arch, and the lower beak is V- shaped. There are two upper and three lower rows of denticles, of which the second upper row is broadly interrupted medially. The body and tail are bluish gray with an iridescent bluish green tint on the belly. The tadpoles occur in open ponds where they orient head-up at an angle of about 45° to the surface. Usually they are found singly or in groups of less than 10. Upon being disturbed they seek ref- uge amidst vegetation or detritus on the bottom. Mating Call. — The call is a soft "click" repeated at intervals of several seconds to three minutes. Analysis of two record- ings made at Santa Cecilia reveals that the notes have a duration of 0.13-0.14 (X=0.135) sec and 180-200 (X=190) pulses per second. The fundamental fre- quency is at about 1500 Hz and the domi- nant frequency at 3000 Hz. Food. — In 20 stomachs orthopterans and homopterans were the most fre- quently taken prey (40% and 35% of the stomachs, respectively). Spiders were present in one-fourth of the stomachs, whereas each of three frogs had eaten a fly, a beetle, and a caterpillar. Remarks. — Although P. palliafa and P. tarsiiis commonly were found at the same breeding sites at the same time, ob- servations at one site throughout one year revealed that gravid females of only one species were present on a given Fig. 103. — PhtiUomedusa tarsius, $, 88 mm SVL (KU 123193). night. The presence of one or the other was sporadic, not seasonal. Phyllomedusa tarsius (Cope) (Figure 103) PitJiccopiis tarsius Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, 20:113 [Type locality.— Rio Amazonas, below mouth of Rio Napo, Departamento Loreto, Ecuador]. PliyUomedusa tarsius — Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.: 428. Duellman, 1974, Herpetologica, 30:108. Material. — Diueno, 1; Santa Cecilia, 105-f8. Identification. — This large species of PhijUonicdusa differs from other mem- bers of the genus by having rounded parotoid glands extending to the mid- body or sacrum, the flanks marked by cream, pink, or pale orange spots, and an orange iris with black reticulations. The species is most easily confused with P. vaiUanti, which has angular parotoid glands bearing a dorsolateral row of white tubercles, flanks that are green above and red below a row of longitudi- nally elliptical pale spots, and a pale gray iris. Other PhyUomediisa have orange or cream flanks with brown markings, and green frogs in other genera in the area have webbing and the first toe shorter than the second (webbing absent and first longer than second in P. tarsius). The snout is inclined anteroventrally in both sexes and is angular above in males and round in females. The paro- 178 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY toid glands arc rounded and extend to the niidbody or sacrum. The skin on the dorsum is sha greened; that on the hind hmbs is tubercuhite in hirge individuals. The discs are moderately large; webbing is absent, and the first toe is longer than, and opposable to, the second. Calcars are absent. The dorsal surfaces and sides of the head are green. The flanks are dull green or bluish green with discreet or confluent cream, pink, or pale orange spots. The anterior and posterior sur- faces of the thighs are brown or dull green with cream, pale orange, or bluish- gray spots. The throat, chest, and ven- tral surfaces of the limbs are dark brown or grayish-brown; the belly is gray or pale orange-brown. Ventral markings consist of a pair of white spots on the throat, a median white spot on the chest, and usually a pair of para-anal white spots. The iris is bright orange with bold black reticulations. Breeding males have grayish brown nuptial excrescences. $ 97 mm; $ 111 mm. Occurrence. — Phyllomcdusa tarsius was found in March through October, with most individuals being observed in April through June. All were found in forest at night. Non-breeding individ- uals usually are perched on limbs 2-6 m alcove the ground. Life History. — This species sporadi- cally breeds after rains. Males have been heard calling from limbs of trees and bushes over ponds at various times throughout the year. Several hundred of these large frogs congregated at a forest pond bordered by spiny palms on the night of 2 April 1972. As many as four individuals were observed on one palm frond. Two nights later only a few indi- viduals were found. The unpigmented eggs are deposited on or between leaves over ponds. Usually the frogs fold leaves over the clutch so as to conceal the eggs. Eight clutches contained 426-650 (X= 551) eggs having diameters of 3.5-4.0 (X^3.7) mm. These clutches hatched in 7.0-8.3 (X=7.4) days; hatchlings had to- tal lengths of 11.5-13.0 (X=12.6) mm. Fifteen females contained 410-750 (X= 549) mature ovarian eggs; thus it seems as though this species usually deposits the entire ovarian complement as a single clutch. Fourteen metamorphosing young snout- vent lengths of 20-29 (X=25.3) mm. Tadpoles. — The following description is based on tadpoles (KU 153260) in state 36 having a body length of 1(S mm and a total length of 50 mm. The body is ovoid, as wide as deep and deepest posteriorly. The snout is truncate. The eyes are large, widely separated, and di- rected laterally. The nostrils are directed anteriorly nearly at the tip of the snout. The opening of the sinistral spiracle is directed posteriorly nearly on the midline and at a point at about midlength of the body. The dextral anal tube is short. The caudal musculature is robust proximally, gradually tapering to a terminal point. A shallow dorsal fin is present only on the posterior two-thirds of the tail. The ven- tral fin is shallower than the musculature and extends to the tip. The tail is about 64 percent of the total length. The small terminal mouth lacks lateral folds. The upper lip is bare; there is one row of small labial papillae laterally and two rows ventrally. The beaks are moderately robust with coarse serrations. The upper beak is a narrow arch, and the lower beak is V-shaped. There are two upper and three lower rows of denticles; the second upper row is broadly interrupted medially. The dorsal surfaces of the body and tail are pale tan; the sides and venter are pale metallic green. The iris is bronze. Tadpoles orient head-up at an angle of about 45° to the surface of the water. Although some have been found singly or in small numbers, two large concen- trations were found in open water. Each congregation consisted of at least 100 individuals; all were oriented in the same way. Mating Call — The call is a moder- ately loud low-pitched "cluck." Analysis of three recordings from Santa Cecilia EQUATORIAL HERPETOFAUNA 179 provides the following data: 2.0-3.2 (X=2.7) notes per minute, SO pulses per second; and notes having a duration of 0.5-0.7 (X=30.5S) sec. The fundamental frequency at 304-522 (X=375) Hz is dominant. Food. — Twent>' of 21 stomachs ex- amined contained only Orthoptera, most of which \\'cre large and only one to a stomach. The largest orthopteran was 43 mm in length. The other stomach con- tained a spider. Taxonomic Comments. — Most refer- ences to the large Phyllomedusa in Ama- zonian Ecuador have been under the specific names P. edenfula Andersson or P. orcesi Funkhouser; Duellman (1974b) showed that those names were junior synonyms of P. tarsius (Cope), a species widespread in the upper Amazon Basin in Colombia, Ecuador, and Peru. Remarks. — The exclusive occurrence of gravid females of P. tarsius and P. pallkita is discussed in the account of P. pal J lata. Phyllomedusa tomopterna (Cope) (Figure 104) Pithccopus tomopteinus Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, 20:112 [T>pe local- ity.— Rio Amazonas, below the mouth of Rio Napo, Departamento Loreto, Peruh PliyUomcdiisa tomopterna — Boulenger, 1882, Cat. Batr. Sal. Brit. Mus.:430. Duellman, 1974, Herpetologica, 30:109. Material — Lago Agrio, l-j-l; Santa Cecilia, 61+2. Identification. — PlujUomedusa tomop- terna is readily distinguished by its col- oration - bright green dorsum, and flanks and hidden surfaces of the thighs orange with purplish brown vertical bars. The only other green frog in the area having brown markings on the flanks and thighs is P. palliata, which differs from P. tom- opterna in having the sides of the head and the flanks cream with brown flecks, smaller digital discs, and no calcars. The snout is truncate in males and rounded in females. The parotoid glands are diffuse and usually not evident. The Fig. 104. — Phyllomedusa tomopterna, $ , 42 mmSVL (KU 139257). skin on the dorsum is smooth. The discs are large; webbing is absent, and the first toe is barely longer than and op- posable to the second. Large calcars are present. The dorsal surfaces and sides of the head are green. The flanks and hid- den surfaces of the limbs are orange with \'ertical purplish brown bars. The throat and chest are white, and the belly is pale orange. The iris is silvery gray. Breed- ing males have brown nuptial excres- cences. $ 48 mm; 9 57 mm. Occurrence. — All individuals of this species were found at night on bushes and low trees in forest from March through November. Life Hi;?^07-f/.— Although P. tomop- terna has an extensive breeding season, it is doubtful if it breeds throughout the >'ear. Data are lacking for January, Feb- ruary, and December, usually the driest months at Santa Cecilia. Males call from bushes and trees over ponds. Five gravid females from April and July contained 60-89 (X=71.4) unpigmented ovarian eees 3.5 mm in diameter; no clutches were found. Tadpoles. — The following description is based on a tadpole (KU 152320) in stage 38 having a body length of 23 mm and a total length of 64 mm. The body is elongately ovoid, as wide as deep and deepest posteriorly. The snout is trun- cate. The large eyes are widely separated and directed laterally. The nostrils are dorsal and directed anteriorly at a point 180 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY one-third the distance from the tip of the snout to the eyes. The opening of the sinistral spiracle is just off the midline and directed posteriorly at about the midlength of the body. The anal tube is short and dextral. The caudal muscula- ture is robust proximally and gradually diminshes to a terminal filament. The dorsal fin is very low; the ventral fin is slightly shallower than the musculature and extends to the tip of the tail. The tail is about 65 percent of the total length. The small temiinal mouth lacks lateral folds. The upper lip is bare; else- where the lips are bordered by two rows of small labial papillae. The beaks are moderately robust and bear fine serra- tions. The upper beak fomis a broad arch, and the lower beak is in the form of a flared V. There are two upper and three lower rows of denticles; the second upper row is broadly interiaipted me- dially. The body and tail are yellow or yellowish tan; the belly is white. A dif- fuse orange spot is present on the pos- terior part of the ventral fin. The iris is silvery white. Mating Call. — The call is a soft "cluck." Food. — Three of six stomachs con- tained a roach each, and the other three each contained an orthopteran, one of the latter also had an homopteran. Remarks. — Phyllomedusa tomopterna, despite its gaudy appearance, is the least conspicuous Phyllomedum in the area. It seems to prefer more dense vegetation than the other species. Half of our speci- mens were obtained from a palm-shroud- ed pond on 2 and 4 April 1972. Phyllomedusa vaillanti Boulenger (Figure 105) Phyllomedusa vaillanti Boulenger, 1882, Cat. Batr. Sal. Brit. Mii.s.:427 [Type locality.— Santarcm, Estado do Pani, Brasil]. Duell- nian, 1974, Herpetologica, 30:109. Material. — Dureno, 1; Puerto Libre, 1; Santa Cecilia, 64+2. Identification. — Phyllomedusa vail- FiG. 105. — Phyllomedusa vaillanti, 9, 49 mm SVL (KU 139252). lanti is a large green tree frog without webbing and characterized by angular parotoid glands bearing a dorsolateral row of small white tubercles. It is most similar to P. tarsius, which has round parotoid glands and no white tubercles, and an orange iris with black reticula- tions (pale gray in P. vaillanti). Other Phyllomedusa in the area have cream or orange flanks with brown markings; in P. vaillanti the flanks are green above and red below a row of pale spots. The snout is angular above and nearly truncate in males, rounded above and in- clined anteroventrally in females. The parotoid glands are slightly elevated, an- gular, and extend to the sacrum; there is a longitudinal row of white granules along the angle of the gland. The skin on the dorsum is rough; it is tubercular on the hind limbs in large individuals. The discs are moderately large; webbing is absent, and the first toe is longer than, and opposable to, the second. Calcars are absent. The dorsum and side of the head are green; the flanks are green above and reddish brown below a row of longitudinally elliptical cream to pale orange spots. The anterior and posterior surfaces of the thighs are reddish brown or lavender with small cream to pale EQUATORIAL HERPETOFAUNA 181 orange spots. The venter is dull grayish brown with a pair of lavender-bordered cream spots on the throat and a pale green spot on the chest. The iris is pale gray with a greenish tint. Breeding males have grayish brown nuptial excrescences. S 58 mm; 9 84 mm. Occurrence. — Individuals of this spe- cies were found in primary and secon- dary forest at night in all months, except November and December. Adults were on bushes and trees to heights of 4 cm. Life History. — Males call from bushes over ponds, but no breeding congrega- tions were encountered. Five gravid fe- males from March, April, June, and Au- gust contained 912-1250 (X=1114) ma- ture ovarian eggs having diameters of 2.0 mm. One female deposited a clutch of 645 unpigmented eggs 2.0 mm in diameter. The eggs are deposited on vegetation above ponds. Eleven meta- morphosing young from February (2), June (1), and July (8) have snout-vent lengths of 21-25 (X=23.8) mm. Tadpoles. — The following description is based on a tadpole (KU 152319) in stage 38 having a body length of 19 mm and a total length of 52 mm. The body is elongately ovoid, slightly deeper than wide. The snout is truncate. The large eyes are widely separated and directed laterally. The nostrils are directed an- teriorly nearly at the tip of the snout. The spiracular opening is directed pos- teriorly just to the left of the midline at a point at about the midlength of the body. The anal tube is short and dextral. The caudal musculature is robust proxi- mally and gradually narrows to a slender • tip. The dorsal fin is very shallow, and the ventral fin is shallower than the mus- culature and extends to the tip of the tail, which is about 63 percent of the total length. The small terminal mouth lacks lateral folds. The upper lip is bare; elsewhere the mouth is bordered by two rows of small labial papillae. The beaks are moderately robust with fine serra- tions. The upper beak is a broad arch with slender alary processes; the lower beak is broadly V-shaped. There are two upper and three lower rows of denticles; the second upper row is broadly inter- rupted medially. The dorsum is olive green, paler on tail than on body. The belly is white, and the ventrolateral sur- faces are iridescent vellowish green. The iris is pale bronze. Mating Call — The call consists of a short, harsh, "cluck." Analysis of one recording from Pilcopata, Departamento Cuzco, Peru, shows 13.3 notes per minute having a duration of 0.5 sec, 240 pulses per second, and a dominant frequency at about 1800 Hz. Food. — Sixteen of 18 stomachs con- tained orthopterans up to 34 mm in length. The other two contained spiders. Taxonomic Comments. — Most refer- ences to this species in the upper Ama- zon Basin ha\'e been under the name of PJiyllomedusa perlafa, a name synony- mized with P. vaiUanti by Duellman (1974b). Remarks. — PhijUomedusa vaiUanti ap- parentK' breeds at sites exclusive of those used by the other three species of FlujUo- medusa. At two ponds where the other species called commonly at various times throughout the year, P. vaiUanti was never found. Instead individuals of P. vaiUanti were scattered throughout the forest or in small groups at temporary ponds. Sphaenorhynchus carneus (Cope) (Figure 106) Hylella carnea Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, 1868:111 [Type locality.— "Napo or upper Maranon" ( ? Peru)]. Sphaenorhijnciius carneus — Duellman, 1974, Occas. Papers Mus. Nat. Hist. Uniw Kan- sas, 27:22. Material. — Santa Cecilia, 19. Identification. — This small green tree frog with a protruding snout has cream (night) or red (day) canthal and dorso- lateral stripes. This color pattern resem- bles that of the larger Hijla punctata which has a round, non-protruding snout and less webbing on the feet. The much 182 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 106. — Sphaenorhtjnchus carneus, $ , mmSVL (KU 146576). 19 larger Sphaenorhynchus eurhostus also has a protruding, acuminate snout, but it lacks red dorsolateral stripes and has more webbing on the hand. All other small green frogs in the area have round- ed snouts, except Eleutherodactylus acu- minatus, which differs from S. carneus in lacking webbing and a tympanum. The snout is narrowly truncate in dor- sal view, acuminate and protruding well beyond the jaw in profile. The anterior edge of the tympanum is evident. The skin of the dorsum is smooth; that on the venter is weakly granular. The fingers are about one-third webbed and the toes three-fourths webbed. At night the dor- sum is pale green with golden cream can- thai and dorsolateral stripes. By day the dorsum is dark green with reddish brown flecks and faint greenish cream canthal and dorsolateral stripes bordered below by reddish brown stripes. The belly is white, and other ventral surfaces are pale green. The axilla and groin are blue, and the webbing is unpigmented. The iris is pale silver, and the bones are green. Breeding males have brown nup- tial excrescences, i 20 mm; 9 23 mm. Occurrence. — All individuals were on emergent vegetation in a pond in pri- mary forest; they were observed there on five nights in April and May, 1972, and on one night in June 1975. Life History. — Males call from emer- gent vegetation. One female contained 143 mature ovarian eggs. No other data on life history are available. Mating Call. — The call is a series of loud clicks. Analysis of three recordings from Santa Cecilia revealed that notes are produced at a rate of 23-180 (X=77) notes per minute. The duration of the notes is 0.02-0.05 (X=0.025) sec, and the pulse rate is 180 pulses per second. The dominant frequency is 4750-5000 (X=4880) Hz. Food. — Six individuals contained only ants, Sphaenorhynchus eurhostus Rivero (Figure 107) Sphaenorhynchus eurJwstus Rivero, 1969, Co- peia ( 1969(4) :701 [Type locality.—"? Brasil"]. Material. — Santa Cecilia, 1. Identification. — This moderate-sized green tree frog has an acuminate snout protruding well beyond the margin of the jaw. Only the much smaller Sphae- norhynchiis carneus has a protruding snout, but that species has cream dorso- lateral stripes and less webbing on the hand. Other green frogs in the area have rounded, truncate, or acuminate snouts that do not protrude beyond the margin of the jaw; furthenuore, all have less webbing on the hand than does S. eurho- stus. The snout is acuminate in dorsal view and pointed and protruding well anterior to the margin of the lip in profile. The tympanum is barely evident. The fingers are half and the toes four-fifths webbed. The skin on the dorsum is smooth; that on the belly is weakly granular. The dorsum is green; the belly, ulnar, tarsal, and anal stripes are creamy white, and the canthal stripe is dark brown. The ventral surfaces of the limbs are bluish green, and the iris is pale creamy bronze. EQUATORIAL HERPETOFAUNA 183 Fig. 107. — Sphaenorhynchus eurhosttis, juve- nile, 21 mm SVL; Belem, Estado do Para, Bra- sil (KU 127981). Breeding males have brown nuptial ex- crescences. 9 48 mm. Occurrence. — The single specimen from Santa Cecilia was on a bush at the edge of primary forest at night in August. Life History. — The single female con- tained 550 mature ovarian eggs. Obser- vations on this species in Brasil and Venezuela indicate that the males call on vegetation overhanging ponds at the edge of forest. Presumably the eggs are deposited in ponds. Tadpoles. — The tadpoles are un- known. Mating Call. — The call consists of a short, low pitched note, folowed or not by a secondary note. Analysis of two re- cordings from Georgetown, Guyana, re- veals that notes are produced at a rate of 20-24 (X=22) notes per minute. The notes have a duration of 0.05-0.10 (X= 0.07) of a second and a rate of 200-240 (X^217) pulses per second. The funda- mental frequency is at 350-410 (X= 391) Hz and the dominant frequency at 1050-1230 (X=1183) Hz. Food. — The single individual con- tained two ants. Remarks. — A male from Limoncocha, Ecuador, has a snout-vent length of 41 mm. CENTROLENmAE Centrolenella midas Lynch and Duellman (Figure 108) Fig. 108. — Centrolenella midas, $ , 20 mm SVL (KU 123219). Centrolenella midas Lynch and Duellman, 1973, Occas. Papers Mus. Nat. Hist. Univ. Kansas, 16:38 [Type locality. — Santa Cecilia, Pro- vincia Napo, Ecuador]. Material. — Lago Agrio, 2; Puerto Libre, 4; Santa Cecilia, 7+1- Idefitification. — This small centrolenid differs from the other species by having a dark green dorsum with scattered gold flecks dorsolaterally on the body. It fur- ther differs from C. resplendens by lack- ing lateral dermal fringes on the fore- arms and feet and by having a truncate instead of a sloping snout. Centrolenella midas also differs from C. munozorum in snout shape (round in C munozorum) and in having the lower two-thirds of the tympanum visible (tympanum en- tirely concealed in C. munozorum). Cen- trolenella midas differs from Hijla gra- nosa in snout shape, dorsal coloration, and smaller size; H. granosa has a round snout and unifonn green dorsum, in ad- dition to having rounded discs on the fingers (truncate in Centrolenella) . The shape of the discs also distinguishes C. midas from species of Sphaenorhynchus, which have a pointed snout protruding anteriorly beyond the margin of the lip. The green Eleutherodactylus acuminatus has no webbing on the hand and has a black canthal stripe. The dorsal surfaces of the head, body, and limbs are dark green with a few small golden yellow flecks dorsolaterally 184 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY on the body (in preservative lavender with white flecks). The hands and feet are dull greenish yellow. The chest is white, and the heart is not visible; the visceral peritoneum is white. The bones are green, and the iris is silvery bronze with black reticulations. $ 19 mm; 9 26 mm. Occurrence. — Two were obtained from tree tops as trees were felled by day. Others were found on low vegeta- tion at night in both primary and secon- dary forest. The specimens from Puerto Libre and three of those from Santa Ce- cilia were on bushes and herbs adjacent to, or overhanging, small shallow streams. Individuals were found in April-August and November. Life History. — Two females (April and August) show no evidence of repro- ductive activity, and two others (April and May) have small ovarian eggs. Four females (June, July 2, November) have 14-29 (X=22) large ovarian eggs. A tadpole tentatively referred to this spe- cies was found in June. Tadpoles. — A single centrolenid tad- pole was found in the shallow stream at Santa Cecilia where numerous Centrole- nella midas were heard and a few col- lected. No other centrolenids were found along the stream, so the tadpole (KU 124218) is tentatively referred to C. midas. The tadpole is in developmental stage 25 and has a body length of 6.2 mm and a total length of 19.0 mm. The body is elongate and depressed; the eyes are small, close together, and directed dorso- laterally. The mouth is directed antero- ventrally and is bordered laterally and ventrally by one row of small papillae. The beaks are robust and serrate. One upper and two lower rows of denticles are present. The caudal fins are low; only on the posterior one-third of the tail are the fins deeper than the robust caudal musculature. The tadpole essentially lacks pigment in the skin. Mating Call. — The call is a high- pitched triple note "peep-peep-peep" re- FiG. 109. — Centrolenella munozorum, 9 , 23 mmSVL (KU 123225). peated at intervals of more than 20 sec- onds. Food. — Two individuals contained food - one spider and one homopteran. Centrolenella munozorum Lynch and Duellman (Figure 109) Centrolenella munozorum Lynch and Duellman, 1973, Occas. Papers Mus. Nat. Hist. Univ. Kansas, 16:40 [Type locality. — Santa Ce- cilia, Provincia Napo, Ecuador]. Material — Lago Agrio, 1; Santa Ce- cilia, 12+1. Identification. — This small centro- lenid difi^ers from the other species by having white bones and a clear parietal peritoneum resulting in the heart being visible through the ventral skin. Further- more, it differs by having a round snout (truncate in C. midas and sloping in C. resplendens) . Centrolenella munozorum has a pale green dorsum with large, dif- fuse yellow spots, contrasting with many small white spots in C. resplendens and few small gold flecks in C. midas. Cen- trolenella munozorum further differs from C. resplendens by lacking lateral dermal fringes on the forearms and feet. Hylids (Sphaenorhynchus and Hyla gra- nosa) differ by having rounded, instead of truncate, discs on the fingers and either uniformly green dorsal surfaces or red dorsolateral stripes ) . The only green Eleutherodactylus (E. acuminatus) lacks EQUATORIAL HERPETOFAUNA 185 webbing on the hand and has a black canthal stripe. The dorsum is pale green with diffuse pale yellow or greenish yellow spots (in preservative, creamy white with minute black flecks). The limbs are pale green, except for the thighs, which are unpig- mented. The iris is pale gold. S 20 mm; 9 21 mm. Occurrence. — One was obtained from the canopy of a large tree felled in pri- mary forest. Others were found on vege- tation in or near swamps in primary or secondary forest at night and along small trickling streams at the edge of the forest. Individuals were found in April through October. Life History. — A female (July) con- tained 18 large ovarian eggs. Males were calling from low vegetation over a rivulet in September and October; tadpoles were found in the stream in October. Tadpoles. — The following description is based on tadpoles (KU 155524) in stage 25 having a body length of 4.2 mm and a total length of 13 mm. The body is elongately and narrowly o\'oid, wider than deep. The snout is bluntly rounded. The eyes are small and directed dorso- laterally. The nostrils are directed an- terodorsally about midway between the snout and the eyes. The opening of the sinistral spiracle is directed posterodor- sally at a point just below the midline at about midbody. The anal tube is long and dextral. The caudal musculature is robust, fonning more than half of the depth of the tail. The fins are about equal in depth to one another, except that the dorsal fin is very low anteriorly and does not extend onto the body. The tip of the tail is rounded. The tail is about 70 percent of the total length. The mouth is small, directed anteroventrally, and bordered laterally and ventrally by one row of small papillae. The beaks are moderately robust and finely serrate. There are two rows of denticles below the beaks and none above. Except for a few dark flecks on the dorsal edge of the caudal musculature, pigment is lacking. Fig. 110. — Ccntrolcnclla resplendens, S, 27 mmSVL (KU 118053). Mating Call — The call is a soft, high- pitched, single note. Food. — One individual contained a spider, and two contained small beetles. Centrolenella resplendens Lynch and Duellman (Figure 110) Ccntrolcnclla resplendens Lynch and Duellman, 1973, Occas. Papers Mus. Nat. Hist. Univ. Kansas, 16:51 [Type locality. — Santa Ce- cilia, Piovincia Napo, Ecuador]. Material. — Santa Cecilia, 1. Identification. — This, the largest cen- trolenid in the area, differs from all other frogs there by having a dark green dor- sum with many white tubercles, a long sloping snout, a U-shaped anal fold, and a scalloped dermal fold along the outer edge of the hand, forearm, and elbow, and another along the outer edge of the heel, tarsus, and foot. Other centrolcnids and green hylids and Eleiitherodactylus have smooth skin on the dorsum, lack scalloped dermal folds on the limbs, and have round or truncate snouts. The only other frog in the area having scalloped folds on the limbs is the larger Hyla niarmorata, which is mottled gray, green, and brown above and black, white, and yellow below. The dorsum is dark green with small white to pale bluish green spots on tu- bercles; the fringes on the limbs and around the anus are white. The venter is yellow, and the iris is gray. S 27 mm. 186 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Occurrence. — The only known speci- men from Santa Cecilia was on the leaf of a bush about 30 cm above the ground in primary forest. It was found during a light rain on the night of 14 June 1967. Life History. — Nothing is known; pre- sumably the species deposits its eggs on vegetation overhanging streams like centrolenids. A juvenile having a snout- vent length of 16.5 mm was found in October at Santa Maria de Sucumbios, Colombia (40 km NNW of Santa Ce- cilia). Tadpoles. — The tadpoles are un- known. Mating, Call. — The call is unknown. Food. — The only known adult, the holotype, was not examined for stomach contents. RANroAE Rana palmipes Spix ( Figure 111 ) Rana palmipes Spix, 1824, Animalia noxa . . . testudinum et rananim . . . Biasilian, p. 5 [Type locality. — "Amazonenfluss"]. Coch- ran and Coin, 1970. Bull. U. S. Natl. Miis., 288:70. Material. — Puerto Libre, 1; Santa Ce- cilia, 97+7. Identification. — This large smooth- skinned frog with fully webbed toes and large tympanum is readily distinguished from all other frogs in the area. The species of Lcptodacfylus have much less, if any webbing on the feet and have a ventral abdominal disc (absent in Rana). The only other frog that might be con- fused with Rana palmipes is Hyla lanci- formis, which differs by having large terminal discs on the fingers and toes. The body is moderately robust, and the snout is acuminate in dorsal view and round in profile. The tympanum is nearly as large as the eye. The fingers are unwebbed and webbing extends to the terminal phalanges of all but the fourth toe. The dorsum varies from bright green to olive-green or olive-tan, usually always with a green snout. Some individuals have irregular dark brown Fig. 111. — Rana palmipes, $, 89 mm SVL (KU 146639). or black spots on the dorsum. The flanks are tan with dark brown or black spots. The venter is creamy yellow with a gray- ish suffusion on the throat and black flecks on the ventral surfaces of the limbs and on the posterior part of the belly. The posterior surfaces of the thighs are black with creamy yellow reticulations. The iris is reddish brown with a bright bronze cast dorsally. i 101 mm; 9 126 mm. Occurrence. — Rana palmipes is wide- ly distributed in a variety of habitats, but SO percent of the specimens were found within 3 m of water. Some juveniles were found under logs or in small streams by day, but adults were found only at night. Thirty-five percent of the frogs were in, or at the edge of, swamps or lakes, and 30 percent were in, or at the edge of streams, and 5 percent were on the bank of the river. The frogs were found in clearings and forest. Adults were found throughout the year. Life History. — Males were heard call- ing throughout the year from lakes, swamps, streams, and the river. Cravid females were found throughout the year, except December and January, when no females were found. Twenty-five females having snout- vent lengths of 103-125 (X=117) mm contained 720-6750 (X= 2860) ovarian eggs. Eggs are deposited in clumps in ponds or in quiet pools in streams. Two metamorphosing young EQUATORIAL HERPETOFAUNA 187 have snout-vent lenc^ths of 48 and 53 mm. Tadpoles. — Tlie following description is based on a tadpole (KU 152332) in stage 35 having a body length of 37 mm and a total length of 92 mm. The body is ovoid, as wide as deep, and deepest posteriorly. The snout is bluntly round- ed. The eyes are moderately large, widely separated, and directed laterally. The nostrils are directed anteriorly at a point about two-thirds of the distance from the eyes to the snout. The opening of the sinistral spiracle is directed pos- terolaterally well below the midline at a point about midlength of the body. The cloacal tube is short and medial. The caudal musculature is moderately robust, gradually tapering to the tip. The fins are subsequal in depth and deepest at about midlength of the tail, where the depth of each is about equal to the depth of the caudal musculature. The fins ter- minate in a rounded tip, and the dorsal fin does not extend onto the body. The tail is about 60 percent of the total length. The mouth is small and directed anteroventrally. The median part of the upper lip is bare, and the median part of the lower lip bears a single row of small papillae; elsewhere there are two rows of papillae. The lips are folded laterally. The beaks are slender and bear fine ser- rations. The upper beak is a broad, flat- tened arch, and the lower beak is a broad curved arch depressed medially. There are two upper and four lower rows of denticles; the second upper and first lower rows are interrupted medially. A short lateral row is present on either side of the beak. The dorsum is greenish brown mottled with dark brown. The belly is cream with gray spots, and the tail is orange-tan. The iris is bronze. Tadpoles are solitarily pelagic in quiet water. Mating Call. — The call is a series of guttural chuckling sounds produced in water. Food. — Examination of 17 stomachs Fic. 112. — Chiasmocleis anatipes, S, 19 mm SVL (KU 146034). revealed a variety of large prey, the larg- est being a mantid 63 mm long in a frog having a snout-vent length of 98 mm. Orthopterans (35% of stomachs), beetles (30%) and spiders (30%) were the most common prey items. Caterpillars were found in two stomachs and each of the following in one: diptcran, hymenop- teran, homopteran, odonate, and odonate n\'mph. MiCROHYLroAE Chiasmocleis anatipes Walker and Duellman (Figure 112) Chiasmocleis anatipes Walker and Duellman, 1974; Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 26:1 [Type locality. — Santa Cecilia, Provincia Napo, Ecuador]. Material. — Santa Cecilia, 13+2. Identification. — Among the microhy- lid frogs in the region, all of which have narrow snouts and small eyes, C. anatipes is the only species having nearly fully webbed feet. The much larger Cteno- phnjne geayi has toes less than one-half webbed, and the ventral surfaces are dark brown with cream flecks. In C. anatipes the venter is white with irregu- lar large black spots, a pattern shared with C. ventrimaculatas, a species in which the toes have only basal webbing. Chiasmocleis bassleri and Hamptophryne holiviana have only basal webbing on the feet, dark throats, and mottled bel- lies. The onlv other microhvlid in the 188 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY area. Syncope antenori has only four toes and a visible tympanum (tympanum concealed in others). The body is relatively slender, and the snout tapers medially, anterior to the eyes, to a rounded tip. The limbs are slender. The fingers bear lateral fringes, and the toes are webbed to the bases or middle portions of the tenuinal discs. The skin is smooth above and below with scattered minute tubercles on the dorsal surfaces of the head, body, and hind limbs. The dorsum is dull olive-green to dull brown with green and/or gold me- tallic flecks; a narrow middorsal tan stripe is present in some specimens. The upper arms are tan or orange. The pos- terior surfaces of the thighs are gray with black flecks. The throat is pale gray with bold black mottling, and the belly and the ventral surfaces of the thighs are white with large, brownish black marks. The iris is reddish brown. Breeding males lack nuptial excrescences. 6 20 mm. Occurrence. — All individuals were found in or near a semi-penuanent pond in primary forest at night. In April 1972 the frogs were on the ground or on leaves of low ( < 25 cm ) herbs, but in May 1975 six males were calling from the shal- low part of the pond. Life History. — With the exception of the presence of calling males in May and tadpoles in April, no observations are available; females have not been found. Tadpoles. — The following description is based on a tadpole ( KU 146836) in stage 37 having a body length of 9.5 mm and a total length of 32.5 mm. The body is as deep as wide, widest anteriorly with a bluntly rounded snout. In profile the snout is round. The eyes are small and directed laterally. The spiracles are paired ventrolaterally; the cloacal tube is dextral. The caudal musculature is slender, tapering gradually to a pointed tip beyond the terminus of the fins, which are deepest at midlength of the tail and about twice the depth of the caudal musculature. The dorsal fin does not extend onto the body. The tail com- prises about 69 percent of the total length. The small, terminal mouth is di- rected anterodorsally and lacks papillae and keratinous structures. The body is olive-tan above and yellowish white be- low. The tadpoles were swimming indi- vidually near the surface of the pond during the day. Mating Call. — The call consists of a short buzz. Food. — Only small ants had been eaten. Remarks. — In addition to the seven specimens reported by Walker and Duellman (1974), six adult males ob- tained by Ronn Altig in 1975 have been examined. They agree in all respects with the type series. Chiasmocleis bassleri Dunn Ciiiasmoclcis hassleri Dunn, 1949, Amer. Mus. Novitates, 1419:9 [Type locality. — Rio Uto- (luinia to Rio Tapiche, Departamento Lo- reto, Peru]. Material. — Lago Agrio, 1; Santa Ce- cilia, 7. Identification. — Like most other mi- crohylid frogs in the region, C. hassleri has a narrow snout, small eyes, and a concealed tympanum; the toes are only basally webbed, a character distinguish- ing C. hassleri from C. anatipes and Ctenophryne geayi, which have more ex- tensive webbing. The throat and chest are brown with pale flecks, whereas the belly and ventral surfaces of the limbs are white with large black spots. In C. ventriniaculata the entire venter is white with black spots. The larger llampto- phryne holiviana has a ventral pattern like that of C. hassleri, except the spots are smaller and more numerous. Fur- thermore, the dorsum in //. holiviana is pale and sharply delineated by dark flanks, whereas in C. hassleri the dorsum is dark and not diflerentiated from the coloration of the flanks. Chiasmocleis hassleri has five toes and a concealed tympanvnn, characters distinguishing it EQUATORIAL HERPETOFAUNA 189 from Syncope antenori, which has four toes and a visible tympanum. The body is robust, and the head ta- pers medially from the angle of the jaws to a pointed snout. The forelimbs are slender, and the hind limbs are moder- ately robust. The fingers lack lateral fringes, and the toes are barely webbed basally. The skin is smooth above and below. The dorsum and flanks are uni- form grayish brown. The dorsal surfaces of the forelimbs are pinkish tan. There are indistinct pink spots on the flanks and anterior surfaces of the thighs. A pair of small black spots is present in the inguinal region. The throat and chest are gray; the belly and ventral surfaces of the hind limbs are white with large black or brownish black spots. The iris is greenish silver with dark brown reticu- lations. Breeding males lack nuptial ex- crescences. S 22 mm; 9 28 mm. Occurrence. — Individuals of this spe- cies were found in equal numbers in primary and secondary forest. With the exception of one individual on an herba- ceous leaf 20 cm above the ground, all were found on the ground. The annual distribution is January (1), May (2), June ( 1 ) , July ( 1 ) , August ( 1 ) , Novem- ber ( 1 ) , and December ( 1 ) . Life History. — No observations on breeding are available. Three females from June, July, and November con- tained 235, 250, and 151 mature ovarian eggs, respectively. Tadpoles. — The tadpoles are un- known. Mating Call. — The call is unknown; presumably it is a buzzing sound like that of other members of the genus. Foorf.^Minute beetles were found in four stomachs and ants in four stomachs; also one individual had eaten two tiny wasps in addition to an ant. Chiasmocleis ventrimaculata (Andersson) Engystoma ventrimaculata Andersson, 1945, Arldv. Zool., 37A:2 [Type locality.— Rio Pastaza, Ecuador]. Chiasmolceis ventrimaculata — Dunn, 1949, Amer. Mus. Novitates, 1419:8. Material. — Santa Cecilia, 9. Identification. — Chiasmocleis ventri- maculata, like other microhylid frogs in the area, has a narrow snout and small eyes. The dorsum is dark brown, and the entire venter is white with large ir- regular black spots - characters immedi- ately distinguishing the species from Chiasmocleis bassleri, Ctenophryne geayi, and Homptophryne holiviana, all of which have dark throats and chests with pale flecks. Furthermore, in Homp- tophryne and Ctenophryne the dorsum is pale and sharply delineated from dark flanks. The ventral coloration of C. ven- trimaculata is shared with C. anatipes, which has nearly fully webbed feet; the toes are webbed basally in C. ventrimac- ulata. The only other microhylid frog in the area is Syncope antenori, a small species having four toes and a concealed tympanum. The body is moderately slender, and the head tapers medially from the angle of the jaws to an acutely rounded tip. The limbs are slender. The fingers lack fringes, and the toes are barely webbed basally. The skin is smooth above and below. The dorsum is dark olive-brown with minute gold flecks. All ventral sur- faces are bluish white with irregular bold black spots. The iris is reddish brown. Breeding males lack nuptial excres- cences. 6 23 mm; 9 22 mm. Occurrence. — All individuals were found at night in primary forest; some were on low herbs, others were on the ground, and one was calling from the edge of a shallow pond. The seven frogs were found in April (1), May (4), June (1), and July (1). Life History. — A male was calling in May, and one female from June con- tained 210 mature ovarian eggs. Recent- ly metamorphosed young were found in April and May. Tadpoles. — The tadpoles are un- known. 190 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Mating Call. — The call consists of a soft buzz. Food. — Only small ants were found in three stomachs. Ctenophryne geayi Mocquard Ctenophnjnc gcayi Mocquard, 1904, Bull. Mus. Hist. Nat. Paris, p. 308 [Type locality.— Rio Sarare Departamento Norte de Santan- der, Colombia]. Dunn, 1949, Amer. Mus. Novitates, 1419:18. Material. — Santa Cecilia, 4. Identification. — This is the largest mi- crohylid frog in the area, and it has a squat, robust body with a small pointed head. The toes are about four-fifths webbed, and the venter is dark with cream flecks - a combination of charac- ters distinguishing C. geayi from all other microhylid frogs in the area. Chias- mocleis anatipes has the toes almost fully webbed; in that species and Chiasmo- cleis ventrimacidata the venter is white with black spots. In Chiasniocleis bass- leri and Hamptophryne holiviana the throat and chest are dark with pale flecks, and the rest of the venter is white with dark spots. Hamptophryne hoUvi- ana and C. geayi are alike in having a pale dorsal color sharply delineated from dark flanks. In C. geayi the dorsal color is continuous onto the ami, whereas in Hamptophryne the pale color on the ami is separated from that on the body by the dark color of the flanks. The only other microhylid in the area is the minute Synecope antenori, which has only four toes and a visible tympanum; C. geayi has five toes and a concealed tympanum. The body is robust, the head tapers medially from the angles of the jaws to a round snout. The limbs are moderately robust. Lateral fringes are present on the fingers, and the toes are webbed nearly to the bases of the terminal discs. The skin is smooth above and below. The dorsum is pale brown with a pale cream middorsal line and minute cream flecks. The dorsal color, which is con- tinuous onto the arms, is delineated by a narrow creamy white line, separating the dorsal color from the darker brown flanks and anterior and posterior surfaces of the thighs. The vocal sac is dark grayish brown; the other ventral surfaces are dark brown with white flecks. The iris is grayish bronze. Males lack nuptial ex- crescences. S 32 mm; 9 54 mm. Occurrence. — Males were calling in May from under leaves at the edge of a semi-permanent pond in primary forest. One female lacking ecological data was collected in August. Life History. — Except for the pres- ence of calling males in May, the only reproductive infonnation available is that a female from August contained 1250 mature ovarian eggs. Tadpoles. — The tadpoles are un- known. Mating Call — The call consists of a long, low-pitched, toad-like trill. Food. — One individual contained re- mains of beetles. Hamptophryne boliviana (Parker) (Figure 113) CJiiasmocleis holiviana Parker, 1927, Occas. Pap. Mus. Zool. Univ. Michigan, 187:3 [Type locality. — Buenavista, Departamento Santa Cruz, Bolivia]. Hamptophryne holiviana. — Carvalho, 1954, Occas. Pap. Mus. Zool. Univ. Michigan, 555:7. Material. — Dureno, 2; Santa Cecilia, 64-f6. Identification. — This robust microhy- lid with a pointed snout has a pale dor- sum sharply delineated from the darker flanks; the dorsal color on the body is not continuous onto the arms. It differs from the larger Ctenopliryne geayi, in which the dorsal coloration is continuous onto the arm, by having a dark throat and chest and the rest of the venter pale with dark spots; in Ctenophryne geayi the entire venter is dark with pale flecks. The ventral coloration of //. boliviana is like that of the smaller Chiasniocleis bassleri, which differs by having a dark dorsum not differentiated from the flanks. Other microhvlids in the area either have EQUATORIAL HERPETOFAUNA 191 Fig. 113. — Hamptophrtjne holiviana, 9, 40 mm SVL (KU 124012). white venters with bold black spots (Chiosmocleis anatipes and C. ventri- macukitus) or a visible tympanum and only four toes (Syncope antenori); H. holiviana has a concealed tympanum and five toes. The body is moderately robust, and the snout tapers medially from the eyes to a narrow truncate tip. The limbs are moderately robust. The fingers and toes have distinct terminal discs and lack lat- eral fringes and webbing. The skin is smooth above and below. The dorsum is tan or grayish tan with a slightly darker, large, middorsal blotch and usu- ally a narrow, faint, creamy white mid- dorsal line. The flanks, side of the head, and anterior and posterior surfaces of the thighs are dark brown. The dark brown on the flanks separates the pale tan on the dorsum of the body from that on the arm. The throat is dark brown with creamy white flecks. The belly and ventral surfaces of the hind limbs are creamy white with brown to brownish black spots and reticulations. The iris pale bronze with fine black reticulations. Breeding males lack nuptial excres- cences. 6 38 mm; 9 44 mm. Occurrence. — About 85 percent of the specimens were in mature forest, others were in secondary forest. Most individuals were on the ground at night, but some were in shallow water in ponds and three were perched on leaves of low herbs. One was on the ground by day. Although the frogs were active through- out the year, 40 percent of the individ- uals were found in April. Life History. — Despite the abundance of this species compared to the other microhylids in the area, few data on its life history are available. Males were calling from shallow ponds in April and May. One female from October con- tained 2186 mature ovarian eggs. Tad- poles were found in April, May, and June, and metamorphosing young were found in June. Tadpoles. — The following description is based on tadpoles (KU 152564) in stage 27 having a body length of 10 mm and a total length of 30 mm. The body is depressed, about twice as wide as deep, and widest at the level of the or- bits. The snout is bluntly rounded. The eyes are small and directed laterally. The spiracle and the cloacal tube are median. The caudal musculature is slender and tapers to a slender tip. The fins are sub- equal in depth to one another and to the musculature, and the fins extend to the tip of the tail; the dorsal fin does not extend onto the body. The tail is about one-third of the total length. The mouth is terminal and directed ventrally; the fleshy upper lip nearly covers the orifice. Keratinous mouth parts are absent. The dorsum and throat are dark brown; the belly is gray with black mottling. A white stripe is present laterally on the proximal one-third of the caudal muscu- lature. Tadpoles were found in forest ponds by day when they were swimming indi- vidually near the surface. Mating Call — The call consists of a moderately long, loud "baaaah." Food. — Stomachs of 25 adults were examined. One contained only a small beetle larva. All others contained ants (as many as 47); two also contained Isoptera, and one contained a small beetle and a Dermaptera. Uamptophryne was observed feeding on ants at night. 192 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Syncope antenori Walker Syncope antenori Walker, 1973, Occas. Pap. Mus. Nat. Hist. Univ. Kansas, 20:3 [Type locality. — Puerto Libre, Proxincia Napo, Ecuador]. Material. — Puerto Libre, 11. Identification. — This minute, dull mi- crohylid frog differs from all other frogs in the area by having only four toes and, additionally, from all other microhylid frogs by having a visible tympanum. This small frog has a robust body and broadly truncate snout in dorsal view. The distinct tympanum is about half of the diameter of the eye. The fingers bear lateral fringes; the four toes (the nomial first toe absent) have terminal discs and lack webbing. The dorsum is dull brown; the venter is grayish brown with small bluish white flecks. Males lack nuptial excrescences. S 12 mm; 9 14 mm. Occurrence. — Eight individuals were in primary forest and three in secondary forest in July. All were found at night - four on leaves of low herbs and seven on the ground. Life History. — Five females contained 5-6 (X=5.4) ovarian eggs 1.2 mm in diameter. The presence of few relatively large ovarian eggs suggests that in this species eggs may be terrestrial and un- dergo direct development. Mating Call. — No call was associated with this species. The absence of vocal slits in adult males perhaps is indicative that this small frog is mute. Food. — Examination of seven stom- achs revealed the presence of ants and mites in six; one of these also contained a small beetle. The seventh individual contained a beetle, a diplopod, and a pseudoscorpion. TURTLES CHELmAE Chelus fimbriatus (Schneider) Tcsiudo fiml}riata Schneider, 1783, Allgem. Naturgescli. Schildkr.:349 [Type locality. — Surinam]. Cliclt/s finiJ)iiata — Boulenger, Cat. Chelon. Rhynchoceph. Crocod. Brit. Mus.:209. Clielus jxmhnatus — Mertens, Miiller, and Rust, 1934, Bl. Aqua.-Terr. Kunde, 45:65. Materia]. — Santa Cecilia, 3. Identification. — This flattened turtle, commonly called matamata, with a broad head and fleshy proboscis is unlike any other Amazonian turtle. The snout and parietal region are orange-tan; the rest of the head and neck are dull dark brown. The limbs are dark brown with a tan stripe along the anterior edge of the forelimb. The carapace is dark brown with a yellowish tan middorsal stripe; the tips of the keels on costal and and marginal scutes are tan. The plas- tron is dull tan. 6 330 mm; 9 345 mm. Occurrence. — All of the specimens were obtained from the Rio Aguarico by Indians. Life History. — No data are available. Mesoclemmys gibba (Schweigger) Emys gihha Schweigger, 1812, Konigsberg. Arch. Naturgesch. Math., 1:299 [Type lo- cality.— unknown]. Mesoclemmj/s ^ihha — Gray, 1873, Ann. Mag. Nat. Hist., (4) 11:306. Material. — Santa Cecilia, 5. Identification. — This side-necked tur- tle has a smooth, broad carapace that is flared laterally. It differs from Vhrynops by having a dark plastron ( light in Fhry- nop.s) and from Platemys by having a smooth carapace (two rounded longi- tudinal ridges in Platemys). Further- more, Mesoclemmys has a shallow round- ed indentation in the posterior margin of the carapace, whereas Phrynops has a deep angular indentation. The carapace and plastron are dark brown to black, and the head and neck are brown. 9 214 mm. Occurrence. — Two were brought in by natives, and two shells were found by a forest pond. Another was in a marsh in a clearing. Life History. — No data are available. Remarks. — The two recently killed females were found by a pond in primary PLATE 1 Colosicthus satili Dendiobatcs pictiis pictus Colostethus marchesianus PlujUobatcs feinoralis Edalorhina perezi © Linda Trueb PLATE 2 Hijla bifurca Htjia IcucophyUata Htjla triauiiuhim Hijla miniita Hyla favosa HtjL Ilia trianguhim Hyla marmorata © Linda Trueb PLATE 3 * Anolis trachyderma ^ Anolis chrtjsolepis scijpheus Anolis transversalis Anolis ortoni Anolis punctatus houlengeri © Linda Trueb PLATE 4 ErythroJainpnis acsciilapii aesculapii Micrurus spixii obscunis Micrtiius surinamensis surinamensis m Micrurus langsdorffi langsdoi'ffi ntUEB '?3 © Linda Trueb EQUATORIAL HERPETOFAUNA 193 forest in April; only shells remained. Conceivably these individuals were seek- ing oviposition sites when they were at- tacked by predators. The proximity of the turtles to the large permanent pond suggests that this species may inhabit such ponds and lakes. Phrynops geoffroanus tuberosus (Peters) Platcmtis tiihcrosa Peters, 1870, Sber. Akad. Wiss. Berlin, 1870:311 [Type locality.— Cotinga River, Mt. Roraima, Guyana]. Phninops pcoffroana tiihcrosa — Miillcr, 1939, Physis, 16:95. Material. — Puerto Ore, 1; Santa Ce- cilia, 1. Identification. — Plirynops ^cojfroanus has a broad, smooth carapace that is flared peripherally. The carapace has a deep angular indentation in the posterior margin. The carapace is black. The plastron and ventral flange of the cara- pace are creamy yellow to orange with black spots. The limbs are dull gray with pale red to creamy white spots. The head and neck are dull olive green with fine black reticulations and a black line from the snout through the orbit and onto the neck. The throat is pale tan with irregular black markings. A pair of long creamy white chin barbels are present. The only other side-necked tin-- tle in the area with a smooth carapace is Mesoclemmys gihha which has a black plastron with a shallow rounded indenta- tion in the posterior margin. 9 316 mm. Occurrence. — Both specimens were obtained from the Rio Aguarico by na- tives. Life Histori/. — No data are available from Santa Cecilia. Medem (1969:334) reported nests containing 10, 11, and 15 e22:s found in March and November in Caqueta, Colombia. Platemys platycephala (Schneider) Testiido platycephala Schneider, 1792, Schr. Ges. Naturf. Freunde Berlin, 10:261 [Type locality. — "East Indies"]. Platcntys platycephala — Boulenger, 1889, Cat. Chelon. Rhynchoceph. Crocod. Brit. Mus.: 227. Material. — Santa Cecilia, 8. Identification. — This brightly colored turtle is readily distinguished from other side-necked turtles in the area by its elongate carapace having two rounded longitudinal ridges, bordering a median depression. The shells of MesocJemmijs and Plirynops lack ridges, and CheJiis has three keeled ridges. The carapace, plastron, and skin are dark brownish black with an orange-tan periphery to the shell and broad orange stripes on the limbs and dorsum of the head, c? 161 mm; 9 156 mm. Occurrence. — A juvenile having a carapace length of 55 mm was in a patch of sunlight in a shallow rivulet in pri- mary forest, and one adult was crawling along a trail by day. One adult was in shallow water in the Rio Aguarico by day, and two adults were lying on the bottom of shallow swamps at night. Life History. — No data are available. KiNOSTERNIDAE Kinosternon scorpioides (Linnaeus) Testudo scorpioides Linnaeus, 1766, Systema Naturae, Ed. 12, 1:352 [Type locality.— Surinam]. Kinosternon scorpioides — Gray, 1831, Synops. Rept., 1:34. Cinosternon scorpioides scorpioides — Sieben- rock, 1907. Sher. Akad. Wiss. Wien, 116: 576. Material. — Santa Cecilia, 9. Identification. — This small turtle with a high carapace with three low, longitu- dinal keels withdraws the neck into the shell. Other species with longitudinal ridges or keels on the carapace ( CheJus and Platemys) have low, flattened shells, and fold the neck laterally under the margin of the shell. The other turtle with a high carapace is Geochelone, which has short limbs with unwebbed toes; the toes of Kinosternon are webbed. The carapace is dull grayish brown; the plastron is dull yellowish brown. The top of the head is yellowish orange with black flecks; the rest of the head, neck, and limbs are dull gray. 6 161 mm; 9 155 mm. 194 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Occurrence. — One individual was in shallow muddy water in swamp forest, one was in a pond in forest, and one was in a shallow muddy stream in primary forest. Life History. — No data are available. Remarks. — Three specimens were found in 1966, five in 1967, and one in 1975. For some inexplicable reason no others were found during the extensive field work in other years. Testudinidae Geochelone denticulata (Linnaeus) Testudo denticulata Linnaeus, 1766, Systema Naturae, Ed. 12, 1:352 [Type locality.— "Virginia"]. Geochelone denticulata — Williams, 1960, Brev- iora, 120:10. Material. — Lago Agrio, 4; Santa Ce- cilia, 3. Identification. — This large tortoise with a domed shell and short, stubby legs is unlike any other turtle in the area. The carapace is dull brown and tan; the plas- tron is tan. The limbs are brown with orange-tan scales, and the neck is dark brown. The head is yellowish to orange- tan with brown spots in the occipital region. Males have a concave plastron, whereas the plastron in females is nearly flat. 6 378 mm; $ 328 mm. Occurrence. — This is a diurnal forest dweller. One was sitting at the base of a stilt palm; two were walking on the for- est floor, and one was moving along the edge of the forest in late afternoon. Life History. — No data are available. Remarks. — Geochelone is eaten by local human inhabitants. On the basis of the number of shells found around Indians' houses, the tortoise must be common in the area. However, with the recent increase in the human population, the number of tortoises can be expected to dwindle rapidly. CROCODILIANS CROCODYLroAE Caiman crocodilus crocodilus (Linnaeus) Lacerta crocodilus Linnaeus, 1758, Systema Naturae, Ed. 10, 1:200 [Type locality.— Guyana (restricted by Mertens and Wer- muth, 1955)]. Caiman crocodilus crocodilus — Miiller and Hell- mich, 1936, Wiss. Ergebn. Deutsch. Gran- Chaco Exped., Amph. Rept.:110. Material. — Santa Cecilia, 8. Identification. — Caiman is readily dis- tinguished from the other crocodilian in the area, Paleosuchus palpebrosus, by having a bony transverse preorbital ridge and two or three transverse rows of oc- cipital scales. Paleosuchus lacks a pre- orbital ridge and has one transverse row of greatly enlarged, strongly keeled oc- cipital scales. Caiman has a dull brown or olive-brown dorsum with black trans- verse marks on the dorsum of the body and sides of the tail. The flanks are yel- lowish brown, and the lips are olive-tan. The venter is creamy white, and the iris is greenish gray. 6 550+350+ mm (in- complete); 9 943+1009 mm (extrapo- lated from skull ) . Occurrence. — All individuals were found at night. One juvenile was in the backwater of the Rio Aguarico. One adult was in a forest swamp, and two adults and three juveniles were in tempo- rary marshes in clearings. One juvenile came from a lake where many adults were observed at night. Life History. — No data are available from Santa Cecilia. Medem (1969:330) reported on a nest of this species on the Rio Caqueta (Amazon drainage) in Co- lombia. The nest, containing 24 eggs, was found in 16 March. Hatchlings had snout- vent lengths of 115.5-123.0 mm. These are noticeably smaller than the smallest juveniles from Santa Cecilia (230, 275, 285 mm). Of these, the one having a snout-vent length of 230 mm was found on 16 March. The other two are from April. In light of Medem 's data it is probable that Caiman has an exten- sive reproductive season. Remarks. — Hunting pressure and wanton killing of Caiman by the increas- ing human population seems to have de- creased the population of Caiman in the EQUATORIAL HERPETOFAUNA 195 lakes at Santa Cecilia. In 1967 scores of Caiman (as determined by their red eye- sliine) were observed on various nights. In 1971 and 1972 only a few could be seen on any given night. Paleosuchus trigonatus (Schneider) Crocodihis trigonatus Schneider, 1801, Hist. Amph., 2:161 [Type locality. — unknown]. Paleosuchus trigonatus — Schmidt, Field Mus. Nat. Hist. Zool., 12:209. Material — Lago Agrio, 3; Santa Ce- cilia, 5. Identification. — Paleosuchus is read- ily distinguished from the only other crocodilian in the area. Caiman croco- dilus, by lacking a bony transverse pre- orbital ridge and by having a single transverse row of greatly enlarged, strongly keeled occipital scales. Caiman has a preorbital ridge and two or three transverse rows of small occipital scales. Paleosuchus is dull brown dorsally with broad, dark brown and orange-tan bars on the tail. The venter and lower jaw are cream with large, brownish-gray spots. The iris is brown. S 465+410 mm; 9 837+732 mm (extrapolated from skull). Occurrence. — All were found at night. One juvenile was in a muddy depression in second growth forest near the river; two were in small streams, and one was in the Rio Aguarico. Ecological data are lacking for the other specimens. Life History. — No data are available. Remarks. — Seemingly the two croco- dilians in the area have diflFerent habitat preferences. Caiman inhabits lakes and swamps, whereas Paleosuchus primarily inhabits streams and rivers with a strong current. These observations corroborate those of Medem (1952) on Paleosuchus trigonatus in Colombia. The large black caiman, Melanosuchus niger, apparently does not occur in the upper Rio Aguarico. According to Medem (1963), it inhabits slowly flowing rivers. Medem reported it from the Rio Putimiayo in Colombia, and there are specimens at The Univer- sity of Kansas from the Rio Napo in Ecuador. LIZARDS Gekkonidae Gonatodes concinnatus (O'Shaughnessy) (Figure 114) Material. — Lago Agrio, 5; Santa Ce- cilia, 27. Identification. — This small gecko has claws that are not covered by a scaly sheath, and the scales on the belly are broadly rounded posteriorly. In these respects it differs from the other small gecko in the area, Pseudogonatodes guianensis in which the claws are retrac- tile into a scaly .sheath, and the scales on the belly are pointed posteriorly. Female Gonatodes can be confused with small Anolis. In the latter, scaly, moveable eyelids are present, the digits are di- lated, and the scales on the top of the head are not granular. Furthemiore, Anolis have much longer limbs and tails. The scales on the dorsal and lateral surfaces of the body are small and gran- ular; the ventral scales are noticeably larger, ovoid, and flat. Striking sexual dimorphism exists in coloration. In both sexes a white collar is evident and usu- ally interrupted dorsally; the collar is narrow in females and much wider, es- pecially ventrolaterally, in males. The head and neck are orange or reddish brown with cream spots in males; the body is olive green with reddish brown reticulations, and the tail is black with white rings. The throat is orange with cream streaks, and the belly is yellowish gray. In females the head, body, and tail are grayish tan with irregular brown markings, some of which are crescent- shaped in a pair of dorsolateral rows; the spots on the tail tend to be fused across, giving the tail a banded appear- ance. The venter is creamy tan. The iris is reddish brov/n with a cream circle around the pupil. Juveniles are colored like females. Individuals having regen- erated tails lack bands or rings on the tail. >-, :•** Fig. 114. — Gontodes concinnatus. Upper: $, 41 mm SVL (KU 146643). Lower: $, 48 mm SVL (KU 146645). Occurrence. — By day, most individ- uals were found on trunks of large trees in primary forest; one was found in leaf litter on the forest floor. All were in shade. At night, individuals were sleep- ing horizontally on leaves or stems of bushes and vines; usually adjacent to tree trunks. One was on a log in a clear- ing at night. Life History. — Six females ( March 1, April 3, May 1, August 1) each contained one egg; the smallest of these females had a snout-vent length of 34 mm. The largest oviducal egg was 10.4 mm in length. The two smallest juveniles (snout- vent lengths of 17 and 19 mm) were obtained in July. These limited data, together with those summarized by Fitch (1970:20), suggest that Gonatodes probably reproduces throughout the year with the possible exception of the drier months ( December-February ) . Food. — A variety of small arthropods was eaten. In both frequency and vol- ume, five items arc about equal ( spiders, beetle larvae, termites, roaches, and he- mipterans). These comprise 61.3 percent of the frequency and 56 percent of the volume in 10 stomachs. Pseudogonatodes guianensis Parker (Figure 115) Material. — Puerto Libre, 6; Santa Ce- cilia, 11. Identification. — This minute lizard, EQUATORIAL HERPETOFAUNA 197 Fig. 115. — Pseudogonatodes guianensis, 2, 26 mm SVL (KU 146646). the smallest species in the area, has claws that are retractile into a scaly sheath; it differs in this respect from the slightly larger gecko, Gonatodes concinnatus, in which the claws are not covered by a scaly sheath. Furthermore, in Pseudo- gonatodes the posterior margins of the belly scales are pointed, whereas they are broadly rounded in Gonatodes. Both Gonatodes and Pseudogonatodes lack moveable lower eyelids, a characteristic shared with a third gecko, Thecadactylus rapicauda, which has greatly dilated toes and large tubercles interspersed among the granular dorsal scales. Gonatodes and Pseudogonatodes have uniform gran- ular scales on the dorsum and cylindrical or laterally compressed toes. The dorsum is dull brown. Usually the pattern consists of an orange-tan trans- verse band on the neck and posterior part of the head and small orange-tan spots in two dorsolateral rows. In some individuals a yellowish-tan dorsolateral stripe is evident on the proximal half of the tail. Some individuals are darker brown or gray and have an obscured pat- tern. Usually two cream bars are present on the lip. The chin and throat are white with brown or gray longitudinal streaks, and the belly is tan or gray. The iris is orange-tan. S 26-]-21 mm; 9 27+23 mm. Occurrence. — This small lizard is ac- tive amidst shaded leaf litter on the for- est floor by day. Nine of 13 individuals were in primary forest; two were in sec- ondary forest, and two were at the edge of the forest. Life History. — One female from April contained one egg 5 mm in length, and one from July contained one egg 6 mm in length. All other females were ob- tained in May, June, and July; five con- tained small ovarian eggs, and one showed no evidence of reproductive ac- tivity. Food. — In 11 stomachs five food items comprised 93.3 percent of frequency and 90.9 percent of volume - spiders, sow bugs, beetle larvae, termites, and small orthopterans. Taxonomic Comments. — The morpho- logical characters of the Ecuadorian specimens support Hoogmoed's (1973: 96) contention that Pseudogonatodes amazonicus Vanzolini is indistinguish- able from P. guianensis. Thecadactylus rapicauda (Houttuyn) (Figure 116) Material. — Lago Agrio, 4; Santa Ce- cilia, 9. Identification. — This large, robust gecko has two rows of greatly expanded pads on the underside of each toe; the dorsum of the body is covered with mi- nute granular scales with larger tuber- cles interspersed. These characters serve to distinguish Thecadactylus from all other lizards in the area. The dorsum is tan, orange-tan, or 198 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY c£r» Fig. 116. — Thecadactijlus rapicauda, $, 53 mm SVL; Barro Colorado Island, Canal Zone (KU 75768). grayish tan with irregular dark brown or black lines on the head and neck and narrow dark markings on the body. The latter markings vary from short dashes to rhomboidal marks enclosing pale areas of the ground color. A quadrangular or diamond-shaped black or brown spot usually is present at the base of the tail, and two or three such markings are present on original tails. The flanks are dark brown or colored like the dorsum; if the latter, a dark brown ventrolateral stripe extends from the eye to the base of the hind limb. The venter is pale yel- lowish tan; the iris is reddish tan. $ 108+91 mm; 9 112+84 mm. Occurrence. — This is the only noc- turnal lizard in the area. One individual was found in a thatched roof at dusk, one was on a box next to a building at night, and others were observed on banana plants at night. Four were obtained from trees when primary forest was being cleared. Life History. — Four females each contained one oviducal egg 18-22 (X^ 19.0) mm in length; the females were ob- tained in May (2), June, and Augu.st. Food. — In seven stomachs, roaches comprised 70 percent of the volume. One individual contained a scorpion. Iguanidak Anolis chrysolepis scypheus Cope ( Plati-; 3 ) Anolis scypJwus Cope, 1864, Proc. Aead. Nat. Sei. Philadelphia 1864:172 [Type locality.— not given]. Anolis cJt)ysoIe))is scyplieus — Vanzolini and Williams, 1970, Arq". Zool., Sao Paulo 19:85. Material. — Dureno, 2; Lago Agrio, 6; Puerto Ore, 1; Santa Cecilia, 167. Identification. — This moderately large anole has a robust body and proportion- ately, a large head with a short snout. The dorsum is predominately brown, and the dewlap is red peripherally and blue medially with white scales. No other anole in the area has a similarly colored dewlap. Females can be distinguished from other anoles by the large head, small keeled supraorbital scales, large keeled ventral scales, and the presence of a small dewlap. Anolis ortonii is the species most easily confused with Anolis chrysolepis; the former has larger smooth supraorbital scales and smooth ventral scales. Furthermore, the dewlap in Ano- lis ortonii is orange with red streaks. The dorsum is variegated brown, red- dish brown, tan, and/or gray. Usually in males and occasionally in females, there is a narrow, yellowish tan, longi- tudinal nu'ddorsal stripe on the posterior part of the body and base of the tail. The dorsal pattern in males usually con- sists of dark brown or reddish brown diagonal marks or semicircles (closed edge dorsally). The top of the head is brown with a dark brown interorbital bar. Most females have a broad middor- EQUATORIAL HERPETOFAUNA 199 sal gray or tan stripe bordered by a broad brown area. The venter is pinkish cream in females and rusty red or burnt orange in adult males. The dewlap is blue medially with white scales and a broad red periphery. The iris is tan. S 84+182 mm; $ 86+170 mm. Occurrence. — This primarily terres- trial anole inhabits lioth primary and secondary forest. Although a few indi- viduals have been observed basking on logs or buttresses, most have been seen actively foraging in the shade. The ma- jority of the lizards were found sleeping at night. Usually the lizards were hori- zontal on a branch of a bush less than 1.5 m above the ground, and some were found head up on sticks or stems close to the ground. Only two were found on branches more than 1.5 m above the ground. Although Anolis chrysolepis climbs on logs or buttresses of trees during its nonnal diurnal activity, it is primarily a terrestrial species. Individuals run short distances on the ground and then remain motionless; their coloration blends well with the leaf litter on the forest floor. Fitch (1968:36) reported body tempera- tures of eight specimens as 24.0-32.8°C (Xm27.5±0.88) with a mean deviation from air temperature of +1.51±0.77°C; he discussed these lizards under the name Anolis scypheus. Life History. — Gravid females have been found from March through Novem- ber; only three specimens of the species were obtained from December through February. Six females each contained one oviducal egg, and seven contained two eggs, length 15.4-19.6 (X=:17.1, N= 20) mm. One individual having a snout-vent length of 29 mm hatched on 31 October from an egg found on 22 September. Juveniles having snout-vent lengths of 28-31 mm were found from March through October. These data and those on gravid females strongly suggest that reproduction occurs throughout the \'ear. Food. — In 25 specimens examined, orthopterans comprised 28 percent of the volume of food, and beetle larvae, 11.2 percent. The other seven categories of food were found in small amounts. One was observed eating a microteiid lizard, Leposoma parietale, and one contained the tail of an Anolis trachyderma. Anolis fuscoauratus fuscoauratus D'Orbigny (Plate 3) Material. — Lago Agrio, 27; Puerto Libre, 15; Puerto Ore, 3; Santa Cecilia, 163. Identification. — This small anole is identifiable chiefly by the absence of any one distinguishing character. All of the head scales are small and keeled; the ventrals are small, smooth, and nearly granular. The dorsum is gray or dull tan; many females have a pale tan middorsal stripe. The chin, throat, and belly are grayish white with brown or dark gray flecks. In males the large dewlap is rose- pink with white scales and a white mar- gin. The iris is brown. Anolis fuscoaura- tus is most easily confused with A. trachy- derma and A. ortonii. The former has larger, flat, smooth scales ventrally and a transverse yellow line on the chin; the small dewlap is orange-red with black scales. Anolis ortonii has flat, smooth ventral scales and smooth head scales. The dorsum of A. ortonii is tan or pale gray with brown mottling; the large dew- lap is orange with red streaks. Juveniles of A. punctatus can be distinguished from A. fuscoauratus by the former hav- ing keeled ventral scales and keeled scales on the tip of the snout, whereas the other head scales are smooth. S 43+85 mm; 9 47+92 mm. Occurrence. — Although this small liz- ard frequently is seen on the forest floor by day, individuals are most easily col- lected at night when they are sleeping horizontally or head-down on grasses, herbs, or limbs of bushes, usually at heights of less than 1 m from the ground. During the day, Anolis fuscoauratus for- 200 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ages in shade on the forest floor; only four were observed sunning - three on logs and one on a low bush. The local distribution of the species is most accu- rately determined by the occurrence of sleeping individuals, most of which are associated with forest - primary forest 26 percent, secondary forest 52 percent, and forest edge 15 percent. Three individuals were found in clearings, and nine in banana groves. Individuals were found throughout the year, but only four were taken in the drier months (December-February). Life History. — The smallest females with mature ovarian eggs have snout- vent lengths of 37 mm or more and have been found in every month except Janu- ary and February. No females were found in January, and the only one taken in February had immature ovarian eggs. More than 40 percent of the females taken in each of five months ( March- July) contained oviducal eggs. Of 52 fe- males containing oviducal eggs, 33 had one egg, and 18 had two eggs. Of 36 females containing large ovarian eggs, 30 had one egg, and six had two eggs. Four females each laid a single egg in captivity. Eggs were deposited on 2 March, 4 October, and 14 and 21 Decem- ber. They required 53-63 (x=57) days to hatch. The eggs varied in size from 9x5 mm to 9.5 X 6 mm. The hatchlings had snout-vent lengths of 16-19 (x= 17.4) mm. Apparently, Anolis fuscoauratus usu- ally deposits single eggs throughout the year. The presence of two equally well- developed eggs in the reproductive tract at a given time indicates that clutches of two may be deposited or that the eggs are laid separately at short intervals. Food. — In 25 stomachs, small spiders and orthopterans comprised half of the volume of food. Ants made up 11 per- cent of the volume, whereas other kinds of food were present in small quantities. Remarks. — The ecological relation- ships of Anolis fuscoauratus and A. trachydcrma are discussed in the account of the latter species. Anolis ortonii Cope ( Plate 3 ) Material. — Lago Agrio, 3; Santa Ce- cilia, 9. Identification. — This small tan or pale gray anole has large smooth ventral scales; the supraorbitals are large and smooth. The scales on the tip of the snout are keeled, but the other head scales are smooth. Both Anolis fusco- auratus and A. trachyderma have keeled dorsal head scales and small keeled su- praoculars; the ventral scales in these species are smaller than those in A. or- tonii. The large dewlap in A. ortonii is orange with red streaks, whereas the dewlap in A. fuscoauratus is large and rose-pink with white scales, and that in A. trachyderma is orange-red with black scales. The dorsal coloration of A. or- tonii is tan or pale gray with brown streaks on the body and a dark brown interorbital bar. The throat is brown with dark brown diagonal bars evident in most adult males. The venter is creamy tan, and the iris is brown. $ 50+64 mm; 9 43+68 mm. Occurrence. — Half of the 12 speci- mens were obtained in June and July; others were found in March ( 1 ) , April (2), May (2), and November (1). Two individuals were found on a thatched roof in a clearing. All others were in for- est (five in primary, two in secondary). One was on a tree trunk, one on the branch of a low bush, and three were on the ground. Two were sleeping on low bushes at night. These limited observa- tions suggest that A. ortonii is primarily terrestrial but does ascend tree trunks and low bushes during its period of ac- tivitv. Life History. — Two females (one from May and one from July) each con- tained two oviducal eggs 10.5 and 10.3 mm in length. Two females from June and July contained large ovarian eggs. No juveniles were found. Food. — Ants formed the most numer- ous food item in seven stomachs and comprised 53.6 percent of the volume. Other food items included spiders and EQUATORIAL HERPETOFAUNA 201 orthopterans, including one mole cricket ( Gryllotalpidae ) . Anolis punctatus boulengeri O'Shaughnessy ( Plate 3 ) Material. — Lago Agrio, 7; Santa Ce- cilia, 10. Identification. — The green dorsum and yellow scales around the eye imme- diately distinguish this large anole from its sympatric congeners. The snout is long, and the scales on the tip of the snout are keeled; the other head scales are smooth, and the supraorbitals are enlarged. The ventral scales are weakly keeled. The dewlap is large and yellow or orange-yellow with pale yellow or white scales. The only other large anole in the area is Anolis transversalis, which has smooth head scales and ventrals and a large orange or pale yellow dewlap with black or dark brown streaks or bars; furthermore, the dorsum of A. transver- salis has diagonal black or dark brown marks. The dorsum of Anolis punctatus is bright green with pale cream or bluish white flecks laterally and dorsolaterally and tan middorsally. The throat and scales around the eye are yellow; the tail is tan distally. The belly and ventral surfaces of the limbs are creamy white to pale tan with brown flecks. The iris is bronze. S 82-f 170 mm; 9 77+202 mm. Occurrence. — Individuals were found in all months, except November through January; seven were obtained in May from trees as they were felled at Lago Agrio. Three individuals were sleeping on bushes at heights of less than 1.5 m above the ground, and one juvenile was sunning on a log. All were found in forest. Life History. — All 11 females con- tained eggs; two females taken in May each had one oviducal egg ( 17.1 and 18.1 mm in length), and one found in August had two oviducal eggs 16.2 mm in length. Females having matru-e ovarian eggs were found in July and October, and females from February through July con- tained immature ovarian eggs. Thus, the limited evidence suggests reproductive activity throughout the year. The only juvenile has a snout-vent length of 49 mm and was obtained in September. Food. — In 14 stomachs, three groups of insects comprised 76.8 percent of the volume of food: orthopterans (30.4%), ants (28.6%), and beetles (17.8%). AnoHs trachyderma Cope ( Plate 3 ) Material. — Lago Agrio, 3; Puerto Libre, 9; Santa Cecilia, 308. Identification. — This small, drab liz- ard is darker colored by day than any other anole in the area. It can be dis- tinguished from the other species by its dark chin with one or two narrow, trans- verse cream or yellow lines. The dorsal head scales are keeled, and the ventral scales are small, flat, and smooth. In males, the base of the tail is greatly swollen, and the dewlap is small and orange-red with black scales. Anolis fuscoauratus and A. ortonii are about the same size, but both lack transverse light lines on the chin; the base of the tail is not swollen in males, and the dewlap is differently colored - rose pink with white scales in A. fuscoauratus and orange with red streaks in A. ortonii. At night Anolis trachyderma is pale tan with little or no evidence of pattern. By day, it is brown with dark brown markings: interorbital bar, two diagonal bars on lips, stripe extending posteriorly from eye, and in some males small chev- rons middorsally on body and base of tail. Narrow, cream diagonal lines are present on the forearm and shank. In many females a reddish brown middorsal stripe is bordered by a narrow, irregular tan line; in some females small yellowish tan vertical bars are evident dorsolater- ally on either side of the middorsal stripe. The brown dorsal color extends onto the ventrolateral surfaces of the body. The belly is cream or yellow, usually flecked with brown. The iris is reddish brown. $ 51+106 mm; 5 58+89 mm. b 202 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Occurrence. — This small anole is ac- tive on the forest floor and in low vege- tation. Approximately twice as many in- dividuals were observed on the ground and in leaf litter as were seen in bushes. None was observed in direct sunlight, nor was the species found in clearings or cultivated areas. Of 265 individuals found sleeping, usually head up or hori- zontally on low vegetation at night, 52 were in primary forest, 195 were in sec- ondary forest, and 18 were in forest-edge situations. Life History. — Reproduction appar- ently takes place throughout the year. No females were obtained in January; two were collected in December, and both contained immature ovarian eggs. Females containing oviducal eggs were found in all of the other months. No females having snout-vent lengths of less than 45 mm contained large ovarian or oviducal eggs, although females having snout-vent lengths of 40-45 mm con- tained small ovarian eggs. Each of eight females contained two oviducal eggs, and 36 had one egg each. Lengths of oviducal eggs were 12.4-15.4 (x=13.6, N=52) mm. Three captive females laid single eggs on 7 February, 15 September, and 22 November; the latter hatched 77 days later on 7 February. One egg found in leaf litter on 19 July hatched on 10 August, and one found in leaf litter on 22 September hatched on 6 November. The eggs varied in size from 10 X 3.4 mm to 14 X 6 mm. The hatchlings had snout- vent lengths of 22.0-23.0 mm. Six- teen recent hatchlings having snout-vent lengths of 23-29 mm were found in March (2), May (l),June (5), July (4), August ( 1 ) , September ( 1 ) , and October (2). The dates for hatchlings seem to reinforce the data on oviducal eggs in providing evidence for acyclic reproduc- tion in this species, but no recent hatch- lings were found in the drier months ( December-February ) . Food. — In 27 stomachs, spiders com- prised 42.2 percent of the volume; nine Table 6. — Comparative Data on Abundance of AnoJis fuscoauratiis and Anolis trachijderma at Santa Cecilia. A. jus- A. trachij- coauratus derma (iV=164) (N=283) Habitat Percentage Occurrence Primary Forest 26% 20% Secondary Forest -- 52% 73% Forest Edge 15% 7% Clearing - 7% 0% Date Frequency of Occurrence March 1967 17 14 June 1967 19 7 June 1968 22 35 August 1971 2 35 October 1971 8 19 Other kinds of arthopods were present in much smaller amounts. Remarks. — In comparison with Anolis fuscoauratiis, A. trachijderma seems to be more restricted in its ecological dis- tribution and a less frequent inhabitant of primary forest. This conclusion is borne out by three sets of data. The ma- jority (73%) of A. trachijderma was taken in secondary forest, whereas only 52 per- cent of the A. fuscoauratiis was found there. Anolis fuscoauratiis was found in clearings, but no A. trachijderma was found there (Table 6). At Lago Agrio when primary forest was being cleared in May 1969, all anoles seen were cap- tured - 27 A. fuscoauratus and three A. trachijderma. In early collections from Santa Cecilia, when most of the forest was undisturbed ( 1967-68 ) , the two spe- cies were found in nearly the same fre- quency, whereas in later collections ( 1971 ) , when much of the primary for- est had been partially lumbered, A. trachijderma was found in far greater abundance than A. fuscoauratus (Table 6). Thus, it seems evident that although A. fuscoauratus has broader ecological tolerances than A. trachijderma, the lat- ter species is increasing in relative abun- dance with the increase in the amount of secondary forest. Anolis transversalis Dumeril (Plate 3) EQUATORIAL HERPETOFAUNA 203 Fig. m .—Emjalioides cofanorum, $, 107 mm SVL (KU 146658). Material. — Lago Agrio, 3. Identification. — This large anole is readily distinguished from all other anoles in the area by having smooth head scales and a distinctive color pattern of transverse markings on the body. The dorsal coloration changes from dark brown to black. The dark transverse markings are diagonal posteroventrally onto the edge of the belly and vary from four broad to six or seven narrow marks on the body and about an equal number on the tail. A dark vertical bar extends ventrally from the eye, and another from the ear. The throat and belly are cream, and the dewlap is yellow with bold black or dark brown vertical bars or narrow gray or bluish gray diagonal lines. Dew- laps are present in both sexes. The iris is bright blue, i 72+104 mm; 9 72+100 mm. Occurrence. — Three specimens were obtained from high branches of large trees as they were felled. Presumably the species lives only in the foliage of the upper story of primary forest. Life History. — The single female ob- tained in May contained one ovarian egg 14.4 mm in length. Food. — All three specimens contained only beetles. Enyalioides cofanorum Duellman (Figure 117) Emjalioides cofanorum Duellman, 1973, Herpe- tologica 29:230 [Type locality. — Santa Ce- cilia, Provincia Napo, Ecuador]. Material. — Santa Cecilia, 11. Identification. — Emjalioides cofano- rum is a moderately large lizard with a large, chunky head, a middorsal crest of enlarged spiny scales, and keeled spinous caudal scales arranged in whorls. The species resembles Enyalioides laticeps festae but differs in that the latter has homogeneous scales on the dorsolateral surfaces of the body; E. cofanorum has a dorsolateral row of distinctly enlarged, conical scales on the body. Plica umbra ochrocollaris superficially resembles both species of Enyalioides, but it is a smaller, more slender lizard with larger scales on the bodv, smaller caudal scales not an- 204 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 118.— Entjalioides laticeps festae, $, 114 mm SVL (KU 122106). ranged in whorls, and a blue lining of the mouth (pink in Enyalioides) . Adults are dark brown with scattered greenish yellow scales on the back and lips. The throat is brown with scattered cream settles, and the belly is tan. Ju- veniles are dark brown with yellowish tan cross bars and black spots along the dorsal crest. There is a yellow diagonal bar from the ear to the shoulder; the head is rusty brown. The chin is grayish white, and the belly is cream. The iris is orange-brown. £ 101+ 118 mm; ? 107 + 120 mm. Occurrence. — Five of nine individ- uals were found in primary forest; the others were in secondary forest. Two were active by day on the forest floor (one was on a log). By night individuals were found sleeping head-up on sticks or horizontally on limbs of bushes, all less than 1.5 m above the ground. Life Histonj. — Of six adult females, two were found in July and one each in March, June, August, and September. One female taken in March contained two ovarian eggs 26.3 mm in length; one taken in July had two oviducal eggs 28.4 mm in length. The smallest juveniles, having snout-vent lengths of 55 and 57 mm, were taken in April and June, re- spectively. Food. — Six individuals contained five kinds of food in nearly the same vol- umes: earthwoiTns, sow bugs, spiders, beetles, and orthopterans. Remarks. — Comparative ecological data on the two species of Enyalioides are given in the following account. Enyalioides laticeps festae Peracca (Figure 118) Material. — Dureno, 1; Lago Agrio, 11; Puerto Libre, 1; Santa Cecilia, 46. Identification. — This moderately large lizard has a chunky head and a middor- sal crest of large spiny scales; the keeled, EQUATORIAL HERPETOFAUNA 205 spinous scales of the tail are arranged in whorls. Enyalioides laticeps festae is most easily confused with two other iguanids of about the same size - En- yalioides cofanorum and Flica umhra ochrocollaris. The latter has a propor- tionately smaller head and weakly keeled caudal scales not arranged in whorls; furthermore, the lining of the mouth is blue in Plica and pink in Enyalioides. The two species of Enyalioides are easily distinguished by the natine of the scales on the body. In Enyalioides laticeps fes- tae the dorsal scales, except the middor- sal row, are of uniform size; in E. co- fanorum there is a dorsolateral row of distinctly enlarged, conical scales. Some preserved specimens of E. laticeps festae appear to have a dorsolateral row of en- larged scales, but in realit\^ this appear- ance is due to a fold of skin, an artifact of preservation. In adults the dorsum varies from dull green to brown with paler, irregular spots dorsolaterally, faintly expanded in- to diagonal bars in some specimens. Usu- ally the top of the head is dull olive green, and the rest of the dorsum is dull olive green with tan or pale orange spots or diagonal bars. The lips are green, and blue or blue-green scales are present ventrolaterally on the body and on the limbs. The chin is pale green. In males the throat is streaked orange, brown, and green; the gular region is dark brown, and the belly is orange. In females the gular region is rusty red, and the belly is pinkish tan. In many specimens there is a pale orange or tan diagonal blotch on the side of the neck. Most adult males have an orange stripe from the corner of the mouth to a point below the ear. The iris is brown with a yellow or greenish yellow ring around the pupil. Juveniles are pale green or tan with brown or red- dish brown diagonal markings on the body; the throat is white (with red streaks in some), and the belly is creamy tan. S 128+200 mm; $ 125+202 mm. Occurrence. — Enyalioides laticeps festae is a diurnal inhabitant of primary forest; 80 percent of the individuals were found in primary forest, whereas the others were in secondary forest. One was observed on the ground by day; 17 others were on vegetation. By day, the species usually is observed on trunks of small trees or palms having a diameter of less than 15 cm; one was found on a bamboo. Especially in the mornings, the lizards sun head-up on the trunks; when dis- turbed they move around and up the trunk. At night the lizards sleep, usually nearly horizontally, on branches or palm fronds, usually at heights of more than 1.5 m above the ground. Some sleep head-up on trunks of saplings. Life History. — No females were found in October through February; females with immature ovarian eggs were found in March through September. One fe- male with mature ovarian eggs was found in March, and four were found in May. These five females have snout-vent lengths of 107-125 (x=114) mm. The number of eggs in each varied from 5 to 7 (x=6.2), and lengths of the eggs were 15.0-16.6 (x=15.6) mm. Adult males were found in March through November. The three smallest juveniles have snout- vent lengths of 43, 45, and 57 mm and were found in March, April, and March, respectively. The reproductive data are inconclu- sive with regard to seasonality. The ab- sence of reproductively active females from October through February prob- ably is due to inadequate sampling, be- cause the smallest juveniles were taken in March and April. Presumably, the species breeds throughout most or all of the year. Food. — In 24 stomachs, three food items comprised 70.4 percent of the vol- ume: spiders (37.5^), caterpillars (20.4%), and beetle grubs (12.5%). The other 29.6 percent was made up of eight different groups of invertebrates, of which orthopterans comprised 8.2 per- cent. Remarks. — Enyalioides cofanorum and E. laticeps festae principally inhabit pri- 206 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 4 w^^; '^t < '■■ >^^^^w<^ Fig. 119.— PZfca umbra ochrocollaris, $ , 89 mm SVL ( KU 146659). mary forest; the former species appar- ently is terrestrial, whereas the latter seems to be primarily arboreal. Thus, the two species seem to be ecologically segregated. Plica umbra ochrocollaris (Spix) (Figure 119) Material. — Lago Agrio, 5; Santa Ce- cilia, 27. Identification. — This lizard is imme- diately distinguishable from all others in the area by having a middorsal crest and large, pointed, keeled scales of unifomi size on the body and tail; the scales on the tail are not arranged in transverse whorls. Plica is most easily confused with EnyaJioides, the only other lizards in the area having middorsal crests, but Enyalioides has small granular scales on the body, caudal scales arranged in trans- verse whorls, a large chunky head, and the lining of the mouth pink. Plica has a proportionately smaller head and the lin- ing of the mouth blue. The dorsal coloration is highly vari- able. Some males are bright green with small black spots in a dorsolateral row; others are dull olive-green or olive-tan, as are females. The tail is olive-tan above and reddish tan below. In all individuals a black mark is present below the eye and another is present at the angle of the jaw. A narrow black line extends pos- teriorly from the eye and connects (or nearly so) with a narrow black collar that is incomplete middorsally. The area below the line is creamy white to reddish brown. The throat is grayish brown, reddish brown, or greenish gray, and the belly is pale grayish brown to reddish brown. The iris is orange or reddish tan. S 89+204 mm; $ 90+181 mm. Occurrence. — All individuals were as- sociated with forest ( 12 in primary, 18 in secondary). One was sunning on a log at the edge of the forest, and five were obtained as trees were felled in primary forest. Twenty individuals were sleeping at night horizontally no bran- ches, palm fronds, or bamboo twigs; one was head-up on a tree trunk. Most sleep- ing individuals were at heights of less than 1.5 m above the ground, but five were at greater heights, one at 4 m. Nine of 20 sleeping individuals were on perches over swamps or streams. Life History. — Three females ( March 2, August 1) each contained two ovi- ducal eggs 23.0-26.7 (x=24.6) mm in EQUATORIAL HERPETOFAUNA 207 Fig. 120.— Polychrus marmoratus, 2, 148 mm SVL (KU 142870). length. One from May contained four maturing ovarian eggs 16.5 mm in length. All other females were taken in April through November and all contained small ovarian eggs. The smallest of these lizards has a snout-vent length of 69 mm. The smallest juvenile has a snout-vent length of 45 mm and was obtained in August. Beebe (1944:208) noted that in Guyana Plica nmhra was in breeding condition in April, May, and October. Food. — Eleven individuals contained only ants in their stomachs. Beebe ( 1944: 208) reported beetles, orthopterans, and centipedes, in addition to ants, in lizards from Guyana. Polychrus marmoratus (Linnaeus) (Figure 120) Material. — Dureno, 2; Santa Cecilia, 7. Identification. — This large, green (brown in preservative) lizard with white and yellow markings and a long tail is easily recognized. The toes are not dilated, and the third and fourth toes are equal in length. The body and tail are covered with smooth, overlap- ping scales of unifomi size; a middorsal crest is absent. The dorsum is green with reddish brown crossbars. The nar- row postorbital and suborbital stripes are black. The pale green venter is separa- rated from the darker green dorsum by irregular ventrolateral white markings on the body, tail, and anterior edge of the hind limb. The iris is reddish brown; the tongue is fleshy pink; and the lining of the throat is black. $ 122+330 mm; 9 148+321 mm. Occurrence. — This arboreal species is a forest inhabitant. Precise ecological data are lacking for most of our speci- mens. In October 1971 one adult female, one adult male, and two juveniles were obtained at the edge of the forest. These were sleeping at heights of 1-2 m; one was head-down on a clump of cane- like grass, and the others were on hori- zontal limbs of bushes. The male and one juvenile were in a swamp. Life History. — Two adult females (July and October) are available; both contain small ovarian eggs. The smallest 208 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 121.— Uracentwnfiaviceps, $, 123 mm SVL (KU 126781). juvenile was obtained in October and has a snout- vent lengtli of 60 mm. In Guyana, Beebe (1944:211) found a fe- male with seven eggs in January and one with eight eggs in August. Food. — In six stomachs the only ex- ception to a variety of grasshoppers was one small beetle. In Guyana, Beebe (1944:210) found a cicada, ants, moths, berries, and seeds, in addition to beetles and orthopterans. Uracentron flaviceps (Guichenot) (Figure 121) Material. — Lago Agrio, 1; Santa Ce- cilia, 1. Identification. — This moderately large lizard with a short, depressed, spinous tail is unlike any other species in the area. The body and tail are black above and below. A narrow pinkish white col- lar is present; the head is pale orange with black flecks. Fugler and Schwaner (1968:253) noted that females have white or yellow flecks on the dorsal and ventral surfaces. S 123+70 mm; $ 117 4-66 mm. Occurrence. — Both specimens were taken from large trees after they were felled. Thus, the limited observations, supported by those of Fugler and Schwaner (1958:253), indicate that this species is arboreal in primary forest. Life History. — Nothing is known. Food. — One specimen contained many beetles. SCINCIDAE Mabuya mabouya (Lacepede) (Figure 122) Material. — Dureno, 9; Santa Cecilia, 35. Identification. — This stocky-bodied skink is easily recognized by the presence of shiny smooth scales of unifonu size on the dorsum. The species can be con- fused with only IpJiisa elegans, a slender lizard having two longitudinal rows of greatly enlarged scales, both dorsally and ventrally. The dorsum of Mabuya is brown with a bronze sheen; the dark brown flanks are separated from the mid- dorsal color by a greenish bronze dorso- lateral stripe. The venter is greenish bronze, becoming pale blue on the tail. S 94-f 132 mm; 9 100+139 mm. Occurrence. — This heliophilic lizard is seldom found in forest, and when it has been found there, individuals usually were in areas of tree falls where sunlight reached the ground. The lizards were observed most frequently in clearings, banana groves, and at the forest edge; they forage on the ground and sun on logs or stinnps. Life History. — Mabuya gives birth to EQUATORIAL HERPETOFAUNA 209 Fig. 122.— Mabuija mabouija, 5, 94 mm SVL (KU 122127). living young; females containing unde- veloped ova were found in January through November. Ten females ( March 2, June 2, and August 6) contained 4-6 (x=i5.2) well-developed embryos. The smallest female with developing embryos has a snout-vent length of 72 mm. The smallest juveniles have snout-vent lengths of 48 and 49 mm; they were ob- tained in October. The presence of ova in specimens collected through the year, possibly is indicative of acyclic breeding in the species. Food. — In 29 stomachs, 66.3 percent of the volume was orthopterans; spiders comprised 13.8 percent, and the remain- der was made up of six other groups of arthropods. Teiidae Alopoglossus atriventris Duellman Alopoglossiis atriventris Duellman, 1973, Herpe- tologica 29:228 [Type locality. — Lago Agrio, Provincia Napo, Ecuador]. Material. — Lago Agrio, 2. Identification. — This small lizard is like Alopoglossus copii and Leposoma parietale in having heavily keeled dorsal scales in oblique rows. Alopoglossus atriventris differs from both by having keeled ventral scales that are cream in juveniles and black in adult males, as opposed to smooth ventral scales that are cream (with black edges in some indi- viduals of A. copii) or pale orange (males of L. parietale). Also, the head shields in Alopoglossus atriventris are slightly rugose, whereas they are keeled in L. parietale and smooth in A. copii. The dorsal surfaces of A. atriventris are silt brown; the sides of the body and tail are black with a white stripe ex- tending from the corner of the mouth to the base of the hind limb. The ventral surfaces are cream in juveniles, cream with black edges to scales in females and subadult males, and black with white flecks in adult males. S 51+84 mm. Occurrence. — The two specimens were obtained in May while primary for- est was being cleared by bulldozers. Data on specimens collected elsewhere indi- cate that this species inhabits the leaf litter in primary forest. 210 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 123.— Alopoglossus copii, 5, 51 mm SVL (KU 122138). Life History. — No reproductive data are available; both males have seemingly fully developed testes. Food. — The stomachs of both speci- mens were empty. Remarks. — This species is known from several localities in the upper Ama- zon Basin in Colombia, Ecuador, and Peru; at some localities it occurs in sym- patry with Alopoglossus hiickJeiji and at others with A. copii. Alopoglossus copii Boulenger (Figure 123) Mflferifl/.— Santa Cecilia, 22. Identification. — This small lizard is like Alopoglossus atriventris and Lepo- soma parietale in having heavily keeled dorsal scales in oblique rows. Alopo- glossus copii differs from both by having smooth, instead of heavily keeled, ven- tral scales. The head shields of A. arti- ventris have longitudinal ridges, and the head shields of L. parietale are keeled. In A. copii the venter is cream (with black edges on the scales in some large individuals). In A. atriventris the ven- tral coloration is the same, except that the venter is black in adult males; in L. parietale the venter is cream in fe- males and pale orange in males. The dorsum of A. copii is dull brown with small, dark brown middorsal spots; the top of the head and sides of the body and tail are dark brown. A dull orange- tan dorsolateral stripe is evident on the anterior half of the body. The belly is yellowish cream, and the iris is dull red. S 56-f90 mm; 9 62+98 mm. Occurrence. — This terrestrial lizard inhabits leaf litter in primary and secon- dary forest. Four individuals were found beneath logs on wet earth near a stream in a clearing, and another individual was sunning on a log in a clearing. Individ- uals have been found March through August; 75 percent of the specimens were obtained in June and July. Life History. — Five gravid females having snout- vent lengths of 53-62 (x= 58.0) mm obtained in June and July each contained two oviducal eggs 11.7 mm in length. One female laid two eggs on 26 June. The eggs were 13X9 mm and weighed 6 gm each. Two eggs of the same size and weight were found under a log on 22 June; they hatched on 13 August. The hatchlings had snout-vent lengths of 25 mm. Food. — In 10 stomachs, spiders (47.0^), orthopterans (30.0%), and ter- mites (11.0%) comprised the bulk of the food. Sow bugs (8.0%) and caterpillars (4.0%) comprised the remainder. Ameiva ameiva petersii Cope ( Figure 124 ) Material. — Puerto Libre, 6; Santa Ce- cilia, 333. Identification. — This moderate-sized lizard with a long tail can be confused only with Kentropyx pelviceps. The lat- ter has small, keeled ventral scales and rugose head shields, whereas Ameiva has large smooth ventral scales in eight lon- gitudinal rows and smooth head shields. Kentropyx has a broad middorsal bright green stripe beginning on the head and EQUATORIAL HERPETOFAUNA 211 Fig. 124.— Ameiva ameiva petersi. Upper: S, 128 mm SVL (KU 109698). Lower: $, 103 mm SVL (KU 109695). changing to tan or reddish brown pos- teriorly. Adult male Ameiva are green posterodorsally. Females and immature males are brown with cream stripes and spots. This coloration resembles that of Tupinamhis tequixin, which is black with creamy spots. In Tupinamhis the smooth ventrals are in 20 or more longi- tudinal rows, and the dorsal scales are granular. Adult males are brilliantly colored. The top of the head is olive-gray; the thoracic region and forearms are brown or reddish brown. The rest of the dor- sum is green with two lateral brown stripes separated from one another and from the dorsal green color by rows of greenish yellow spots. The tail is brown above and laterally; the hind limbs are green with small blue spots. The chin is white or pale orange. The belly is bluish white, dark blue laterally. Adult females have an olive-brown head and pale brown dorsum, limbs and tail. A dorsolateral and lateral creamy yellow stripe borders a dark brown lateral band. In some individuals the stripes, especially the dorsolateral one, are fragmented into spots. The stripes continue onto the base of the tail. The ventral surfaces are grayish white. Juveniles are colored like females, except that the middorsal area is marked by three longitudinal rows of tan spots, i 142+303 mm; 9 125+215 mm. Occurrence. — This active, heliophilic 212 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY lizard primarily inhabits clearings, where individuals usually are seen only on sun- ny days or when the air temperature exceeds 26°C. On warm days individ- uals were seen even in light rain. Field notes on 312 specimens reveal the follow- ing sites of capture: clearings (175), sand bar of island in river (119), forest edge (14), and secondary forest (4). Three were found beneath logs in clear- ings on cool cloudy days. Fitch (1968:36) reported body tem- peratures of 38 Ameiva ameiva petersii from Santa Cecilia in March 1967. He found body temperatures of 34.0-40.0 °C (x=r37.0=t0.27) and deviation from air temperatures of +3.8-11.6°C (x=6.86± 0.33). Temperature data were recorded for an additional 22 individuals in No- vember and December 1971: body 34.7- 39.0 (x=:37.3), air 26.5-32.4 (x=29.6), deviation of body from air -)-4.2-12.1 (x=7.76)°C. Life History. — Ameiva ameiva is the only lizard at Santa Cecilia for which a seasonal reproductive cycle apparently can be demonstrated (Simmons, 1975). Ninety-three adult females are available from throughout the year, except Janu- ary. Females having oviducal eggs were found in July (3), August (2), October (1), November (2), and December (1). Mature ovarian eggs were found in one female from May and two from July. Mating was observed on 29 June. Sim- mons observed females digging nests in the sand bar on the island on 11 No- vember and 12 December and found a clutch of four eggs on 3 December. The eggs ( 19-22 X 12 mm ) hatched 140 days later on 22 April. Clutch sizes based on numbers of oviducal and matiu'e ovarian eggs are 2-6 (x=3.7, N=12). The hatch- lings from the clutch raised had snout- vent lengths of 39-41 mm. Juveniles having snout-vent lengths of 39-41 mm were found in March (15), April (1), November (3), and December (6). No Ameiva were obtained in January, and only six were collected in February, of which one was a juvenile having a snout- ' vent length of 51 mm. These data and the seasonal develop- ment of the fat bodies in females (Sim- mons, 1975) seem to demonstrate that at Santa Cecilia Ameiva ameiva has a cyclic breeding season lasting about six months; eggs mature in July through December, and young hatch in October through April. Food. — Examination of the stomach contents of 106 individuals revealed that Ameiva ameiva is a general opportunistic feeder (Table 7). As a group, orthop- terans were the most common food items eaten; these included a variety of grass- hoppers, crickets, and roaches, and mole crickets ( Gryllotalpidae ) . Cateipillars are abundant in cleared areas, and these comprised the second most common food item in Ameiva. Six individuals con- tained only maggots. These lizards were collected on the island in the Rio Agua- rico, where Hy-blown fish are not uncom- mon on the shore. One individual con- tained an adult gecko (Gomitodes con- cinnatiis); another contained the tail of a skink ( Mahmja mahoinja ) , and one con- tained two small lizard eggs. Remarks. — At the time of our first field work at Santa Cecilia in November 1966, Ameiva was observed only along the banks of the Rio Aguarico and on the island in the river. As more land was cleared, Ameiva expanded its range, fol- lowing trails, roads, and cultivated land. Apparently, without the aid of human activity in clearing land, Ameiva follows major river courses, where sand banks and associated serai vegetation provide suitable non-forest habitat. Thus, Amei- va probably reached Puerto Libre by following the Rio Aguarico. In May 1969, when primary forest was being cleared at Lago Agrio, no A7neiva was found. At that time Lago Agrio was sep- arated from the Rio Aguarico by about 3 km of primary forest. In 1971-72 Lago Agrio was connected by road to Santa Cecilia and the Rio Aguarico; many Ameiva were observed at Lago Agrio. EQUATORIAL HERPETOFAUNA 213 Fig. 125.— Arthrosaura reticulata, 9 , 70 mm SVL (KU 109823; The clearing of land at Santa Cecilia has resulted in the creation of at least marginal habitat for two species that originally were ecologically exclusive - Ameiva ameiva petersii and Kentropyx pelviceps. The ecological relationships of the two species are discussed in the account of the latter species. Arthrosaura reticulata (O'Shaughnessy) (Figure 125) Material — Lago Agrio, 1; Santa Ce- cilia, 8. Identification. — This dull brown nii- croteiid has the scales on the body ar- ranged in transverse rows; the scales on the dorsum of the body are strongly keeled and pointed posteriorly, and the posterior tips of the scales overlap the scales in the following row. The dorsal scales are separated from the ventral plates, which have rounded posterior margins, by one or two rows of small, granular scales. Prionoclactyhis argulus and P. manicatus differ from Arthrosaura by having 8-12 rows of granular scales between the dorsals and ventrals. Arthro- saura reticulata is most easily confused with Ptychoglossus ])revifrontaUs, in which the dorsal scales have low rounded keels not projecting posteriorly and non- overlapping scales in the following row. Furthermore, Arthrosaura has three su- praoculars and anterior chinshields that are larger than the second pair of chin- shields, whereas Ptychoglossus has four supraoculars and anterior chinshields that are shorter than the second pair of chinshields. The dorsum is dark reddish brown. In small individuals there are six rows (paravertebral, dorsolateral, and lateral) of small black spots with brown centers; the dorsolateral rows continue onto the tail. In some larger individuals the spots are not discrete, and the pattern consists of irregular black flecks on the dorsum. The lips are barred black and tan; the chin and throat are cream with dark brown or black spots laterally. In ju- veniles and females the belly is cream; in adult males it is salmon. The entire tail is orange in juveniles, but in adults only the ventral caudal surface is orange; the dorsum is brown. The iris is orange- brown, i 61+109 mm; 9 70+125 mm. Occurrence. — All individuals were found amidst leaf litter on the ground in primary forest by day, save one, which apparently was frightened from shelter in leaf litter at night. Life History. — Two females (one from June and one from August) each contained two oviducal eggs 13.0 and 13.6 mm in length. Food. — In six stomachs, sow bugs formed 33.3 percent and mole crickets 31.7 percent of the volume. Beetles and roaches were the other identifiable items. Bachia trinasale trinasale (Cope) ( Figure 126) Ophiognomou trinasale Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, 1868:100 [Type lo- cality.— Napo or Alto Maranon, Ecuador]. 214 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 126. — Bachia trinasale trinasale, 9 , 79 mm SVL (KU 122195). Bachia trinasale trinasale — Dixon, 1973, Misc. Publ. Mus. Nat. Hist. Univ. Kansas 57:29. Material. — Lago Agrio, 21; Puerto Libre, 2; Santa Cecilia, 1. Identification. — This slender lizard differs from all others in the area by having small forelimbs and hind limbs that are reduced to small stubs. The dor- sal surfaces of the body and tail are orange tan; the head, flanks, and dorsal stripes are dark brown, and the venter is dull brown, c^ 72+85 mm; 9 79+ 120 mm. Occurrence. — The two specimens from Puerto Libre were obtained in July; one was dug out of sandy soil, and the other was found in a trench. Both were in secondary forest near the Rio Agua- rico. The single specimen from Santa Cecilia was obtained in May from under a rotting log in a clearing. All of the material from Lago Agrio was obtained between 28 April and 4 June 1969. The lizards were unearthed by bulldozers clearing primary forest. The soil in which those were found at Puerto Libre is sandy, but the soil at Santa Cecilia and Lago Agrio is lateritic. Individuals were not observed in undisturbed forest at Lago Agrio; presumably the lizards are active in the soil or at least under the leaf litter. Life History. — Both specimens from Puerto Libre are mature females; one ob- tained on 6 July contained two oviducal eggs 12.9 mm in length, and one ob- tained on 9 July contained two large ovarian eggs 9.1 mm in length. No ju- veniles were found. Food. — Six individuals contained beetle larvae; four contained earth- worms, and one contained a centipede. Remarks. — The variation in, and dis- tribution of, Bachia trinasale, which is known only from the Amazon Basin in Ecuador and Peru, were discussed in detail by Dixon (1973). Dracaena guianensis Daudin (Figure 127) Material. — Santa Cecilia, 3. Identification. — This large lizard with a blunt head resembles a crocodile with a short snout. The dorsal scales are small with large conical scales arranged in paravertebral and diagonal lateral rows on the body, and several rows on the tail, of which the dorsolateral rows are the most prominent. The only other lizard of equal size in the area, Ttipi- nanihis tequixin, has unifonuly small dorsal scales, a long snout, and entirely different coloration. The dorsum of Dracaena guianensis is dull brown or grayish brown with irregular, yellowish tan spots on the flanks. In adult males the throat is mottled orange and black, and the belly is yellowish brown; in adult females the ventral surfaces are gray. The scales around the eye are orange or red, and the iris is dull brown, c^ 371 mm; 5 412 mm. Occurrence. — The three specimens came from near the banks of the Rio Atiuarico. Two other individuals were observed on the ground near the river; one of these took refuge in a hole in the river bank. Life History. — No data are available. Food. — Dracaena feeds on gastro- EQUATORIAL HERPETOFAUNA 215 Fig 127 — Dracaena guianensis, S, 293 mm SVL: Limoncocha, Provincia Napo, Ecuador (KU 122171). pods; it crushes the shells with the large flattened teeth and spits out the frag- ments of the shells (Conant, 1955; Van- zolini 1961). No specimens from Santa Cecilia contained food in their stomachs. Iphisa elegans Gray (Figure 128) Material — Lago Agrio, 2; Puerto Libre, 1; Santa Cecilia, 4. Identification. — This slender lizard with large, smooth scales can be con- fused only with Mahuija mahouija. In the latter species all of the scales on the body are approximately the same size, but Iphisa has two longitudinal rows of greatly enlarged scales both dorsally and ventrally. The head and body are brown with orange-tan dorsolateral stripes; the flanks are dark brown. The tail is gray in juveniles and brown in adults. The venter is gray in juveniles, creamy gray in adult females, and salmon pink in adult males. 6 58+93 mm; 9 54+93 mm. Occurrence. — All individuals were found amidst leaf litter on the forest floor, except one, which was under a log. One was in secondary forest; the others were in primary forest. Life History. — One female obtained in August contained two ovarian eggs. Food. — One individual had eaten a roach. Remarks.— Fitch (1968:37) reported the body temperature of this species as 28.2°C; he erroneously used the name Calliscincophis agilis. Kentropyx pelviceps Cope (Figure 129) Material— Dureno, 2; Lago Agrio, 2; Puerto Libre, 2; Santa Cecilia, 134. Identification. — This moderate-sized lizard with a long tail can be confused only with Ameiva anieiva petersii. The latter has large, smooth ventral scales and smooth head shields, whereas Ken- tropyx pelviceps has small, keeled ven- tral scales and rugose head shields. Fur- 216 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 128.— Iphisa elegans, ^, 51 mm SVL (KU 122172). Fig. 129.— Kenfropyx pelviceps, juvenile, 50 mm SVL (KU 122181). thermore, Kentropyx is the only lizard in the area having a broad middorsal bright green stripe beginning on the head and continuing to the midlength of the body, where it changes to reddish brown or tan. Adult males are brightly colored, and in addition to the middorsal stripe, they have a brown head and limbs with tan flecks on the latter. Bordering the mid- dorsal stripe is an irregular dark brown stripe expanded laterally to form short, transverse bars extending midway onto the flanks, which otherwise are marked by small, pale blue spots. The throat and chest are grayish white; posteriorly, this blends into salmon-orange on the belly and ventral surfaces of the limbs. Adult females usually have a broad, dark brown field on either side of the mid- dorsal stripe, which is greenish tan. The dark brown field is not so distinctly broken into bars as in males, and the lower flanks are uniform brown. The venter is pale gray. Juveniles are bronze- tan with a pale green middorsal stripe bordered by dark brown. A greenish yellow stripe extends from the eye, above the ear opening, and onto the flanks, where it persists as a series of faint dashes in half-grown individuals. $ 115 +244 mm; 9 116+209 mm. Occurrence. — Kentropyx pelviceps is a diurnal inhabitant of forest, where it usually is found in patches of sunlight on the forest floor. Individuals have been observed actively foraging amidst leaf litter, on logs, around bases of trees, and to heights of 3 m in tangled vegetation. Only one was taken in a large man-made EQUATORIAL HERPETOFAUNA 217 clearing, although natural small clear- ings, such as those created by tree falls, commonly are inhabited by this lizard. Nearly equal numbers (48 and 53) were found in primary and secondary forest; five each came from a banana grove and forest edge. Inactive individuals were found beneath logs in the forest by day, and one juvenile was sleeping on a bush about 1 m above the ground at night. Fitch (1968:36) reported on body temperatures of 36 specimens from Santa Cecilia; the temperatures were 31.0- 40.3°C (x=35.75±0.39) with a devia- tion of +6.95±0.44°C from air tempera- tures. The heliophilic behavior of this lizard precludes activity on cloudy or rainy days. Life History. — Despite the large sam- ple size, few egg-bearing females were obtained. The smallest female contain- ing ovarian eggs has a snout-vent length of 91 mm. Twenty-three females collect- ed in March through November con- tained immature ovarian eggs. One fe- male from May contained five oviducal eggs 20.1 mm in length; two from March contained four and six large ovarian eggs. Four adherent eggs were found in a cavity in a bamboo about 1 m above the ground on 8 May; the eggs were about 23 mm in length and hatched on 18 May. The hatchlings had snout-vent lengths of 38-40 mm. Juveniles having snout-vent lengths of 35-43 mm were found in March, May, June, October, and Novem- ber. Although the reproductive data are inconclusive, the presence of ovarian eggs and hatchlings from March through November indicates that some eggs also must hatch in January. Adult females are absent from our collections made in December through February, but the above data suggest that reproduction occurs throughout the year. Food. — Examination of 84 stomachs revealed that orthopterans constitute about two-thirds of the volume of food eaten. Of these, one-third are roaches Table 7. — Cnmparati\-e Food Analysis of An^eiva amciva and Kcntropyx pelviccps. ( Percent frequency of occurrence of prey/percent volume. ) Food Ameiva amciva (N=106) Kentroptjx pelviceps {N=84) Isopoda 4, Arachnida 6 Coleoptera 7. Dermaptera 3, Diptera (larvae) — 3, Hemiptera 13< Lepidoptera (larvae).. 17 Ortlioptera 31 Misc. Insecta 11 Amphibia ( Anura ) ..__ Reptilia (Sauria) 2 0/6.8 .3/3.5 17.8/13.4 ,0/6.0 0.7/0.1 ,3/2.5 ,0/5.7 ,4/4.9 .3/20.5 0.7/0.8 .3/33.3 65.7/62.3 .9/21.9 13.7/21.9 0.7/10.7 .6/0.9 0.7/0.8 and the others grasshoppers and crickets. Spiders make up about 18 percent of the volume (Table 7). One individual con- tained a frog (Eleutlicrockictylus ocken- deni) and one a lizard {AnoUs trachy- derma ) . Remarks. — At the beginning of our field studies at Santa Cecilia in 1966, Kentropyx pelviceps and Ameiva ameiva petersii were ecologically exclusive; the latter was found only along the edge of the Rio Aguarico and the airstrip, where- as Kentropyx was confined to primary forest. With continued human disturb- ance of the forest, habitats at least mar- ginally suitable for both species were created. Consequently, it is now possible to find the species together in partially cleared forest and in banana groves. Op- timal habitat for Kentropyx is receding, while optimal habitat for Ameiva is in- creasing. No interactions between the species have been observed. Both feed heavily on orthopterans, but the abun- dance of these insects, especially in par- tially cleared forest, probably precludes competition for food. Leposoma parietale (Cope) (Figure 130) 218 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 130.— Leposoma parietale, <5 , 34 mm SVL (KU 112209). Material. — Lago Agrio, 2; Puerto Libre, 28; Puerto Ore, 1; Santa Cecilia, S3. Identification. — This small lizard re- sembles Alopoglossus atriventris and A. copii in having heavily keeled dorsal scales in oblique rows. It differs from both by having keeled dorsal head scales. The head scales of AJopo^losstts copii are smooth, and those of A. atriventris have low longitudinal ridges. The ventral scales of L. parietale and A. copii are smooth, whereas those of A. atriventris are heavily keeled. The dorsum is dull reddish brown; the head and sides of the body are dark brown. A pale tan dorso- lateral stripe is evident on the anterior half of the body in most specimens. In females the chin is white, and the belly is cream; in males the chin and belly are orange. The iris is orange-tan. S 38-|-55 mm; 9 39+55 mm. Occurrence. — Leposoma parietale is active by day amidst leaf litter on the forest floor. Sixty-two percent of the specimens were found in primary forest; the remainder was about equally divided among secondary forest, forest edge, and clearings. Most individuals were active on sunny days; the lizards forage in the leaf litter and sun on the litter or small logs. A pile of wood chips at the base of a stump at the forest edge harbored a large population. Two individuals were found moving on the ground at night - one in a clearing and one in secondary forest; presumably both were frightened from their nocturnal refuges by the ob- servers. Life History. — Females produce two eggs at a time; the largest oviducal eggs are 8 mm in length. Twelve females with oviducal eggs were obtained in March, June, July, and August. Ten others from July and August have immature ovarian eggs, and three from June and July show no evidence of reproductive activity. The limited data indicate that perhaps repro- duction in Leposoma parietale is cyclic. Food. — A large variety of small in- sects are eaten. In 38 stomachs, ants and hemipterans each accounted for 21.4 per- cent of the volume. Spiders and sow bugs each amounted to less than 6 per- cent, whereas small orthopterans com- prised 14.6 percent. One individual con- tained the head of a juvenile Leposoma. Neusticurus ecpleopus Cope (Figure 131) Material. — Lago Agrio, 5; Puerto Ore, 1; Santa Cecilia, 71. Identification. — This small lizard has the appearance of a minute, short-headed crocodilian. The snout is pointed, and the dorsal scutellation consists of small scales with four longitudinal rows of en- larged keeled scales forming four ridges on tlie back and two on the tail. Other similarly colored small lizards in the area are Leposoma parietale, Alopoglossus atriventris, and A. copii, but these have uniformly large, keeled scales on the body. Dracaena fiiiianensis and Thcca- dactylus rapicauda are the only other liz- ards in the area that have large scales mixed with small ones on the dorsum. ThecadactijJus has small, granular scales on the head, a short thick tail, and broad- ly expanded toes, whereas Nciistictirtis has large plates on the top of the head, a slender tail, and slender toes. Dracaena EQUATORIAL HERPETOFAUNA 219 r'T"^riBC3BC3s*r»i ^^ Fig. 131. — Neiisticurus ecpleopus, $ , 65 mm SVL (KU 126807). has a large, blunt head and scattered large scales amidst small scales on the dorsum. Adult Neiisticurus ecpleopus are dull brown above with the flanks more olive- brown. Faint quadrangular tan spots are arranged in alternating rows on the dor- sum. In females there is a single black spot with a cream center above the in- sertion of the arm; males have three ad- ditional spots - one on the neck and two on the flanks. The belly is cream (with rusty brown edges on the scales in larger individuals). The labials and chin are pale yellow in some individuals and are marked by brown or black diagonal bars. In juveniles the dorsum is pinkish tan; the head and flanks are dark brown, and the throat and belly are black and white. $ 64+112 mm; 9 67+101 mm. Occurrence. — Neusticurus ecpleopus seems to be restricted to the immediate vicinity of small streams. All individuals were found within 1 m of water. The lizards always were associated with wet ground and active amidst leaf litter at the edge of the water. Some were under logs at the edge of the water. When dis- turbed, the lizards readily entered water and sought refuge under detritus on the bottom of the stream. One was observed in the water at night, but probably it inadvertantly had been displaced from its nocturnal retreat. Only two individ- uals were found in clearings; all others were in forest. The lizards were associ- ated only with small rivulets, usually less than 1 m wide and only a few cm deep; most such streams where Neusticurus were found had a good current and sandy bottoms. Except for the two speci- mens found in clearings, all were in heavy shade. Fitch (1968:37) reported body tem- peratures of 15 individuals as 24.5-30.4°C (x=26.8d=0.5l) with the mean devia- tfon from air temperatiue being +0.66±: 0.34°C. All of our specimens were ob- tained in March through August. Most (42 specimens) were obtained through the diligent efi^orts of Henry S. Fitch in March 1967. Because no specific attempt was made to collect these lizards other than in March and June, their absence in collections made in September through February probably more accurately re- flects the nature of collecting than sea- sonality of the lizards. Life History. — Two females (one from March and one from April) each contained two oviducal eggs 11 mm in length. Juveniles, presumably recent hatchlings, having snout-vent lengths of 23-25 mm were found in March, June, and July. Thus, the species apparently has a long breeding season, if indeed it does not reproduce throughout the year. Sherbrooke (1975) demonstrated breed- ing throughout the year in this species at Tingo Maria, Peru. Pood. — Examination of 25 stomachs revealed a diversity of food items, of which orthopterans (30.8% volume) were the most numerous, followed by spiders (19.7%), ants (14.4%), and beetle larvae ( 10.0%). One contained a centipede, and three contained lizards - Anolis trachy- derma, Leposoma parietale, and Neusti- curus ecpleopus. Remarks. — Despite the similarity in 220 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 132.— Prionodactylus argxihis, $ , 35 mm SVL (KU 126834). habits, size, and coloration of Neiisfi- curiis ecpleopiis and Echinosaura hor- ricla, the former, unhke tlie latter, does not become rigid when handled, turn over, nor wrap its tail around objects, as described by Uzzell ( 1965:89), and Levi- ton and Anderson (1966:160). When handled, Neiisticurus ecpJeopus writhes strenuously and bites ferociously. Prionodactylus argulus (Peters) (Figure 132) Material. — Lago Agrio, 12; Puerto Libre, 4; Santa Cecilia, 44. Identification. — This small microteiid has keeled dorsal scales in transverse rows; the dorsal scales arc separated from the ventrals by a zone of 8-12 small, granular scales. The broad zone of lat- eral granules immediately distinguishes this species from all other lizards in the area, except Prionodactylus manicatus, which has one frontonosal scale (two in P. ar'^ulus). Furthermore, the larger P. manicatus has uniform dark brown Hanks and a white ventrolateral stripe extend- ing from the lip to the hind limb; P. argidus has brown or reddish brown flanks with black ocelli with cream cen- ters. The dorsal surfaces of the head and body are brown to olive-brown, becom- ing paler posteriorly and changing to reddish tan or orange on the tail; the darker anterior coloration continues pos- teriorly on the body as a series of dashes or narrow longitudinal lines. The dorsal surfaces of the limbs are tan to orange with yellow spots. A dorsolateral tan stripe extends the length of the body. The flanks are orange-tan to reddish brown with a row of black ocelli with white to yellow centers. The lips are pinkish white to creamy yellow; a ven- trolateral stripe of the same color con- tinues posteriorly on the body, bordered below by a tan stripe in some specimens and blending with the venter in others. The chin, throat, and belly are metallic yellow, and the underside of the tail is orange. The iris is bronze. $ 43-f-98 mm; 9 47-|-94 mm. Occurrence. — Although Prionodacty- lus is principally a terrestrial inhabitant of primary forest, where the majority of individuals were observed moving about in leaf litter by day, the species does occur elsewhere. Ten were obtained in secondary forest, and three each in clear- ings and banana groves. Three individ- uals were observed sunning on logs, but most were in the shade. Two were ob- served on buttresses of trees; one was on the branch of a bush, and two juveniles were on herbaceous plants. Ten were sleeping at night on leaves and stems of herbs and bushes; all were at heights of less than 1.5 m, except one on a Heli- conia leaf about 2 m above water in a swamp. Life History. — All females were ob- EQUATORIAL HERPETOFAUNA 221 tained in March through September; some individuals in each of these months, except April, contained small ovarian eggs. Four females from July contained oviducal eggs 8.9-9.2 (x=9.06) mm in length; three had two eggs and one had one egg. The presence of small juveniles at various times of the year suggests a more lengthy breeding season than do the data from gravid females. The seven smallest juveniles have snout-vent lengths of 18-21 (x=20.0) mm; these were found in January (1), July (3), September (1), and December (2). Food. — In the 20 stomachs examined, spiders were the most frequent food item (46.0% frequency, 36.2% volume), fol- lowed by orthopterans (27.0% and 30.0%) . Ants, hemipterans, beetle larvae, and sow bugs comprised the rest of the iden- tifiable remains. Remarks. — The limited data on the diurnal activity of this species in bushes and other vegetation above the ground are supported by the presence of indi- viduals in the stomach of the diurnal arboreal snake OxijbeJis argenteus. Fitch (1968:38) commented on the body tem- peratures of the species under the name Prionoclactylus oshaughnessyi. He noted a body temperature of 28.8°C and air temperature of 26.7°C. Prionodactylus manicatus (O'Shaughnessy) (Figure 133) Material. — Lago Agrio, 1; Puerto Libre, 5; Santa Cecilia, 18. Identification. — This moderately small microteiid has keeled dorsal scales in transverse rows. The dorsal scales are separated from the ventral scales by a zone of 8-12 granular scales. The broad zone of granular scales distinguishes this lizard from all others in the area, except Prionodactylus argidiis, which has two frontonasal scales (one in P. manicatus) . Furthermore, the smaller P. argidus has brown or reddish brown flanks with black ocelli with cream centers; P. mani- catus has uniform dark brown flanks and Fig. 133. — Prionodactylus manicatus, hatchling, 26 mm SVL (KU 122196). a white ventrolateral stripe extending from the lip to the hind limb. The dorsum is dull olive-brown or brown with or without two rows of ir- regular dark brown spots; a bronze dor- solateral stripe extends from the orbit to the base of the tail. The flanks are dark brown, and the sides of the head are darker brown or black. A light labial stripe continues posteriorly on the ven- trolateral surface of the flank to the base of the hind limb. The stripe is white or yellowish white anteriorly and golden yellow or bronze posteriorly. The chin and throat are iridescent white, and the undersides of the forelimbs and the belly are iridescent yellow. The ventral sur- faces of the hind limbs and tail are orange. In juveniles the dorsum is dull brown, and the tail is rusty orange; the forelimbs are white, and the belly is creamy yellow. $ 65-f 93 mm; 9 70-(- 99 mm. Occurrence. — All specimens were in, or at the edge of, primary forest. One was 35 cm above the ground on the stem of a small bush; one was sunning on the base of a stilt palm, and two were sun- ning on logs. Most of the others were moving about amidst leaf litter. Several were observed to take shelter in the roots of stilt palms. About one-third of the specimens collected and more than half of those seen, but not collected, were in sunlight. Life History. — One female collected in March contained two oviducal eggs 222 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 134.— Pttjchoglosstis brevifrontalis, 9, 58 mm SVL (KU 122197). 15.0 mm in length. A female from March and one from May contained small ovar- ian eggs, and another from March showed no evidence of reproductive ac- tivity. An egg found beneath a log on 8 July hatched the following day; the hatchling had a snout-vent length of 26 mm. Eight other juveniles have snout- vent lengths of 27-33 (x=28.9) mm and were collected in May ( 1 ) , July ( 6 ) , and August (1). Food. — Identifiable remains in 11 stomachs consisted of spiders (47.6% fre- quency, 59.5% volume) and orthopterans (42.8% and 22.3%). Remarks. — Although, in comparison with Prionodactyhis arg^ulus, this species seems to be restricted to primary forest, it is heliophilic; most P. argulus were found in shade. Ptychoglossus brevifrontalis Boulenger (Figure 134) Material. — Puerto Libre, 1; Santa Ce- cilia, 1. Identification. — This long, slender microteiid has the scales on the body arranged in transverse rows; the scales on the dorsal surface of the body have low, rounded keels with straight poste- rior margins. The dorsals are partially separated from the ventrals by one in- complete row of granular scales. The ventrals are smooth and have straight posterior margins. The dorsals are par- tially separated from the ventrals by one incomplete row of granular scales. The ventrals are smooth and have straight posterior margins. Among other micro- teiids having transverse rows of scales on the body, Bachia trinasale has the hind limbs reduced to small stubs and Priono- dactyhis argulus and P. manicatus have 8-12 rows of granular scales laterally separating strongly keeled dorsals from the ventrals. Ptychoglossus brevifron- talis is most easily confused with Arthro- saura reticulata, in which the dorsal scales are strongly keeled with pointed pos- terior margins overlapping scales in the following row and the ventral plates have rounded posterior margins. Fur- thermore, Ptychoglossus has four supra- oculars and anterior chinshields that are shorter than the second pair of chin- shields, whereas Arthrosaura has three supraoculars and anterior chinshields that are longer than the second pair of chinshields. In adult females, the dorsum is red- dish brown with a faint orange-tan dor- solateral stripe and dark brown flanks. The lips are cream with brown bars; small cream spots are present on the sides of the neck. The chin is white with large, brown spots laterally. The chest is yellow, becoming orange posteriorly. The ventral surface of the tail is pinkish white. The iris is reddish brown. $ 59-|- 93 mm. Occurrence. — One was found beneath a stone at the edge of the forest, and another was amidst leaf litter in primary forest. Life History. — A female obtained in August contained two oviducal eggs 10.5 mm in length, and one found in July had two ovarian eggs. EQUATORIAL HERPETOFAUNA 223 Fig. 135. — Tupinambis teqiiixin, $, 292 mm SVL; Belem, Estado do Para, Brasil (KU 128264). Food. — Both individuals lacked food in the stomachs. Taxonomic Comments. — PfycliogJos- stis nicefori was diagnosed by Loveridge (1929) and Dunn (1944) as having 32 transverse rows of dorsal scales and lack- ing dorsolateral stripes, as opposed to P. brevifrontalis, which has 34 rows of scales and a dorsolateral light stripe. Of the two specimens from Puerto Libre and Santa Cecilia, one has 32 and the other 31 rows of scales. The faint dorso- lateral stripe evident in life is not visible in preservative. On the advice of James R. Dixon, I refer our specimens to Pty- choglossus hrevifrontaUs; P. nicefori ( Loveridge ) is considered to be a junior synonym. Tupinambis tequixin (Linnaeus) (Figure 135) Material. — Santa Cecilia, 2. Identification. — This is the largest liz- ard in the area. It has small, flat dorsal scales and 20 or more longitudinal rows of larger, smooth scales on the belly. The dorsum is black with small pale creamy white spots dorsolaterally and tan flecks overall. The adults, because of their size, can be confused only with Dra- caena guianensis - a large, brown lizard having large conical scales interspersed among small granular scales on the dor- sum. Juveniles of Tupinambis might be confused with Ameiva ameiva and Ken- tropyx pelviceps; the latter has keeled ventral scales, and Ameiva has smooth ventrals in eight longitudinal rows. Both species have small, granular dorsal scales - round in Ameiva and keeled in Ken- tropyx. The only adult from Santa Ce- cilia is a female having a snout-vent length of 307 mm, but adults of this spe- cies are known to reach snout-vent lengths in excess of 400 mm. Occurrence. — Although this large liz- ard sometimes is seen in the forest, most individuals have been observed in clear- ings, where they forage around clumps of brush and sun on logs. Life History. — A female obtained in February contained three ovarian eggs 33.7 mm in length. Beebe (1945:26) found clutches of Tupinambis eggs in termite nests in Guyana. Food. — One individual contained a caterpillar and a grasshopper. Beebe (1945:25) in reporting on the species in Guyana stated: "The tegu is an omniv- orous feeder in the widest possible mean- ing of that temi. No phylum of animals is refused, whether the organisms are small or large, living or long dead. A large number of plants, leaves and ber- ries are taken." In light of these state- ments, the food data from Santa Cecilia must be taken only for what they are - contents of one stomach. Taxonomic Comments. — I follow Presch (1973:741) in placing Tupinam- bis nigropunctatus Spix, 1825, in the sy- nonymy of T. tequixin (Linnaeus, 1758). Remarks. — During early field work at Santa Cecilia (1966-68), this large lizard rarely was seen; perhaps one was ob- served each month or so. With increased 224 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY clearing of the forest, the Hzards either have become more numerous or at least more easily observed. On sunny days in 1971-72 Tupinamhis was readily ob- served in clearings and partly cut-over areas; as many as five could be seen in one morning. AMPHISBAENIANS Amphisbaenidae Amphisbaena fuliginosa Linnaeus Material. — Lago Agrio, 3; Santa Ce- cilia, 3. Identification. — This amphisbaenian is easily distinguished from all other liz- ards by the complete absence of limbs, short blunt tail, non-imbricate scales of uniform size arranged in transverse rows around the body, and the eye covered by a large scale. The last two characters distinguish Amphisbaena from snakes. These reptiles are dull grayish white above and below, with a slight pinkish tint in some specimens; the markings consist of black transverse bars irregu- larly arranged, usually offset dorsally and less conspicuous ventrally. $ 310 mm; 9 367 mm. Occurrence. — Individuals have been ol:)served by natives on the forest floor by day. Three specimens were unearthed by bulldozers at Lago Agrio ; one of these was in the roots of a stilt palm. Life History. — No reproductive data are available. The smallest juveniles, having snout-vent lengths of 204 and 211 mm, were obtained in June and July, re- spectively. Food. — Three individuals contained food: one a centipede, one an earth- worm, and one ants. Taxonomic Comments. — Vanzolini ( 1951 ) recognized five subspecies of Amphisbaena fidiginosa; these were de- fined primarily on coloration (head spot- ted or not, venter as densely pigmented as the dorsum, and arrangement of the dorsal markings). In the specimens from Lago Agrio and Santa Cecilia, the head is spotted in two, immaculate in four; the venter is much less densely pig- mented than the dorsum in three; the dorsal spots form a checker-board pat- tern in five and transverse bars in one. These specimens seem to possess char- acters found in three subspecies - ama- zonica to the east, bassJeri to the south, and varia to the north. With such varia- tion in the small sample at hand, it is futile to recognize subspecies in the area. SNAKES Aniliidae Anilius scytale (Linnaeus) ( Plate 4 ) Materia]. — Lago Agrio, 4; Santa Ce- cilia, 3. Identification. — This cylindrical snake has a short, blunt head and tail; the ven- tral scutes are only slightly wider than the dorsal scales. The small eye is be- neath a single large scale, and there are only five enlarged plates on the top of the head. In these characters, Anilius scytale differs from all other snakes in the area. The coloration of red and black rings resembles patterns of Micrurus langsdorffi, Atractus elaps, and Oxyrho- ptis melanogenys, but all of these have enlarged ventral scutes, more than five dorsal head shields, the eye surrounded by several scales, and the tail long and tapered. The dorsum is dull dark red, and the venter is paler red. About 50 black rings, equal in width to the red interspaces, are on the body; one or two rings are on the tail. The black rings usually are com- plete across the venter. Some black rings are irregular, bifurcate laterally, or inter- connected middorsally . 6 650-f21 mm; 9 964+23 mm. Occurrence. — One individual was found on the ground in a clearing at night. Four were obtained during clear- ing of primary forest by bulldozers; pre- sumably, the snakes were in leaf litter or in the soil. Life History. — One female collected in June contained small ovarian eggs. EQUATORIAL HERPETOFAUNA 225 Fig. 136. — Boa constrictor constrictor, Santa Cecilia (Photo by Terry D. Schwaner). BOIDAE Boa constrictor constrictor Linnaeus (FiGTJRE 136) Material. — Santa Cecilia, 3. Identification. — This heavy-bodied boid does not have a conspicuously lat- erally compressed body. All scales on the top of the head and side of the snout (loreal region) are small; there are two rows of large scales between the tip and the orbit. Labial pits are absent. With the exception of Boa and Eunectes, all other boids in the area have labial pits; of these, Coralhis caninus and emjdris have laterally compressed bodies, and Epicrates cenchria has a round body. Eunectes murimis differs from Boa con- strictor by having enlarged scales on the anterior part of the head - paired inter- nasals, prefrontals, supraoculars, loreals, and preoculars. Also, the top of the head is dark brown with a pale, broad, diag- onal postorbital stripe. The top of the head is tan with a dark brown middorsal, stripe extending from the snout onto the neck. A broad, dark brown mark extends from the lip nearly to the nostril and posteriorly to the eye; a narrow, vertical, brown bar is present below the eye, and a dark brown postorbital stripe is confluent with the first transverse dark mark on the body. The body is tan or grayish tan anteriorly with rich brown transverse markings de- limiting ovoid tan blotches; rounded el- liptical brown blotches laterally enclose small cream spots. On the posterior third of the body, the dark blotches are reddish brown outlined with dark brown; the tan areas change to cream and are much narrower than those anteriorly. The belly is cream with irregular brown or black flecks and spots anteriorly, and large reddish brown blotches posteriorly. All of our specimens are juveniles, the largest having a body length of 876 mm and a tail length of 98 mm; we have seen individuals in excess of 2 m from the area. Occurrence. — Boas are uncommon at Santa Cecilia; the three juveniles were found on the airstrip - one in late after- noon and two at night. None was ob- served during clearing operations at Lago Agrio. Life History. — We have no reproduc- tive data for this viviparous species from Santa Cecilia. The smallest juvenile (body length 538 mm) was found in November. Corallus caninus (Linnaeus) (Figure 137) Material. — Lago Agrio, 1; Santa Ce- cilia, 7. Identification. — Corallus caninus is one of two arboreal boids in the area which have laterally compressed bodies. Adults are green with white markings, whereas juveniles are reddish brown to yellow with white markings. On the top of the head, the paired internasals are fol- lowed by small scales. There are four lo- reals, two or three preoculars, and deep pits in all of the upper and lower labials. The dorsal scales are smooth and in about 70 rows at midbody. The other arboreal boid is Corallus emjdris, which has two loreals, one preocular, and pits present only in posterior labials; the dorsum is brown or gray, usually with distinct dark brown or black saddles outlined with cream. Other boids in the area do not have strongly compressed bodies. 226 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 137. — Corallus canimis, $ , 952 mm SVL (KU 121833). Adults (body length of 570 mm or more) are dark green above with white markings that usually consist of a mid- dorsal stripe, short transverse bars ex- tending laterally from the middorsal stripe, and short white bars laterally; the ventral surfaces are bright yellow. Two small individuals ( 440 and 4S7 mm ) are orange-tan and cinnamon-brown dorsally with white markings. Two others (477 and 540 mm) are pale yellow dorsally. i 905+170 mm; 9 104+190 mm. Occurrence. — Most specimens were found by local residents, but I found a juvenile coiled on a limb in second growth forest and an adult coiled on the ground in direct sunlight on the airstrip. Despite these limited observations to the contrary, Corallus caninus is known to be primarily nocturnal and arboreal. Life History. — No reproductive data are available for this viviparous species. The three smallest juveniles ( < 500 mm ) were found in June, July, and Au- gust. Remarks. — The striking ontogenetic change in color from reddish brown to yellow and green has not been studied. Corallus enydris enydris (Linnaeus) Material. — Santa Cecilia, 1. Identification. — Corallus enydris is the least conspicuous of the arboreal boids having laterally compressed bodies. Adults are brown or gray, usually with distinct dark brown or black saddles out- lined with cream. On the top of the head, small scales lie posterior to the paired internasals. There are two loreals, one preocular, and deep pits in the posterior upper and lower labials. The dorsal scales are smooth and in about 45 rows at midbody. The other arboreal boid, Corallus caninus, has four loreals, two or three preoculars, pits in all labials, and about 70 rows of dorsal scales at mid- body; the dorsum is green (reddish brown to yellow in juveniles ) with white markings. Other boids in the area do not have strongly compressed bodies. The single specimen from Santa Ce- cilia is a male having a body length of 997 mm and a tail length of 247 mm. It is peculiar in having pale coloration and no conspicuous pattern. The dorsum is pale greenish tan with faint salmon blotches and dark brown X-shaped mark- ings with cream borders. The venter is cream with pale brown and salmon markings. The iris is pale green. Occurrence. — The only individual was found at night on a branch 3.5 m above the ground in secondary forest. Life History. — No data are available on this viviparous species. Remarks. — The pale coloration of the specimen resembles that of Corallus an- nulatus (Cope) but the scutellation dif- fers from that species and is like typical C. enydris from other localities in Ama- zonian Ecuador. Epicrates cenchria cenchria Linnaeus Material. — Lago Agrio, 1; Santa Ce- cilia, 8. Identification . — This r o u n d - b o d i e d boa has a distinctive coloration, consist- ing of three longitudinal dark stripes on the top of the head - one middorsal from EQUATORIAL HERPETOFAUNA 227 snout to occiput, and paired dorsolateral stripes beginning above the eyes and extending onto the neck, where they may or may not be confluent with the border of the first middorsal spot. A lateral brown stripe extends from the snout, through the eye, to the angle of the jaw. The scales on the top of the head are small except for a pair of enlarged inter- nasals. Pits are present in the lower and anterior upper labials, and the upper la- bials enter the orbit. The dorsal scales are smooth and in 49 rows at midbody. Other boids in the area have one or two rows of scales separating the la- bials from the orbit. The two species of Coralhis have laterally compressed bodies, and Boa and Eunectes lack labial pits. No other snake in the area has three longitudinal dark stripes on the top of the head and blotches on the body. The dorsum is brown to reddish brown changing to pale gray or tan lat- erally; round middorsal orange-tan spots are enclosed with black. Large, round, black, lateral spots enclose cream cres- cent-shaped marks. The venter is creamy white, and the iris is grayish tan. Char- acteristically, Epicrates cenchria has a blue iridescence in sunlight. S 1650-[- 205 mm; $ 1705+215 mm. Occurrence. — Two were in primary forest, four in secondary forest, and one in a forest-edge situation. Three were active on the ground by day and one at night; one was crawling in a bush and one in a tree at night, and one was coiled in a tree, 5 m above the ground at night. These limited observations sug- gest that Epicrates cenchria probably is nocturnal and terrestrial. Life History. — No data are available on this viviparous species. Food. — A juvenile 620 mm in body length contained the remains of a rodent. Remarks. — Our collections contain only three adults; the other six specimens are juveniles having body lengths of 457- 670 mm. Eunectes murinus murinus (Linnaeus) Material. — None. Identification. — This large boid has narrow ventrals and small, smooth dor- sals. There are two rows of scales be- tween the labials and the orbit. The scales on the anterior part of the head are differentiated into plates: paired in- ternasals, prefrontals, supraoculars, pre- oculars, and loreals. Pits are absent in the labials. All other boids in the area have broad ventrals and no more than one row of scales between the labials and the orbit. The top of the head is dark brown with a pale orange-tan diagonal bar, bordered below by a narrow brown bar, from the eye to the jjosterolateral corner of the head. The dorsum is gray to gray- ish tan, with or without a dull greenish tint, and with alternating pairs of dark brown to black, round spots; laterally there are two or three rows of small cream spots, narrowly outlined with black. The belly is dirty creamy white with irregular black marks. This giant snake, the anaconda, attains a length of 11.4m (Pope, 1961). Occurrence. — Charles M. Fugler and Ildefonso Mufioz B. (pers. com.) ob- served individuals in the backwaters of the Rio Aguarico behind the island at Santa Cecilia. Ronn Altig obtained one there on 16 February 1974; it had a total length of 2050 mm. Cofan Indians oc- casionally find the snakes along the same river. I have heard no reports of the anaconda occurring in the lakes or swamps. Life History. — No data are available on reproduction of this viviparous species in the upper Amazon Basin; Pope ( 1961) summarized existing data from through- out the range of the species. COLUBRIDAE Atractus elaps (Giinther) ( Plate 4 ) Material— hago Agrio, 4; Santa Ce- cilia, 23. 228 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Idenfificofion. — This predominately red and black snake is highly variable in color pattern and is easily confused with several other snakes having coral snake patterns. The presence of complete rings around the body, or at least across the venter, distinguishes Atractus ehips from Oxijrhopus meJanofiemjs and O. petola, both of which have 19 rows of dorsal scales (15 in A. elaps). Anilius scijtale differs by having the eye beneath a single scale, only five enlarged dorsal head shields, and ventral scutes only slightly wider than the dorsal scales, whereas Atractus claps has the eye sur- rounded by several scales, seven en- larged dorsal head shields, and enlarged ventral scutes. Atractus elaps has a lo- real; this scale is absent in most Micrti- rus, but present in Erythrolamprus aes- culapii, which has a color pattern almost identical to that of some Atractus elaps. However, some consistent differences in scutellation serve to distinguish the two species: A. elaps has a single anal plate, one pair of chinshields, and one post- ocular; E. aesculapii has a divided anal plate, two pairs of chinshields, and two postoculars. The dorsal coloration usually consists of red and black rings; the latter normal- ly are somewhat narrower than the red rings, which vary from 8 to 13 (\==:11.6, N=26) on the body. Scales in the red areas are tipped with black; in some in- dividuals the black pigment is so exten- sive as to neai^ly obliterate the red. Two individuals have red rings about ten times as wide as the narrow black rings, and three have black rings arranged in triads separated by narrow yellow rings, thus having a pattern of red-black-yel- low-black-yellow-black-rcd. The red areas vary from orange-red to vermillion and dark red; usually the color is more intense ventrally. Yellow bars border the black rings ventrally. Usually a narrow yellow or cream transverse line is present on the anterior edges of the prefrontals; in some individuals an incomplete cream or yellow transverse bar is present on the posterior part of the head, which other- wise is black, i 470+70 mm; 9 572+ 54 mm. Occurrence. — All individuals were found by day; two were moving in clear- ings; one was coiled in a building, one was under a log in mature forest, and the others were moving on the ground and amidst leaf litter in mature forest. Life History. — Two females having body lengths of 506 and 572 mm collected in May each contained six oviducal eggs 29 mm in length. Eight females from May, June, and July contained small ovarian eggs, and two females from April and July showed no evidence of repro- ductive activity. The smallest juveniles (body lengths of 167 and 182 mm) were found in May and June. Food. — One individual with a body length of 572 mm contained the remains of a giant earthworm. Remarks. — Notable sexual dimor- phism occurs in the numbers of ventrals and subcaudals: ventrals in males 140- 152 (x= 146.6, N=13), in females 150- 162 (x = 154.4, N=13); subcaudals in males 29-34 (x=31.3, N=13), in females 19-24 (x=22.6, N=10). Other features of scutellation are constant: upper la- bials 6, lower labials 7, loreal 1, post- ocular 1. The amount of black pigment on scales in the red areas is not corre- lated with size. Some small individuals have more black pigment than do some large ones. Two specimens (KU 126066- 07) have narrow black rings and very wide red rings, but in scutellation these specimens are like the normally pat- terned specimens. The ventral surface of the tail of Atractus elaps is red with one black bar. Two individuals were observed to ele- vate and coil the tail so that the bright ventral surface was visible from above. The presumably aposematic tail-coiling behavior is common in many species of snakes having red ventral caudal sur- faces (see Myers, 1965, for further dis- cussion ) . EQUATORIAL HERPETOFAUNA 229 Fig. 138.— Atractus major, 2, 347 mm SVL (KU 125987). Atractus major Boulenger (Figure 138) Material. — Lago Agrio, 11; Santa Ce- cilia, 14. Identification. — Atractus major is a predominantly brown snake with darker brown dorsal blotches. All other snakes in the area having a similar color pattern have dorsal scales in other than 17 rows, keeled scales, or the vertebral scale row enlarged, except Liophis cohella. The latter has straight-edged large, dorsal blotches that usually extend across the venter and a single anal plate. Atractus major has irregular dorsal blotches that do not extend onto the ventrals, and a divided anal plate. The top of the head is brown, dull gray, or black. In juveniles (body length to 180 mm ) the posterior part of the head is grayish white; in some larger individ- uals (body length to 307 mm) the area is dull red, whereas in larger specimens it is brown. A dark brown or black nape band is present. The dorsal ground color varies from tan to brown, sepia, or red- dish brown, with or without scattered black flecks. The dorsal blotches are brown to rust, usually outlined with cream, and highly variable in number and shape (Fig. 139). There are 21-52 (x=31.4, N=2l) blotches on the body; the blotches are 1-5 (xrr2.8, N=21) scales in length. In most specimens the venter is cream or pale yellow with scat- tered brown or black flecks, usually con- centrated in two medial rows anteriorly and lateral rows posteriorly, but in some individuals of all sizes the venter is pre- dominately brown or dull gray. The tongue is gray with a white tip. S 442+ 91 mm; 9 650-f 73 mm. Occurrence. — All individuals were ac- tive by day on the ground or in leaf litter, except one, which was concealed be- neath a palm frond. One was found in secondarv forest, and two were in clear- ings; all others were in primary forest. Life History. — Immatiu'e ovarian eggs were present in three females collected in May and one in June. A female with a body length of 650 mm contained 12 eggs. The smallest juveniles have body lengths of 145, 150, and 157 mm and are from May, March, and June, respectively. Food. — One female having a body length of 542 mm contained earth wonn. Remarks. — Notable sexual phism occurs in the numbers of ventrals and subcaudals: ventrals in males 156- 168 (x=161.3, N=9), in females 163-180 (x=172.0, N=13); subcaudals in males 41-45 (.x=43.1, N=9), in females 27-36 giant dimor- 230 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 139. A B -Dorsal color pattern at midbody of Atractus major. A. KU 125991, B. KU 125998. (xz=31.1, N=:14). Seven upper and seven lower labials occur in all speci- mens, except one male having eight up- per labials on one side and one male having eight lower labials on each side. Atiactus occipitoalbus (Jan) (Figure 140) Material. — Lago Agrio, 7; Puerto Libre, 8; Santa Cecilia, 2. Identification. — This small snake with 15 rows of smooth dorsal scales is poly- morphic in coloration. Most individuals are dark gray above and below except for a pale chin and occipital collar. Oth- ers are brown above with a row of pale spots laterally and a pale belly with a midventral row of dark spots, fused into an irregular stripe in some specimens. The gray individuals superficially resem- ble Ninia hudsoni, which has keeled dor- sal scales in 21 rows. Two other small snakes with smooth dorsal scales are similar to the brown phase of Atractus occipitomaculatus, which lacks a pre- ocular and has a long, narrow loreal. Tantilla tnehinocepJiala lacks a loreal and has a preocular and a divided anal; Rha- dinaea brevirostris has a loreal, preocu- lar, divided anal, and dorsal scales in 17 rows at midbody, reducing to 15 rows posteriorly. Both Tantilla and Rhadinaea have longitudinal dark stripes dorsally, and neither has a midventral row of dark spots. Fig. lAQ.— Atractus occipitoalbus, $, 174 mm SVL (KU 121847). EQUATORIAL HERPETOFAUNA 231 The dark color morphs are dark gray to bLack above and below; the chin is cream or white, and the occipital collar is dull orange to pinkish orange. The brown morph is grayish brown above with salmon pink spots on the fourth and fifth rows of dorsal scales; the belly is pale yellow, changing to pale orange posteriorly in males, with dark brown spots. 6 211+30 mm; 9 221 + 15 mm. Occurrence. — Three individuals were found under stones at the edge of the forest; five were on the ground in secon- dary forest, and the others were in pri- mary forest. All were found by day, either moving on the ground or amidst leaf litter, or under rocks and logs. None was in sunlight. Life History. — One female from May having a body length of 219 mm con- tained three oviducal eggs 23 mm in length. One female each from March, April, and July contained small ovarian eggs. The smallest juveniles, having body lengths of 69, 80, 85 mm, were found in July, May, and June, respectively. Food. — One individual contained an earthworm. Taxonomic Comments. — The range of variation in numbers of ventrals and sub- caudals of the brown morph (5 speci- mens) is encompassed within the range of the more common dark morph (12); ventrals in males 130-137 (x=132.8, N=7), in females 142-150 (x= 145.8, N=10); subcaudals in males 21-26 (x= 23.3, N=7), in females 12-18 (x=14.4, N=10). Savage ( 1955) named a striped speci- men from Loreto, Ecuador, as Atractus orcesi; in the same paper he referred a specimen with pale lateral spots from Sucua, Ecuador, to Atractus duhoisi Bou- lenger. The brown morph from the Santa Cecilia area is the same as the specimen called Atractus duhoisi by Sav- age. Savage ( 1960) placed both of these names of the synonymy of Atractus oc- cipitoalhus. Chironius carinatus (Linnaeus) (Figure 141) Material. — Santa Cecilia, 10. Identification. — The presence of dor- sal scales in 12 rows on the anterior half of the body distinguishes Chironius ca- rinatus from all other snakes in the area, except C. multiventris, which differs by having more ventrals and subcaudals and a proportionately longer tail (Table 8). Moreover, the two species exhibit slight differences in adult coloration. In C. carinatus the dorsum is greenish olive anteriorly and olive-brown posteriorly, and the upper labials are yellow; in C. multiventris the dorsum is dark olive- brown, and the upper labials are tan. The other two species of Chironius in the area have dorsal scales in 10 rows and single anal plates; the anal plate in C. carinatus is divided. Adults are dull greenish olive an- teriorly and dull olive-brown posteriorly; the belly and upper labials are yellow. The iris is grayish brown, and the tongue is salmon-red with a gray tip. Juveniles are more brightly colored. The dorsum is olive-green anteriorly with a bluish tint laterally and olive-brown posteriorly. The lips, ciiin, throat, and anterior one- sixth of the body are bright yellow; the rest of the venter is greenish yellow. S 681+384 mm; 9 745+405 mm. Occurrence. — All were sleeping at night. Two were on bushes, and one was 4 m up in a tree in secondary forest; one was 3 m above the ground in a tree and five were in bushes at the edge of the forest. Life History. — No data are available on reproduction. Eight juveniles were found in April and May. Pood. — An analysis of stomach con- tents indicates that juveniles eat hylid frogs - adults of HyJa hifiirca, garhei, punctata, and ruhra. One contained three metamorphosing young of Hyla marmorata, and another contained two metamorphosing young of an unidenti- fiable Hyla. Chironius carinatus never 232 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 141.— Chironius carinahis, $, 1100 mm SVL; Limoncocha, Provincia Napo, Ecuador (KU 107038). has been found active at night; thus, it probably finds these nocturnal frogs while they are hidden in dense vegeta- tion by day. Chironius fuscus (Linnaeus) (Figure 142) Material. — Puerto Libre, 2; Santa Ce- ciHa, 12. Identification. — This is one of two snakes in the area having dorsal scales in 10 rows and a single anal plate. The other is Chironius scurriilus, which dif- fers from C. fuscus by having more ven- trals, fewer subcaudals and a propor- tionately shorter tail (Table S). The paravertebral scale rows are smooth in C. scurrulus and weakly keeled in adult C. fuscus. Both juveniles and adults of the two species differ in coloration. Ju- veniles of C. scurrulus are green with rusty orange blotches in larger individ- uals, and adults are black above and rust below. Juveniles of C. fuscus are tan or gray with brown blotches; adults are brown above and yellow below. The other two species of Chironius in the area have 12 rows of dorsal scales. Considerable ontogenetic change in coloration obtains in Chironius fuscus. The smallest juveniles (225 mm) have black-edged brown bands three scales long on the dorsum, separated by gray bands one scale long; the top of the head is olive-brown. With increased size the bands become faint and finally disappear at a body length of 350-400 mm. In small juveniles, the chin and throat are white, becoming progressively darker posteriorly until deep gray on the under- side of the tail. In larger individuals the chin and throat are yellow, and the belly is tan; in large adults the chin and throat are bright yellow, and the belly is creamy yellow. The iris varies from gray to brown, usually palest dorsally. S 826-f- 448 mm; 9 758-f 399 mm. EQUATORIAL HERPETOFAUNA 233 ■■^-^^cx^osi- Fig. 142.— Chironius fiiscus, juvenile, 654 mm SVL (KU 121856). Occurrence. — Ecological data are available on 11 of 14 specimens. All were sleeping at night on bushes and low trees - seven at heights of less than 1.5 m, and four higher, to 4 m. Seven were in sec- ondary forest and two each in primary forest and at the forest edge. Usually the snakes were loosely coiled on the top of a bush or on a tangle of vines at night. At the slightest disturbance of the bush, the snake dropped to the ground and rapidy crawled away. On two such oc- casions, snakes were observed to ascend nearby bushes and again assume the sleeping posture. Life History. — No data are available. Remarks. — Despite observations on 34 individuals of the four species of Chironius, no ecological differences are apparent. All C. multiventris were on the ground by day. All C. carinatus and C. fuscus, and half of the C. scurrulus were sleeping on bushes or trees at night. Only one Chironius {scurrulus) was ac- tive in a tree by day. All individuals of C. carinatus and C. fuscus were associ- ated with forest; 25 percent of the C. multiventris and 57 percent of the C. scurrulus were found in clearings. Chironius multiventris Schmidt and Walker (Figure 143) Material. — Lago Agrio, 3; Santa Ce- cilia, 2. Identification. — This is one of two species of snakes in the area having dor- sal scales in 12 rows and a divided anal plate. The other is Chironius carinatus, which differs from C. multiventris by having fewer ventrals and subcaudals and a proportionately shorter tail (Table 8). Furthermore, adults of the two spe- cies differ in coloration. In C multiven- tris, the dorsum is dark olive-brown, and the upper labials are tan; in C. carinatus the dorsum is olive-green anteriorly and 234 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 143. — Chironhis multiventris, $ , 2570 mm SVL (KU 121855). olive-brown posteriorly, and the upper labials are yellow. The other two species of Chironhis in the area have 10 rows of dorsal scales. Adults are dark olive-brown above; the upper labials and side of the head anterior to the eye are olive-tan. The venter is bright yellow. The iris is dark brown, and the tongue is pink with a gray tip. Juveniles are more olive than brown and also have tan labials and yel- low venters. $ 1660+910 mm; $ 1420 +840 mm. Occurrence. — All individuals were found on the ground by day. One was in a clearing and three in primary forest. Life History. — No data are available. Taxonomic Comments. — As noted by Schmidt and Walker (1943a: 282), Chi- ronius multiventris differs from C. carin- atus principally by having more ventrals and subcaudals; their two specimens from southern Peru had 183 and 178 ven- trals and 172 and 202 subcaudals. Peters and Orejas-Miranda (1970:61) noted that the species was also known from Surinam and Benjamin Constant in west- ern Brasil. In addition to the specimens listed above, there are in The University of Kansas collections specimens from Limoncocha, Provincia Napo, Ecuador, and Belem, Estado do Para, Brasil. Chironius scurrulus (Wagler) (Figure 144) Material. — Santa Cecilia, 12. Identification. — The presence of dor- sal scales in 10 rows anteriorly on the body and a single anal plate distinguishes C scurrulus from all other snakes in the area except C. fuscus. The latter differs by having fewer ventrals, more subcau- dals, a proportionately longer tail, and faint keels on the paravertebral scale rows in adults ( Table 8 ) ; all dorsal scales are smooth in C. scurrulus. Furthermore, the two species differ in coloration. Ju- veniles of C. scurrulus are green with rusty orange blotches in larger individ- uals, and adults are black above and rust below. Juveniles of C. fuscus are tan or gray with brown blotches; adults are brown above and yellow below. Other species of Chironius in the area have 12 rows of dorsal scales. The smallest specimen (336 mm) is uniform emerald green above and paler green below. In half-grown individuals (600-700 mm) the dorsum is avocado green, slightly darker middorsally; the top of the head is dull brown, becoming green on the temporals and parietals. The chin is white; the rostral, edges of Table 8. — Comparison of Scutellation and Proportions in Four Species of Chironius in Amazonian Ecuador. Character C carinatus C. fuscus C. multiventris C. scurrulus N ZI 14 15 6 13 Dorsal Scale Rows .-. 12 10 12 10 Keeled Paravertebral Rows - XXX Anal Plate 2 12 1 Ventrals _.. ..- 137-152 145-154 180-188 153-163 Subcaudals 114-137 118-131 180-197 107-117 Ventrals + Subcaudals 266-281 269-281 362-377 267-279 Tail/Body 0.509-0.608 0.495-0.616 0.548-0.650 0.422-0.522 EQUATORIAL HERPETOFAUNA 235 Fig. 144.— Chrionius scurrulus, 5, 889 mm SVL (KU 121850). labials and chinshields, and anterior part of belly are rusty orange. Rusty spots are present on various scales throughout the length of the body, especially laterally. In large adults the dorsum is black, and the venter is rust; the edges of the labials are orange. The iris changes from bronze in juveniles to gray to black in adults. $ 1575+757 mm; 9 1302+550 mm. Occurrence. — We have ecological data on only seven of the 12 specimens. Four were in clearings and one each was in primary forest, secondary forest, and at the forest edge. By day, one was mov- ing on the ground in primary forest; one was eating a frog in a clearing, and one was found 3 m high in a small tree in a clearing. All others were sleeping on bushes at night. Life History. — No data are available. Food. — Four individuals contained frogs: Leptodactylus mijstaceus, L. wag- neri, Hijla lanciformis, and H. rubra. The Hyki lanciforms was being eaten by day, when the snake was found amidst a pile of palm fronds in a clearing. Remarks. — Peters and Orejas-Miran- da (1970) did not record this species from Ecuador. In addition to Santa Ce- cilia, there is one specimen in The Uni- versity of Kansas collections from Mision Bomboiza, Gualaquiza, Provincia Mo- rona-Santiago, Ecuador, 840 m. Clelia clelia clelia (Daudin) Material. — Santa Cecilia, 8. Identification. — This large snake with smooth scales in 19 rows at midbody ( 17 posteriorly) undergoes an extraordinary ontogenetic change in coloration. Juve- niles (head-body length 600 m) have a black snout and black or dark brown head with a yellow band on the neck and posterior part of the head, followed posteriorly by a wider black band. The rest of the dorsum is red. In the smallest juveniles the dorsum is bright red; in larger ones the red is duller and the tips of the scales, especially mid- dorsally, are dark brown or black. Indi- viduals 600-900 mm in head-body length are dull reddish brown to brownish black; in these the yellow nape band be- comes suffused with brown and then black. In large adults the entire dorsum is black. The dorsal color extends onto the edges of the ventrals and subcaudals, both of which are immaculate cream throughout development; the chin, lower 236 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY labials and ventral parts of the upper labials also are cream. Juveniles are easily confused with Drepanoides anomahis, Vseudohoa coro- nata, and Oxyrhopus formosus. Drepa- noides has 15 rows of dorsal scales, and Pseiidohoa has 17 rows; in both of these, only the snout is black, and the head from the eyes posteriorly onto the neck is creamy yellow. In Oxyrhopus formo- sus the tip of the snout is white; in adults, which are colored like Clelia, the top of the head and nape are brown or black, whereas in juveniles, which have bands on the body, the top of the head is brown or black, and the nape is cream. Further- more, Oxyrhopus formosus has fewer than 215 ventrals; Clelia has more than 215. Adults of Clelia clelia can be con- fused with the uniform black Drymolu- her dichrous; the latter species has dor- sal scales in 15 rows and a single anal plate, whereas Clelia has 19 rows and a divided anal plate. The middorsal scale row in Clelia is slightly enlarged. Variation in scutella- tion is given in Table 9 (see accounts of Oxyrhopus). The iris is reddish brown. c5 1057+157+ mm (incomplete); 5 2070+280+ mm (incomplete). Occurrence. — Four specimens lack ecological data. One was moving on the ground in a clearing in the early morn- ing. One was in water in an open marsh at night, and two were moving on the ground at night in a clearing. These limited data suggest that the species may be primarily nocturnal and definitely ter- restrial. Life History. — A female collected in May, and 2070 mm in body length, con- tained 20 immature ovarian eggs. Food. — One individual had eaten a diurnal snake, Leimadophis reginae. Another contained an unidentified rodent and when found at night was eating an Ameiva ameiva, which evidently had been discovered in a nocturnal retreat. Dendrophidion dendrophis (Schlegel) Material. — Santa Cecilia, 2. Identification. — Only two snakes in the area have keeled dorsal scales in 17 rows at midbody. Dendrophidion den- drophis and Drymohius rhomhifer. In the latter the anal plate is divided, whereas in Dendrophidion it is single. Both species have dark brown or black dorsal blotches; in Dendrophidion the blotches have straight, transverse edges and extend laterally to the first scale row, whereas in Drymohius the blotches are ovoid, much wider middorsally than lat- terally, and extend laterally to the second or third scale row. Juveniles of Drymo- Table 9. — Scutellation in Species of Clelia and Oxyrhopus in Amazonian Ecuador. ( Means in parentheses below observed ranges. ) Character C. clelia N 3$, 59 Preocular in contact with frontal Upper Labials .— 7-8 (7.6) Lower Labials 8 Ventrals — $ $ 218-225 (221.5) 9 $ 228-242 (234.2) Subcaudals _____ _ ,5 $ 94-96 (95.0) 9 9 77-86 (82.5) O. formosus 1$, 59 O. petola 115, 119 O. melano- genijs 95, 59 X X 8 8-9 8-9 (8.1) (8.1) 8-10 10-11 9-10 (8.7) (10.4) (9.9) 191 206-221 187-198 (214.5) (191.9) 194-208 209-225 201-206 (199.2) (216.7) (204.0) 95 110-122 82-94 (116.7) (88.0) 74-80 99-112 76-84 (76.4) (103.8) (78.8) EQUATORIAL HERPETOFAUNA 237 liiher dichrous have a body pattern simi- lar to that of young Dendrophidion, but in the latter the head is unicolor brown, whereas in Drijmoluher the head is tan or gray with dark brown markings. Fur- thermore, DnjmoJuher has smooth dorsal scales in 15 rows. The top of the head is olive-brown. The body has 49-55 dark brown blotches 3 scales in width, separated by narrow (/2 scale in width) creamy tan inter- spaces on the anterior half of the body. The anterior edge of each blotch is dark brown or black posteriorly the pale in- terspaces do not extend across the body, so the blotches are delimited only by their dark anterior borders. Laterally the interspaces are orange. The dorsal scales on the anterior part of the body have faint blue edges. The chin and throat are white, and the belly is lemon yellow. The tongue is gray. 9 670-f- 90-f- mm (incomplete). Occurrence. — Ecological data are available for only one specimen, an adult female which was sleeping at night on a bush in primary forest. My observations on this species in Central American and elsewhere in the Amazon Basin show conclusively that it is an active diurnal, terrestrial species, inhabiting forest. Life History. — A female from June contained six ova 30.2 mm in length. Food. — The adult female contained an adult frog, Eleutherodactylus conspi- cillatus. Taxonomic Comments. — The taxo- nomic status of Amazonian Dcndrophid- ion is unsettled, but the late James A. Peters, who was studying Dendrophid- ion, examined the two specimens from Santa Cecilia and tentatively referred them to D. dendrophis. Remarks. — These snakes have long tails which break readily. Most adults in museum collections have incomplete tails. A juvenile from Santa Cecilia has a body length of 318 mm and a tail length of 246 mm. Assuming approxi- mately the same ratio in the adult, the adult female from Santa Cecilia must have had a tail length of about 515 mm. Dipsas catesbyi (Sentzen) (Figure 145) Material. — Lago Agrio, 2; Puerto Libre, 1; Santa Cecilia, 28. Identification. — Of the three species of Dipsas, the only snakes in the area having dorsal scales in 13 rows and an enlarged vertebral row, this is the only species having a loreal and a preocular. Dipsas indica has a preocular and no loreal, and D. pavonina has a loreal and no preocular. Furthermore, D. catesbyi has only one anterior temporal, whereas the other species of Dipsas have two. Other snakes having an enlarged verte- bral scale row have the dorsal scales in 15 {Imantodes lentiferus), 17 (Iman- todes cenchoa) or 19 (Leptodeira and Tripaniirgos) rows. The head is black with a white line across the snout; usually the line con- tinues across the chin. The dorsum is marked by paired elliptical dark brown blotches, meeting or not, middorsally. The dark chocolate brown blotches are bordered with white, especially anterior- ly, and stand out in marked contrast with the reddish brown ground color. The venter is white with longitudinal black marks and flecks, especially posteriorly. The iris is reddish brown. S 489+191 mm; 9 526+200 mm. Occurrence. — All active individuals were found at night. Twelve were in primary forest, seven in secondary forest, and one in a banana grove. Individuals most frequently were observed crawling slowly along branches of bushes and trees, some as high as 4 m above the ground. Two were retrieved from the tops of large trees ( > 20 m ) when the trees were felled by day. One individual was crawling on the forest floor at night. Life History. — Three females (April 2, June 1) contained 1, 4, 1 oviducal eggs, the largest 35.3 mm in length. The small- est juvenile (193 mm) is from June. 238 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 145.—Dipsas catesbyi, 5 , 514 mm SVL (KU 146736). Three others have body lengths of 216 mm (March), 224 and 230 mm (July). Remarks. — By comparison with Pe- ters' ( 1956 ) analysis of variation in Dip- sas cates])i/i, our material from Santa Ce- cilia is rather uniform. There is variation of one or two preoculars and postoculars; in one individual the loreal is fused with the preocular on both sides of the head. No ecological differences are appar- ent among the three species of Dipsas in the area. I have the general impression that Dipsas indica is more commonly found in dense, shrubby vegetation than the other species, but there are too few observations on that species to substan- tiate the impression. Perhaps the three species feed on different species of snails; Dipsas indica has a proportionately larg- er head and heavier body than the other species and thus possibly feeds on larger snails. Dipsas indica ecuadorensis Peters (Figure 146) Material. — Puerto Ore, 1; Puerto Libre, 1; Santa Ceciha, 11. Identification. — This blunt-headed snake with dorsal scales in 13 rows and the enlarged vetrebral row differs from other species of Dipsas by lacking a loreal and having a single preocular. Dipsas catesbyi has a loreal and preocu- lar, and D. pavonina has a loreal and no preocular; furthermore D. pavonina has three postoculars, whereas D. indica has two. Moreover, D. indica is the only species of Dipsas in the area having a dark venter. Other snakes having an enlarged vertebral scale row have the dorsal scales in 15 (Imantodes lentifer- FiG. 146. — Dipsas indica ecuadorensis, $ , 821 mm SVL; Bermejo, Provincia Napo, Ecuador (KU 121871). EQUATORIAL HERPETOFAUNA 239 Fig. 147. — Dipsas pavonina, $ , 477 mni SVL; Pilcopata, Departamento Cuzco, Peru ( KU 139422). us), 17 (Imontodes cenchoa), or 19 (Leptodeira and Tripanurgos) rows. The head is brown with pale yellow and dark brown longitudinal streaks; the body is gray or grayish brown with dark brown blotches that are wider ventrally than dorsally. The throat is pale yellow, suffused with gray in some individuals, and small yellow spots are present ven- trolaterally. The belly is dark brown. The iris is grayish green, i 510+197 mm; 9 732+242 mm. Occurrence. — All individuals were on branches or leaves of bushes and trees at night. Six were in second growth forest; the others were in primary forest. Two were on vegetation higher than 1.5 m above the ground; the others were on lower vegetation. Three were obtained in July, two each in April, May and Au- gust; no more than one was found in any other month. Life History. — None of the three adult females contained eggs. The small- est juvenile ( 264 mm ) was found in Jan- uary. Food. — One individual contained the remains of the body of a snail. Dipsas pavonina Schlegel (Figure 147) Material. — Puerto Libre, 1; Santa Ce- cilia, 2. Identification. — This slender Dipsas with dorsal scales in 13 rows and an en- larged vertebral row differs from the other Dipsas in the area by having three postoculars, a loreal, and no preoculars. Dipsas cateshiji has a loreal and preocu- lar, and D. indica has a preocular and no loreal; both species have two postoculars. Furthermore, D. pavonina differs in col- oration; the belly is unifonnly pale tan, whereas the belly is dark brown in D. indica and white with black streaks in D. cateshiji. In the latter, the body blotches are ellipitical, narrower dorsally and ventrally than laterally, whereas in D. pavonina the blotches extend across the body and are wider dorsally than ventrally. Other snakes having an en- larged vertebral scale row have the dor- sal scales in 15 (Imantodes lentiferus), 17 {Imantodes cenchoa), or 19 (Lepto- deira and Tripanurgos) rows. The head is black with a narrow white line across the snout that extends onto the lower labials but not across the chin. The nape band and first four or five spaces between the black blotches are white. The interspaces become progres- sively more tan posteriorly, but a narrow white border persists around the blotches. The blotches are much broader than the interspaces, extend across the body to the first row of dorsal scales, 240 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 148.— Drtjmobius rhombifer, 9, 1111 mm SVL (KU 121878). and are slightly wider dorsally than lat- erally. The venter is nnifoniily tan. The iris is dull brown. £ 446+206 mm; $ 381+176 mm. Occurrence. — All were active at night - two in primary forest and one in sec- ondary forest. One was crawling up a tree trunk about 2 m above the ground; one was crawling along a vine about 3 m above the ground, and the other was on a limb about 1.5 m above the ground. Life History. — No life history data are available. Drepanoides anomalus (Jan) Material. — Dureno, 1. Identification. — This red snake with a black snout, creamy white head, and black nape band differs from other simi- larly colored species by having the dorsal scales in 15 rows. Fseudohoa has 17 rows, and Clelia and OxyrJiopus have 19 rows. The snout, including all scales anterior to the eye, the anterior part of the frontal, and the anterior part of the chin, is black. The rest of the head and nape are creamy white. The nape color is bordered posteriorly by a black band six scales wide. The rest of the dorsum is red with black on the posterior tips of the scales. The dorsal color does not extend onto the cream venter. The single specimen is a male having 173 ventrals, 78 subcaudals, single anal plate, six upper and lower labials, one preocular, two postoculars, and no loreal. c^ 330+109 mm. Occurrence. — One was crawling on the ground in a clearing at night. Life History. — No data are available. Drymobius rhombifer (Giinther) (Figure 148) Material — Puerto Libre, 2; Santa Ce- cilia, 4. Identification. — This is one of two snakes in the area having keeled dorsal scales in 17 rows at midbody. The other, Dendrophidion dendropJiis has a single anal plate and straight-edged, transverse EQUATORIAL HERPETOFAUNA 241 Fig. 149.— Drymoluber dichrous, juvenile, 334 mm SVL (KU 121882). body blotches extending to the first scale row. Dnjmohius rhomhifer has a divided anal plate and ovoid body blotches that are much wider middorsally than lateral- ly and extend laterally to the second or third scale row. The color pattern of juvenile Dnjmohius is similar to that of juveniles of Xenodon severus and Both- rops atrox. The latter has keeled dorsal scales in 25 rows at midbody and a large pit between the nostril and the eye. Xenodon has smooth dorsal scales in 21 oblique rows. Juveniles are grayish tan with darker blotches, which are chocolate brown peripherally and reddish or grayish brown centrally. The top of the head is olive-tan with dark brown spots on the parietals; the side of the head is grayish white. The chin and throat are pale gray; the rest of the venter is cream with irregular, dark brown marks laterally. The coloration has less contrast in adults. The dorsal ground color is darker, so the blotches are not as distinctive; blotches on the head are indistinct. The venter is grayish white. The iris is bronze. $ 907+361 mm. Occurrence. — Four individuals of this diurnal, terrestrial snake were in primary forest, and two were in secondary forest. Two individuals were sleeping in low bushes at night. Two were taken in July and one each in February, March, April, and May. Life History. — No data are available. Food. — One individual contained re- mains of an unidentifiable lizard. Drymoluber dichrous (Peters) (Figure 149) Material. — Puerto Libre, 2; Santa Ce- cilia, 5. Identification. — The combination of smooth dorsal scales in 15 rows, no en- larged vertebral scale row, two anterior temporals and a single anal plate dis- tinguishes Drymoluber from all other snakes in the area, except Drepanoides anomahis, which has a white head, with a black snout and nape band, and a red body. Adult Clelia clelia are colored like adult Drymoluber, but Clelia has 19 rows of dorsal scales and a divided anal plate. The ontogenetic change in coloration involves a juvenile blotched pattern dor- sally that changes to uniform black in adults. In juveniles, the top of the head is pale gray with two transverse brown marks and a pair of large brown parietal spots. The side of the head is grayish white with brown vertical markings. The dorsal ground color is reddish tan with dark reddisli brown blotches becoming darker posteriorly. The blotches are 2-3 scales wide, have straight edges, and ex- tend onto the edges of the ventrals. An- teriorly the interspaces are 2-3 scales wide; posteriorly they become progres- 242 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY sively narrower until they disappear on the proximal part of the tail. The chin and throat are white with black marks; the rest of the venter is creamy gray. Adults are uniformly black above and cream below. The edges of the ventrals are black, and the upper labials are cream. The iris is tan dorsally with the remainder dark reddish brown, and the tongue is black, c^ 900+322 mm; 2 725+145 mm. Occurrence. — This diurnal, terrestrial snake sleeps in low vegetation at night. Three individuals were in primary forest and one in secondary forest. Life History. — No data are available. Food. — One individual disgorged two lizards - Neusticurus ecpleopus and Ano- lis (unidentifiable to species). Erythrolamprus aesculapii aesculapii (Linnaeus) ( Plate 4 ) Material. — Lago Agrio, 2; Santa Ce- cilia, 3. Identificatiot^. — Three genera of snakes in the area have dorsal scales in 15 rows and patterns of red, yellow, and black rings. Of these, Atractus elaps dif- fers by having a single anal plate and one pair of chinshields. ErythroJatnprus and the four species of Micnirus have divided anal plates and two pairs of chin- shields. Erythrolamprus differs from Mi- crurus by having two (instead of one) secondary temporals and by lacking a poison fang anteriorly on the maxillary. The coloration of the head of Erythro- lamprus differs from that of all Micrurus, except M. lemniscatus. In the latter, the snout is black, and there is a yellow band anterior to the eyes. In Erythrolamprus the snout is yellow with a black band enclosing the eyes and a yellow band posterior to the eyes. The snout is yellow-tan, followed by a broad black band enclosing the eyes, and a broad yellow band extending near- ly to the posterior tips of the parietals. The nape is black, interrupted or not, by a narrow yellow ring. The body pattern of rings (red-black-yellow-black-red) commences posterior to the nape. The black rings are four or five scales wide middorsally and only two or three scales wide ventrally. The yellow rings are two or three scales wide, and the red rings are five or six scales wide. The posterior tips of the scales in the yellow and red rings are black. One individual has the first red ring 27 scales wide; the rest of the pattern is noiTnal. $ 534+64 mm; 9 667+89 mm. Occurrence. — All individuals were on the forest floor by day; one was in secon- dary forest, and the others were in pri- mary forest. Life History. — Females from May and October contained immature ovarian eggs. Food. — Two individuals had eaten snakes: Atractus occipitoalhus. Remarks. — When disturbed, one in- dividual elevated the head and spiraled the tail upward. It is questionable why this presumably aposematic behavior was displayed by Erythrolamprus aescu- lapii, because the ventral surface of the tail is red like the dorsum. Helicops angulatus (Linnaeus) (Figure 150) Material. — Puerto Libre, 1; Santa Ce- cilia, 7. Identification. — The small eyes, dor- sal nostrils, and small, single internasal distinguish snakes of the genus Helicops from others in the area. Helicops angu- latus has keeled dorsal scales in 19 rows at midbody (17 rows posteriorly) and one anterior temporal, whereas Helicops petersi has 21 rows at midbody ( 17 rows posteriorly) and two anterior temporals. The two species also differ in color pat- tern. Helicops angulatus has about 20 distinct, dark brown blotches across the dorsum and extending onto, or continu- ous across, the venter. Helicops petersi has about 40 indistinct middorsal dark blotches alternating with smaller lateral blotches. The dorsal ground color varies from EQUATORIAL HERPETOFAUNA 243 Fig. 150.— Helicops angulatus, $,666 mm SVL (KU 121885). olive-green to dull brown with dark brown blotches that are reddish brown centrally in some individuals and nar- rowly bordered by black in some speci- mens. The top of the head is olive-brown with a dark brown inteorbital bar evident in some individuals. The chin is white to lemon yellow, and the belly is creamy yellow to orange with dark brown to black extensions of the dorsal blotches. The ventral surface of the tail is banded black and creamy gray in juveniles and dull gray in some adults. The iris is dull olive-brown. 6 315+162 mm; 5 475+ 191 mm. Occurrence. — One individual was at the edge of the river by day. All others were found in water - small pools or streams at night. Life History. — One female from June having a body length of 475 mm con- tained 11 ovarian eggs. Food. — One individual contained the remains of an unidentifiable tadpole. Remarks. — These snakes depress the body, flatten the neck and bite viciously. Helicops petersi Rossmann Helicops petersi Rossmann, 1976, Occas. Pap. Mus. Zool. Louisiana State Univ., 50:2 Type locality. — Mishualli River, 1 mi NE Tena, Provincia Napo, Ecuador]. Material — Puerto Libre, 1; Santa Ce- cilia, 2. Identification. — This snake, like Heli- cops angulatus, has small eyes, dorsal nostrils, and a single internasal, thereby differing from all other snakes in the area. Helicops petersi has keeled dorsal scales in 21 rows at midbody (17 rows posteriorly) and two anterior temporals, whereas Helicops angulatus has 19 rows at midbody (17 rows posteriorly) and one anterior temporal. Furthermore, H. 244 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 151. — Imantodes cenchoa cenchoa, juvenile, 551 mm SVL (KU 121890). petersi differs from H. angulatus in col- oration; the former has about 40 indis- tinct middorsal blotches alternatincf with smaller lateral blotches and a dark vent- er, whereas the latter has about 20 dis- tinct middorsal blotches extending onto, or across, the pale venter. The dorsum is dull yellowish tan to grayish tan with olive-brown to dark gray blotches; the venter is gray to black with cream to tan transverse streaks. The iris is tan. s 444+225 mm; 9 340+29+ mm. Occurrence. — One individual was partially buried in the sand at the edge of the river by day; the others were in a back water of the river at night. Life History. — A female collected in July has a body length of 340 mm and contained 12 small ova. Remarks. — The limited data suggest that H. petersi is limited to major rivers, whereas H. angulatus is more wide- spread, inhabiting small streams, swamps, and temporary pools. Imantodes cenchoa cenchoa (Linnaeus) (Figure 151 ) Material. — Lago Agrio, 2; Puerto Libre, 1; Santa Cecilia, 32. Identification. — This long, slender snake with a small, l)lunt head and lat- erally compressed body with a greatly enlarged vertebral scale row has the dorsal scales in 17 rows. No other snake in the area has enlarged vertebral scales and dorsal scales in 17 rows. Imantodes lentiferus has 15 rows of scales and small- er reddish tan dorsal blotches, whereas J. cenchoa has larger dark brown blotches. Of the other genera having an enlarged vertebral scale row, Dipsas has dorsal scales in 13 rows and Leptodeira and Tripanurgos in 19 rows. The top of the head is dark brown; the edges of the scales are cream. The transverse body blotches are dark choco- late brown, and the interspaces are cream to tan. The belly is cream to yellowish tan with small brown flecks. The iris is creamy tan, with a greenish tint in some EQUATORIAL HERPETOFAUNA 245 individuals. $ 811+357 mm; 9 827+ 345 mm. Occurrence. — This nocturnal, arbo- real snake is about equally abundant in primary and secondary forest. Two- thirds of the individuals were found at heights of less than 1.5 m above the ground; others were found to heights of 4 m. One individual was coiled in a bamboo stump by day. These snakes are capable of extending more than half their total length into space and require only a flimsy branch on which to rest the chin before proceeding to move from one branch to another. Life History. — Four of nine (44.4%) of the adult females are gravid. Females with oviducal eggs were found in March, June, July, and November. Three con- tained two eggs, and one had one egg. The largest egg was 29.6 mm in length. Small juveniles (314-347 mm) were ob- tained in May (2), June (1), July (1), and August (1). These data support Fitch's ( 1970:152) contention that Iman- todes cenclioa breeds throughout the year. Food. — Two individuals contained lizards - Anolis fuscoouratus and A. punctatus. Remarks. — Both Imantodes cenclioa and /. lentiferus are active primarily in low vegetation at night. The latter has been found only in secondary forest, whereas 7. cenchoa is equally abundant in primary and secondary forest. With increased human disturbance of the for- est at Santa Cecilia, I. lentiferus seems to be increasing in abundance relative to 7. cenclioa. The ratio of specimens of 7. cenclioa to 7. lentiferus in collections made in 1968-69 is 4:1, in 1970-71 1:1. The limited data on stomach contents re- veals definite differences in food prefer- ences - lizards in 7. cenchoa and frogs in 7. lentiferus. Imantodes lentiferus (Cope) Material. — Puerto Libre, 3; Santa Ce- cilia, 24. Identification. — The presence of a greatly enlarged vertebral scale row, slender laterally compressed body, and small blunt head distinguish this spe- cies from all other snakes in the area, except Imantodes cenchoa. The latter has 17 rows of dorsal scales, whereas 7. lentiferus has 15 rows. Furthemiore, 7. lentiferus has smaller reddish tan body blotches, and 7. cenchoa has larger dark brown blotches. Of the other genera having an enlarged vertebral scale row, Dipsas has dorsal scales in 13 rows and Leptodeira and Tripanurgos in 19 rows. The top of the head is tan with three dark brown spots, fused into one spot in larger individuals. The body is pale tan with brown to reddish brown blotches, narrowly outlined with dark brown, much narrower laterally than dorsally, and extending laterally to the second or third row of scales. The chin is white or cream; the belly is pinkish tan with small black flecks. The iris is yellowish tan. S 682+308 mm; 9 710+291 mm. Occurrence. — All individuals were found in secondary forest; three-fourths of the individuals were crawling in bushes and herbs at heights of less than 1.5 m above the ground, whereas others were found higher in bushes and trees. Life History. — One female (667 mm) deposited three eggs ( 10 mm in diameter and 20, 23, and 25 mm long) on 14 Au- gust. Two eggs hatched on 25-26 No- vember. John Simmons (field notes, 25 November 1971) described the hatching: 0820 hr, 22° C, head appeared in slit 5 mm long. 1200 hr, 28°C, head still in slit, now about 7 mm long. 1310 hr, head withdrawn into shell. 1600 hr, head again in slit. 1707 hr, 27 °C, second slit on opposite side of shell. 1730 hr, 25.5°C, approximately 25 mm of head and neck extended through second slit and at an- gle of about 45° from axis of egg; flicking of tongue. 1800 hr, dusk, snake rapidly emerged from shell and crawled onto plants. The hatching of the second egg on 26 November corresponded to the 246 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY \ X Fig. 152.— Leimadophis reginae, S, 575 mm SVL (KU 121324). first. Again, two slits were made. The first was noted at 0740 hr; the snake emerged from the egg near dusk, 1745 hr. Perhaps the time of emergence was pur- posefully delayed until dusk, at which time these snakes nomially become ac- tive. The young had body lengths of 197 and 204 mm and tail lengths of 82 and 87 mm. Two slightly larger juveniles (243 mm) were found in July. Food. — Stomach contents of four in- dividuals consisted only of frogs: Hyla calcarata, Ihjla geographic a, Eleuthero- dactijhis lacrimosus, and Eleutherodac- tylus sp. Remarks. — See the account of Iman- todes cenchoa cenchoa for a discussion of the ecological relationships of the two species of Imantodes. Leimadophis reginae (Linnaeus) (FiGunii 152) Material — Durcno, 4; Lago Agrio, 1; Santa Cecilia, 8. Identification. — The presence of smooth dorsal scales in 17 rows at mid- body and a divided anal plate distinguish this species from all others in the area except Leimadophis sp., RJiadinaea brev- irostris, and Liophis coI)eUa. The latter has dark bands encircling the body, and Rhadinaea has distinct dark brown and tan stripes dorsally and an immaculate venter. Both species of Leimadophis have black transverse marks or a check- erboard pattern on the belly and lack distinct dorsal stripes, at least anteriorly. In Leimadophis reginae the upper lip is cream, whereas in L. sp. the edge of the upper lip is dark, delimiting a white la- bial stripe. Leinmdophis sp. has 157 ventrals, and L. reginae has fewer than 150 ventrals. The dorsum is tan with a trace of green anteriorly on the first two scale rows. A narrow dark brown line extends from the nostril through the lower edge of the orbit to the angle of the jaw. A faint dark brown stripe usually is evident posteriorly (beginning about two head lengths anterior to the anus) on the EQUATORIAL HERPETOFAUNA 247 third scale row and extends to the tip of the tail. The chin and tliroat are im- maculate cream; the belly and underside of the tail are yellow with irregular transverse black bars. The iris is gold above and reddish brown below, and the tongue is dark gray, i 391+145 mm; 9 436+127 mm. Occurrence. — Most were found active on the forest floor by day; one was in secondary forest and the others in pri- mary forest. One was active in a clear- ing by day. Four were found in August, three in May, two in July, and one each in February, April, and September. Life History. — One female collected in February contained three ovidu- cal eggs 27.5 mm in length, and one from April contained six maturing ovar- ian eggs 22.9 mm in length. Fitch (1970:137) noted that "Gravid females of L. reginae are represented in the Bass- ler Collection from Iquitos, Peru, for each month of the year, except for May, July, and October when no adult fe- males were collected, and January and April, each represented by only one adult female." Food. — One individual contained an adult Leptodactylus wagneri and numer- ous, small tadpoles of Hyla lanciformis. Leimadophs sp. Material. — Santa Cecilia, 1. Identification. — This is one of four snakes in the area having smooth dorsal scales in 17 rows at midbody and a di- vided anal plate. Of these, Liophis co- hella has dark bands encircling the body, and Rhadinaea brevirostris has distinct dark brown and tan stripes dorsally and an immaculate venter. Leimadophis sp. and L. reginae have transverse black marks or a checkerboard pattern of black marks on a cream venter. In Leima- dophis sp. the edge of the upper lip is dark, delimiting a white labial stripe, and in L. reginae the entire upper lip is cream. Leimadophis reginae has fewer than 150 ventrals, whereas L. sp. has 157. The top of the head and middorsum of the body and tail are dark brown; on the body the dark brown fades to tan on scale rows 3-5 and to cream on scale rows 1-2. Posteriorly, the scales on the fourth scale row are edged with dark brown, thereby giving them a darker appearance and an illusion of a tan stripe on the fifth scale row. The upper labials are white with a dark brown lower edge. The lower labials and chin are black and white, and the belly is cream with irreg- ular, black transverse marks. The under- side of the tail is immaculate cream. $ 250+57+ mm (incomplete). Occurrence. — The single specimen was dead on a trail in the forest by day. Life History. — No data are available. Taxonomic Comments. — Charles W. Myers has studied this specimen (KU 112275) and is unsure if an available name exists. Leptodeira annulata annulata (Linnaeus) (Figure 153) Material. — Puerto Ore, 1; Santa Ce- cilia, 43. Identification. — Leptodeira anmdata is one of two species of snakes in the area having the dorsal scales in 19 rows and the vertebral row enlarged, although in some individuals of this species, the ver- tebral row is only slightly enlarged. The other is Tripanurgos compressus. Lepto- deira differs from Tripanurgos by having a divided anal plate (single in Tripan- urgos), one anterior temporal (two), and broad middorsal brown blotches on the body ( narrow transverse blotches ) . Oth- er snakes having laterally compressed bodies and enlarged vertebral scale rows have fewer rows of dorsal scales: Iman- todes cenchoa (17), Imantodes lentiferus (15), andD//wa.s' (13). The head is brown, with or without a dark brown spot or stripe on the nape; a broad, dark brown postorbital stripe is present. The dorsum is tan, grayish tan, or reddish tan with dark brown middor- sal blotches extending to the fourth or 248 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 153. — Leptodeira annulata annulata, $ , 568 mm SVL (KU 109840). fifth row of scales. The blotches usually are connected on the midline, especially anteriorly, where the confluent blotches are no more than five scales in width; thus, in many individuals the anterior pattern consists of a broad, zig-zag mid- dorsal stripe. The venter is creamy white to yellowish tan anteriorly, changing to orange-tan or pinkish tan posteriorly. The underside of the tail is brown or grayish tan. The iris is orange-tan to pale brown. In juveniles the occipital region is white. S 560-f 171 mm; ? 578-fl82mm. Occurrence. — This nocturnal, arbo- real, snake is most abundant in secondary forest; 78 percent were found there, whereas only 17 percent were found in primary forest. Others were found in buildings or at the edge of the forest. Most ( 65% ) were on vegetation less than 1.5 m above the ground; others were found up to heights of 6 m. Active Lep- todeira most frequently are observed in herbs and bushes in and around ponds where frogs are calling; 65 percent of the snakes were captured within 3 m of water. Only one individual was found by day; it was in a bromeliad on a tree limb about 2.5 m above the ground. The monthly distribution of speci- mens is biased by the inequable distribu- tion of field work, but even so there is a heavy concentration of snakes in May- July. The number of Leptodeira ob- tained throughout the year is: February (2), March (4), April (4), May (9), June (12), July (10), August (1), Oc- tober ( 2 ) . Three of the snakes from July are hatchlings. Because these snakes feed almost exclusively on frogs, it might be anticipated that they are most active at times of greatest anuran breeding ac- tivity - April-May, October-November. Duellman (1958:119) showed bimodal seasonal incidence of captures of 104 Leptodeira at Iquitos, Peru; peaks were in March and August. Life History. — One female collected in May contained six ovarian eggs. Three eggs from a clutch found on 5 April hatched on 10 July. The hatchlings had body lengths of 142, 145, and 146 mm. The eggs were part of a group of 15 found in a cavity of a bamboo about 12 m above the ground. No obvious differ- ences in size or shape were noted, so it was a surprise when some hatched into Leptodeira and others into Leptophis ahuetulla. The limited data on reproduc- tion at Santa Cecilia does not refute Duellman's (1958:115) supposition that Leptodeira annulata breeds throughout the year at Iquitos, Peru. Food. — Stomach contents revealed only anuran remains: Hyla marmorata (2), Hyla lanciformis (1), Hyla parvi- ceps (1), Eleuthcrodactylus hrevicrus (1), Hyla hokermanni (1), and Phyllo- mcdusa eggs (3). Remarks. — The relative abundance of Leptodeira in primary and secondary forests may be more apparent than real. Most of the snakes were found at sites EQUATORIAL HERPETOFAUNA 249 Fig. 154. — Leptophis ahuetuUa nigromarginatus, 9, 896 mm Pastaza, Ecuador (KU 107041). SVL; Limoncocha, Provincia where frogs were breeding. Those sites that were worked most intensively were in secondary forest. Leptophis ahuetulla nigromarginatus (Giinther) (Figure 154) Material. — Santa Ceciha, 15. Identificafion. — Leptophis ahuetuUa is the only long, slender snake in the area that is primarily green and has smooth dorsal scales in 15 rows at midbody. Other colubrids that are green (Leima- dophis reginae) or have green stripes (Oxyhelis argenteus) have dorsal scales in 17 rows. The various species of Chi- ronius can be distinguished from Lep- tophis by having dorsal scales in 10 or 12 rows. The dorsum is bright green; each of the scales on the body and plates on the head is margined with black. The first three scale rows are bronze or copper colored. The venter is metallic salmon. The iris is yellow, and the tongue is bluish green with a black tip. The oblique scale rows and diminish- ing size of the scales dorsolaterally give a false impression of an enlarged verte- bral scale row. In some individuals the dorsal scales on the posterior part of the body are weakly keeled. S 662-|-433 mm; 5 620+398 mm. Occurrence. — We have limited eco- logical data from Santa Cecilia, but ob- servations there and elsewhere reveal that L. ahuetuUa is diurnal, and arboreal, mostly in low, dense vegetation in pri- mary and secondary forest. Life History. — A female collected in April contained one egg 16.2 mm in length, and a female from July contained three eggs 18.1 mm in length. Fifteen eggs were found in a cavity of a bamboo about 12 m above the ground on 5 April. Four of these hatched between 10 and 17 May; four others hatched between 26 June and 1 July. Three of the entire clutch hatched into Leptodeira. The remaining eggs spoiled. Apparently the composite nest contained not only eggs of two species but also clutches of Lep- tophis eggs laid more than a month apart. Body length of eight hatchlings is 142-146 {k=l57.5) mm, and tail length of 92-105 (x:=98.4) mm. The limited reproductive data from Santa Cecilia supports Oliver's (1947:5) contention that this species reproduces throughout the year at Iquitos, Peru. Food. — One individual had eaten two Hyla alb 0 guttata. Taxonomic Comments. — Juveniles and some adults have extensive coppery coloration on the body; thus, they re- semble Leptophis cupreus (Cope), as defined by Peters and Orces-V (1960). Examination of 18 specimens from pro- vincia Napo revealed the following vari- ation: ventrals, $ S 147-155 (x= 149.9, N=7), 9 9 149-157 (x=153.3, N=:ll); subcaudals $ $ 144-154 (x=148.8, N= 7), 9 9 137-159 (x=148.9, N=8); cau- dal reduction from six to four rows at level of subcaudal 2-6 (x=5.0, N=18); 250 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 155.— Liophis cohella, $,522 mm SVL (KU 126043). upper labials 8-9 (x=8.5, N=18); pre- ocular in contact with frontal in two of 18 specimens. In all of these characters the sample from Santa Cecilia is much more like L. ahuetuUa than L. ciipreus (see Peters and Orces-V, 1960, for diag- noses ) . Liophis cobella (Linnaeus) (FiGxmE 155) Material. — Santa Cecilia, 2. Identification. — Liophis cobella is a predominantely tan snake with dark brown dorsal blotches, some of which extend across the venter. It is most eas- ily confused with Atractus major which has a single anal plate (divided in Lio- phis) and dorsal blotches that do not extend across the venter. Other similarly patterned snakes are the species of HeJi- cops, and Drymobius, which have keeled scales in 19 or 21 rows at midbody. Other snakes having smooth dorsal scales in 17 rows and a divided anal are Rhadi- naea and Leimadophis, neither of which has large dorsal blotches extending across the venter. The dorsal ground color is tan; about 40 dorsal dark brown blotches are pres- ent on the body; most of these alternate but connect with large lateral blotches which extend as black bars one or two ventrals wide across the belly. The lips are grayish brown; the chin and throat are cream. The belly is tan with a suf- fusion of salmon, and the underside of the tail is cream. The iris is reddish brown, and the iris is red with a black tip. 9 525+109 mm. Occurrence. — One was on the ground in a clearing at night and one in a water- filled, grassy depression in a clearing at night. Life History. — A female from April contained six oviducal eggs 25.1 mm in length. Ninia hudsoni Parker (Figure 156) Material. — Puerto Libre, 1. Identification. — This small snake is the only species in the area having dor- sal scales keeled and in 21 rows through- out the length of the body. The much larger Pseiistes sidphureus has 21 rows at midbody; the number of rows is re- duced to 13 or 15 anterior to the anus. In coloration, Ninia hudsoni is unique in having a dark gray dorsum with a pale gray nape band and a creamy white ven- ter. The pattern of some Atractus occipi- toalbus is similar, but in these the belly also is dark gray. Furthemiore, A. oc- cipitoalbus has smooth dorsal scales in 15 rows. ? 326+83 mm. Occurrence. — The single specimen was beneath a fallen palm frond in pri- mary forest by day. Life History. — The single female col- EQUATORIAL HERPETOFAUNA 251 Fig. 156.— Ninia hudsoni, ? , 409 mm SVL (KU 121913). lected in July contained two oviducal eggs 20.6 mm in length. Remarks. — As in the case of many other secretive snakes, it is unknown whether Ninia hudsoni is diurnal or noc- turnal. In reference to Ninia sehae sehae (Dumeril and Bibron) in Alta Verapaz, Guatemala, Stuart (1948:77) stated: "This snake seems to pass its entire life in concealment, and it was only occa- sionally that it could be found on the surface of the ground even at night." Oxybelis argenteus (Daudin) (Figure 157) Material. — Santa Cecilia, 50. Identification. — This elongate, slen- der snake with a round body and long, pointed snout is easily distinguished from other snakes in the area by its shape and coloration - dull green and tan stripes dorsally and green and white stripes ven- trally. No other snakes are so colored. Leptophis ahuetuUa also is slender and round, but it is predominantly green dor- sally and has dorsal scales in 15 rows ( 17 in Oxybelis). The various nocturnal tree snakes, (Dipsas, Imantodes, Leptodeira, Tripanurgos) with slender bodies have laterally compressed bodies, blunt heads, and blotched dorsal patterns. The top of the head is brown, and the dorsum of the body is tan with three dull green stripes. The throat, lower la- bials, and anterior part of the belly are pale lime green with brown or black flecks. The rest of the venter is white with two green stripes, which are dark- est posteriorly. The iris is pale yellow with a median brown streak. The tongue is olive-brown above and yellow below. ^ 774+508 mm; ? 787+548 mm. Occurrence. — This is the most abun- dant snake in our collections. Specimens were obtained every month except De- cember; five or more were collected in Februaiy, March, June, July, and Au- 252 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 157. — Oxtjhelis argenteus, juvenile, 412 mm SVL (KU 112280). gust. Only three individuals were found by day - in grassy clearing, on low bush, and on vine-covered tree trunk. All others were sleeping on bushes and trees at night. Most of these (77%) were on vegetation less than 1.5 m above the ground. Although some sleeping snakes were loosely coiled on leaves or draped on branches or vines, one sleeping pos- ture was most often seen. The snakes coil rather tightly around a small vertical branch or twig rising from a horizontal branch. Approximately 80 percent of Oxy])elis were found in second growth forest; the others were in primary forest (12%) or forest edge (8%). Apparently with increased human disturbance resulting in more secondary type forest the species is increasing in abundance. We have ecological data and dates of collection for 41 Oxy])elis from Santa Cecilia. The number of specimens obtained in each year is shown as the number in primary forest: secon- dary forest: forest edge - 1966 (0:0:1), 1967 (2:0:1), 1968 (4:0:0), 1969 (0:0: 1), 1971 (0:8:1), 1972 (0:17:0), 1973 (0:4:0). Life History. — One female from July contained four oviducal eggs 33.4 mm in length, and one from August contained three maturing ovarian eggs 23.7 mm in length. In the second week of July a female deposited six adherent eggs ( 12- 14, x=12.8 mm in diameter; 31-36, x= 32.7 mm in length). The smallest juve- niles, having body lengths of 143 and 191 mm, were found in April and October, respectively. Food. — The stomachs of 10 individ- uals contained food representing two species of lizards and three of frogs. The most abundant food items were Priono- (hictylus argulus and Eleuthcrodactylus varia])ilis (3 each). Also eaten were Anoli.s truchy derma (2), Dendrophyrnis- cus minutus (1), and Eleutherodactyhis pseud oacuminatus ( 1 ) . Although Pri- onodactylus ar^uhis is primarily a leaf- litter inhabitant, individuals do ascend bushes, where some fall prey to Oxy- helis. Anolis trachyderma and the two species of Eleutherodactyhis are bush in- habitants. Although the Eleutherodac- tyhis are primarily nocturnal, occasional individuals were active after diurnal rains, and E. variabilis calls from shaded areas in the late afternoon. Per- haps Oxyhelis finds sleeping individuals by day. Dendrophryniscus lives on the forest Hoor; the presence of that species in the stomach of an Oxyhelis is indica- EQUATORIAL HERPETOFAUNA 253 tive that these arboreal snakes occasion- ally feed on the gronnd. Also, perhaps some of the Prionodactylus were eaten on the ground. Oxyrhopus formosus (Weid) Material. — Lago Agrio, 1; Puerto Libre, 1; Santa Cecilia, 3. Identification. — Among the several species of colubrid snakes in the area that have a black and white ( or yellow ) head and red body, at least in some stage of growth, Oxyrhopus formosus is unique in having banded juveniles and unicolor adults. In juveniles ( < 250 mm body length), the tip of the snout is cream, the top of the head is tan, and the head posterior to the eyes and the nape are creamy yellow. The body and tail are creamy yellow with 17-20 broad black blotches that are slightly narrower laterally than middorsally and extend onto the edges of the ventrals and are complete across the ventral surface of the tail. The blotches are 5-10 scales in length; the intervening cream areas are 2-3 scales in length. Half-grown indi- viduals (400-500 mm body length) have a cream snout including the entire area anterior to the eyes; the top of the head is brown, and the cream nape band is suffused with brown. The posterior tips of all dorsal scales are dark or black. The blotches are dull reddish brown, and the interspaces are pinkish cream. In adults ( > 620 mm in body length ) the snout anterior to the eyes and the upper la- bials are creamy yellow; the rest of the head and the nape are dark brown or black. The dorsum of the body and tail is pale salmon red; the tips of the scales are black. The venter in juveniles is cream; in adults it is pinkish salmon. Juveniles are easily confused with Oxyrhopus petolo, but in that species the snout is black and the body blotches are longer (8-12 scales). Furthermore, O. petoJa has more ventrals and subcaudals (Table 9) and does not have the preocu- lar in contact with the frontal. The col- oration of adult O. formosus resembles that of Clelia clelia, Drepanoides anoma- lus and Pseudo])oa coronata. CleJia has a black snout and more ventrals and sub- caudals (Table 9). Drepanoides has dor- sal scales in 15 rows and Pseudohoa in 17 rows; both have black snouts and broad cream bands across the head pos- terior to the eyes. One specimen (KU 126017) has 21 rows of scales at midbody; all others have 19 rows. The iris is reddish tan to rusty red, and the tongue is black. S 621+198 mm; 9 830+197 mm. Occurrence. — One was moving on the ground in primary forest at night, and one was obtained during clearing of primary forest by day. Life History. — Two females from May having body lengths of 778 and 830 mm contained 17 and 13 matin-ing ova, respectively. Food. — One individual contained a diurnal, teiid lizard Arthrosaura reticu- lata. Oxyrhopus melanogenys (Tschudi) Material. — Santa Cecilia, 12. Identification . — This predominantly black or dark gray snake has smooth dor- sal scales in 19 rows at midbody. The head is black bordered posteriorly by an orange or red nape band. The body pat- tern consists of broad (6-13 scales), black l^ands that extend well onto the ventrals, especially posteriorly. The bands are separated anteriorly by nar- row (1-2 scales), sometimes incomplete, white or pale yellow interspaces. On the posterior half of the body the pattern is tricolor : black-yellow-black-red-black- yellow-black. The red bands vary from orange to dull red and are 3-5 scales wide. The red bands are suffused with black; in large adults the bordering black bands meet middorsally, thereby restrict- ing the red band to a pair of lateral spots. One or two red bands are present proxi- mally on the tail, which is black pos- teriorly. The chin and belly are dull 254 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY cream; the underside of the tail is black. In one individual the normally white in- terspaces are lime green. The iris is red- dish brown, and the tongue is dark gray with a white tip. $ 629+187 mm; $ 819+199 mm. The only other Oxijrhopus likely to be confused with O. mcJano^cniis is O. petoki. However, that species has black and red (yellow in juveniles) bands throughout the length of the body and more ventrals and subcaudals (Table 9). Triads of black, red, and vellow rings occur in Afractiis elaps, EnjthroJamprus aesciilapii, and some species of Micrurus, but in all of these the pattern is present throughout the length of the body, and the rings are continuous across the ven- ter. Occurrence. — Five were active on the ground in clearings by day; two were moving on the ground in clearings at night. One was coiled in the crotch of a small tree at night, and another was under a log in a clearing by day. These observations and the nature of the food found in the stomachs indicate that this species is primarily diurnal. Life History. — One female from April with a body length of 819 mm contained 12 small ova. Food. — Stomach contents consisted of one small rodent and three diurnal teiid lizards, Arthrosaura reticulata., Iplii- sa elegans, and Lepsoma parietale. Oxyrhopus petola digitalis (Reuss) Material. — Puerto Libre, 1; Santa Ce- ciha, 20. Identification. — This snake has smooth dorsal scales in 19 rows and a pattern of broad black blotches on the dorsum of the body and tail. The snout and head are black; there are 12-17 (x=15.1, iV=: 18) black bands 8-12 scales wide on the body. The bands extend onto the edges of the ventrals and are separated by pale interspaces 2-3 scales wide middorsally and 4-5 scales wide laterally. In most individuals, some black bands are offset at the midline, especially at midbody. The chin, lower labials (but not upper labials), ventrals, and subcaudals are creamy yellow. In juveniles ( < 300 mm body length) the nape band and inter- spaces are pale yellow. The interspaces become orange and then red progressing anteriorly in larger individuals. Snakes having body lengths of 350-450 mm have red interspaces posteriorly, orange ones at midbody and anteriorly, and a yellow nape band. Snakes with body lengths less than 525 mm have red interspaces and a yellow nape band; in larger indi- viduals the nape band is red. The red is darkest in larger individuals. In juve- niles the ventral surface of the tail is immaculate, but in larger individuals an irregular, mid ventral black line is pres- ent. The iris is brown, and the tongue is gray. $ 622+218 mm; 9 557+211 mm. This snake can be confused with ju- veniles of Oxijrophus forniosus, but in that species the snout is cream, the black bands are shorter (5-10 scales), and the interspaces are creamy yellow. Further- more, O. formo.sus has fewer ventrals and subcaudals (Table 9) and does not have the preocular in contact with the frontal. Oxyrhopus petola can be dis- tinguished readily from O. melanogenys in that the latter has narrow ( 1-2 scales) yellow interspaces separating broad (6- 13 scales) black bands anteriorly and a tricolor pattern posteriorly: black-yel- low-black-red-black-yellow-black. More- over, O. melanogenys has fewer ventrals and subcaudals (Table 9). Occurrence. — Five individuals are from forest, the others from clearings. Five were moving at night; one was coiled in a bush in a swamp and another on a low bush in forest at night. Others were active on the ground l:)y day. Ap- parently this species is primarily noc- turnal. Life History. — No data are available from Santa Cecilia. Fitch (1970:154) re- ported seven gravid females from Iqui- tos, Peru, having 5-10 (x=7.3) eggs col- EQUATORIAL HERPETOFAUNA 255 lected in March, September, and Novem- ber. Food. — Two individuals contained diurnal, terrestrial teiid lizards, Kcntro- pyx peJviceps and Prionodactylus inani- catus. Pseudoboa coronata Schneider Material. — Santa Cecilia, 7. Identification. — This red snake with a black and cream head differs from other similarly colored snakes by having single, instead of divided, subcaudals. Other snakes in the area with similar color patterns are Clelia clelia, Oxyrho- pus formosus, and Drepanoides anoma- lus. The latter has dorsal scales in 15 rows, and the others have 19 rows, whereas Pseudoboa has 17 rows. The anterior part of the head, includ- ing the eyes, is black; the rest of the head and anterior part of the neck are creamy white. A diffuse back band separates the creamy white neck from the red body. The tips of the dorsal scales are black, more so in adults than in juveniles. The venter is cream. 9 816+245 mm. Occurrence. — One was in water in a muddy depression in secondary forest at night. Two were active on the ground in clearings by day and one at night. The stomach contents suggest that this spe- cies may be primarily diurnal. Life History. — One female collected in September with a body length of 761 mm contained four oviducal eggs 34.2 mm in length. Food. — Two individuals contained diurnal, teiid lizards, Ameiva ameiva, and one contained remains of two juve- nile rodents. Pseustes sulphureus sulphureus (Wagler) Material. — Santa Cecilia, 1. Identification. — This large snake has keeled dorsal scales in 21 oblique rows at midbody, that are reduced to 13 rows anterior to the anus, and a single anal plate. Two other snakes in the area have keeled dorsal scales in 21 rows at mid- body. ISJinia hiidsoni is dark gray with a pale band on the head, and Helicops petersi has fused internasals and about 40 middorsal dark blotches alternating with lateral blotches. Pseustes has paired internasals and no blotches anteriorly on the body and indistinct transverse blotches posteriorly. Xenodon severus also has 21 oblique rows of dorsals at midbodv, but in Xenodon the scales are smooth, and the anal plate is divided. The dorsum is dull olive-green, and the skin between the scales is yellow anteriorly and gray posteriorly. Trans- verse marks on the posterior part of the body are black. The chin and throat are bright lemon yellow tinged with olive. The anterior fourth of the belly is dull olive tinged with yellow, and the rest of the venter is gray tinged with olive. The iris is dull brown, and the tongue is pink with an orange tip. <^ 1328+455 mm. Occurrence. — The single specimen was brought in by an Indian, who said he found it on the forest floor by day. Life History. — No data are available. Food. — The specimen contained a rodent. Rhadinaea brevirostris (Peters) ( Figure 158 ) Material — Lago Agrio, 1; Santa Ce- cilia, 7. Identification. — The presence of smooth dorsal scales in 17 rows at mid- body and a divided anal plate distin- guish Rhadinaea from all other snakes in the area, except Leimadophis and Lio- pJiis. Rhadinaea has dark and light brown dorsal stripes and an immaculate belly. Leimadophis is tan or brown with or without lateral stripes posteriorly and with black spots on the belly. Liophis has dark brown bands complete around the body. The coloration of Rhadinaea is similar to that of TantiUa melanoceph- ala, which lacks a loreal and has a nar- row ( 1 scale row wide) middorsal brown stripe. Rhadinaea has a loreal and a middorsal brown stripe three scales in width. 256 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 158.— FUiadinaea brevirostris, 9, 419 mm SVL (KU 1 19391). The head is dark brown with reddish tan spots; the upper hibials are creamy tan with brown flecks. The dorsum of the body is pale brown or grayish tan. Five dark brown stripes extend the length of the body - lateral pair on edges of ventrals, dorsolateral pair on lower edge of fifth scale row, and dorsal on vertebral and paravertebral rows. In some individuals the dorsolateral stripe is narrowly bordered above by a nar- row creamy tan stripe. The chin is red- dish tan to cream, and the belly is creamy white (pale green in one of five adult females). The iris is pale brown with a narrow red ring around the pupil, c^ 300+91 mm; 5 269+86 mm. Occurrence. — This small, diurnal snake was found in a variety of terres- trial situations - primary and secondary forest, clearing, and banana grove. Three are from May, two from August, and one each from June, July, and Novem- ber. Life History. — One female collected in May contained two oviducal eggs 26.2 mm in length. Food. — One individual contained a teiid lizard, Prionodactylus maniccitus. Siphlophis cervinus (Laurenti) Materia]. — Santa Cecilia, 2. Identification. — Smooth dorsal scales in 19 rows at midbody, a single anal plate, and two anterior temporals dis- tinguish Siphlophis from all other snakes in the area. Furthermore, the species has a distinctive color pattern. The sides of the body are pale yellow with 75-80 ir- regular, vertical black marks, each slight- ly wider than adjacent interspaces. A diffuse, orange-red middorsal stripe ex- tends the length of the body. The top of the head and the iris are brown. The throat is pale gray, and the rest of the venter is pale yellow. The tongue is pale pink, s 551+188 mm. Occurrence. — One individual of this nocturnal snake was under a rock in a clearing by day; the other specimen lacks ecological data. Life History. — No data are available. Food. — One had eaten a teiid lizard, Bachia trinasale. Tantilla melanocephala melanochephala (Linnaeus) (Figure 159) Material. — Lago Agrio, 3; Santa Ce- cilia, 2. EQUATORIAL HERPETOFAUNA 257 Fig. 159. — Tantilla melanocephala melanocephala, $, 309 mm SVL (KU 109848). Identification. — This small brown snake has smooth dorsal scales in 15 rows throughout the length of the body. The anal plate is divided, and the loreal is absent. The only other snake in the area with a divided anal plate, no loreal, and smooth dorsals in 15 rows is Lepto- phis ahuetuUa, a long green snake with only 11 or 13 rows of scales posteriorly. Tantilla superficially resembles Rhadi- naea hrevirostris, which has 17 rows of scales at midbody, a loreal, and a broad (3 scales in width) dark brown middor- sal stripe. In Tantilla the middorsal stripe is only one scale wide. Tantilla with dull patterns might be confused with the brown phase of Atractus occipi- toallms which has a single anal, no mid- dorsal dark stripe and has a midventral row of dark spots. The top of the head is brown. A broad dark brown nape band is bordered laterally by cream spots. A narrow ( 1 scale wide) dark brown stripe extends from the nape band to the tip of the tail. A dark brown lateral stripe on dorsal scale rows 1 and 2 is bordered above on the anterior part of the body by a nar- row tan stripe; the rest of the dorsum is brown. The side of the head is dark brown with white labial spots anterior and posterior to the orbit. The lower labials are dark brown and white. The chin and throat are white, and the belly and underside of the tail are pale green or greenish cream. The iris is orange-tan above and grayish brown below; the tongue is brown with a white tip. $ 253+82 mm; $ 207+16+ mm (incom- plete ) . Occurrence. — All specimens were ac- tive on the ground or amidst leaf litter in primary forest by day. Three were ob- tained in May and one each in March and August. Life History. — The single female col- lected in May showed no evidence of re- productive activity. Taxonomic Comments. — Schmidt and Walker (1934b:31S) referred popula- tions of Tantilla melanocephala from coastal Peru and the upper Maranon val- ley of Peru to T. melanocephala capi- strata Cope and distinguished the two subspecies by the contact of the pre- frontal with the second upper labial in capistrata and no contact in melanoceph- ala. Peters and Orejas-Miranda (1970: 295) noted that in capistrata the tip of the snout is white and in melanocephala, brown. All of the specimens from Santa Cecilia and Lago Agrio have brown snouts; in two, the prefrontals are in contact with the upper labials on both 258 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 160.— Tripanurgos compressus, $,582 mm SVL (KU 148395). sides, and in one, there is contact only on one side, whereas in the others the prefrontal is separated from the upper labials. Tripanurgos compressus (Daudin) (FlGUl^E 160) Material. — Lago Agrio, 1; Santa Ce- cilia, 1. Identification. — This is one of two snakes in the area with dorsal scales in 19 rows and an enlarged vertebral scale row. The other is Leptodeira annulata, which differs by having a divided anal plate (single in Tripanurgos), one an- terior temporal (two), and broad mid- dorsal brown blotches on the body ( nar- row transverse blotches). Other snakes having laterally compressed bodies and enlarged vertebral scale rows have fewer rows of dorsal scales: Imantodes cen- choa (17), Imantodes lentiferus (15), and Dipsas ( 13 ) . The head is dark brown bordered posteriorly by a pale yellow nape band and a dark brown band. The dorsum of the body is reddish brown with about 45 narrow dark brown marks one or two scales in length and extending laterally to the first or second scale row. The venter is creamy tan. The iris is burnt orange. 9 612+87 mm. Occurrence. — One specimen was re- trieved from a large tree when it was felled in primary forest by day in May. The other was crawling in low vegetation in secondary forest at night in March. Life History. — No data are available. Food. — One individual contained the remains of a microteiid lizard. Xenodon severus (Linnaeus) (Figure 161) Material. — Puerto Libre, 2; Santa Ce- cilia, 6. Identification. — Xenodon severus is the only snake in the area having smooth dorsal scales in 21 oblique rows. The body is heavy and depressed; the anal plate is single, and one anterior temporal is present. Other snakes in the area with 21 rows of scales at midbody (Helicops, Ninia, and Pseustes) all have keeled scales, except that some specimens of Oxtjrhopus formosus have 21 rows of smooth scales. Oxyrhopus formosus is a EQUATORIAL HERPETOFAUNA 259 Fig. 161.— Xenodon severus, $ , 880 mm SVL (KU 121927). slender, round-bodied snake with two anterior temporals, dorsals in diagonal rows, and a color pattern entirely differ- ent from that of Xenodon. Ontogenetic change in coloration re- sults in a nearly unifomily colored adult from a banded juvenile. Small juveniles ( < 300 mm body length ) have a pattern of diagonal dark brown and grayish white markings on the head. The tip of the snout is brown, outlined in white. A brown bar is continuous across the inter- nasals and thence posteriorly through the orbit to the ventrolateral surface of the neck; it is bordered by white bars, the upper of which continues posteriorly on the side of the neck. A dark brown interorbital bar is narrowly connected middorsally on the frontal to the anterior end of a large ovoid brown blotch that continues onto the anterior part of the body. On the nape the middle of the blotch has a large tan spot. The dorsum of the body is tan with 6-8 brown blotches outlined in black. The blotches are 10-14 scales in length and extend laterally onto the first scale row. The interspaces are tan and 3-4 scales in length. The venter is dark brown with creamy yellow spots laterally. In larger individuals ( < 700 mm ) , the markings become diffuse, and the dorsum is olive-tan or yellowish tan. The ovoid spot with a pale center on the head and neck persists; it is especially evident when the body is flattened, and the neck is spread. In large adults the body is dark gray or black with yellowish tan scales foiming incomplete, irregular transverse lines posteriorly in some speci- mens. The head is brown, and the lower labials and throat are yellow. The belly is creamy yellow to orange-tan. The iris is brown, and the tongue is flesh-colored with a gray tip. S 807+139 mm; ? 1060+152 mm. Occurrence. — This diurnal, terrestrial snake inhabits both primary and secon- 260 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY dary forest. Two were obtained in each of May, June, and July; one was obtained each in February and August. Life History. — One female collected in May contained 22 ova. Fitch (1970: 173) noted that the species "seems to have year-round breeding in the tropical equatorial lowlands." Food. — Xenodon serverus is a pro- digious frog eater. Two individuals con- tained Bufo marinus, and two contained Leptodactylus pentadactyhis. One indi- vidual (body lentgh 1050 mm) was found in the process of eating a L. pento- dactylus having a snout-vent length of 173 mm; subsequent examination re- vealed one adult L. pentadactylus al- ready in the stomach. Just how Xenodon ferrets out large frogs by day is not known. Neither the feeding individual, nor others containing food, was taken immediately after heavy rains. Xenopholis scalaris (Wucherer) Material. — Lago Agrio, 1; Puerto Libre, 1; Santa Cecilia, 1. Identification. — This small snake is distinctive in having the prefrontals fused into a single large shield. The dorsal scales are smooth and in 17 rows. The dorsum is reddish brown becoming pinkish orange or orange-tan laterally. There are 24-30 narrow black crossbars on the body; these extend laterally onto the third or fourth scale row. In some in- dividuals the bars are offset middorsally and/or connected by a narrow middor- sal line. The crossbars are 1-2/2 scales wide. Small black flecks are present on the first and second rows of dorsal scales. The ventral surfaces are immaculate creamy white. The iris is dark reddish brown, and the tongne is pink, c^ 244-|- 54 mm; 9 317+56 mm. Occurrence. — All were found by day. One was moving in a clearing; two were in primary forest - one under a palm frond and one in leaf litter. Life History. — A female obtained in May contained three oviducal eggs 23.9 mm long. Food. — One individual contained a terrestrial frog, Ischnocnema quixensis. Elapidae Leptomicrurus narduccii (Jan) (Figure 162) Material. — Lago Agrio, 1; Santa Ce- cilia, 4. Identification. — This long, slender snake has a round body, blunt head, and short tail. It is unifonnly black above except for a bright yellow band around the head. Ventrally it is black with large yellow spots; the spot (or spots) on the ventral surface of the tail is orange or red. In freshly preserved specimens, the spots on the belly are red. Only two other snakes in the area are dark above and have a pale band on the head or neck. Of these, Atractus occipitoalbus is like Leptomicrurus in having smooth dorsal scales in 15 rows, but in Atractus the venter is uniform gray and the band on the occiput is dull orange to pinkish orange. The other species is Ninia Jitid- soni, which has keeled dorsal scales in 21 rows, a pale gray band across the neck, and a creamy white venter. $ 893+43 mm; 9 798+34 mm. Occurrence. — One was beneath a log in primary forest, and another was plowed up by a bulldozer during the clearing of matiu'e forest. Life History. — One female collected in May contained small ovarian eggs. Remarks. — When disturbed, one in- dividual spiraled the tail dorsally, re- vealing the ventral red markings. Micrurus langsdorffi langsdorffi Wagler ( Plate 4 ) Material. — Lago Agrio, 1; Santa Ce- cilia, 1. Identification. — This distinctive coral snake differs from other Micrurus and coral snake mimics in the area by having numerous (36-38) narrow black rings EQUATORIAL HERPETOFAUNA 261 Fig. 162. — Leptomocnirus narcluccii, 9 , 605 mmSVL (KU 126057). not arranged in triads, and bordered by narrow white rings. The red rings are not more than five scales wide. The snout and top of the head are black. The other species of Micrurus have black rings in triads: black-yellow-black-red- black-yellow-black. In M. lang^sdorjfi the rings are black-white-red-white- black. Some Atractus ekips have a pat- tern like that of M. langsdorffi, but the former has much wider red rings, fewer than 20 black rings, single anal plate, and only one pair of chinshields; M. langsdorffi has a divided anal plate and two pairs of chinshields. Anilius scytale, which is red with about 50 black rings, differs by having the eye beneath a single scale, only five enlarged head shields, and ventral scutes only slightly wider than the dorsal scales, whereas M. langsdorffi has the eye surrounded by several scales, seven enlarged dorsal head shields, and enlarged ventral scutes. The white rings usually are no more than one-half scale in width. The red rings are dark red, and the scales in the red rings are tipped with black. The chin is white. S 483+91 mm; 9 676+ 65 mm. Occurrence. — One individual was crawling on the ground in a clearing by day; another was killed by natives by day. Life History. — A female obtained in May contained small ovarian eggs. Micnuais lemniscatus helleri Schmidt and Schmidt ( Plate 4 ) Material. — Lago Agrio, 1; Santa Ce- cilia, 2. Identification. — Three species of Mi- crurus in the area have a pattern of rings arranged in triads: black-yellow-black- red-black-yellow-black. Of these, M. lemniscatus is the only species having a ringed pattern on the head; the snout is black, followed successively by yellow, black, and yellow rings, and a black nape. In M. spixii the head and nape are black; in M. surinamensis the head scutes are black and red. Erythrolamprus aesculapii has a ringed pattern on the head; the snout is yellow followed by a black ring, a yellow ring, and a black nape. In Erythrolamprus the yellow ring is posterior to the eyes, whereas in M. lemniscatus the yellow ring is anterior to the eyes. Some individuals of Atractus claps have rings in triads, but this species has a black head with or without a creamy yellow ring posteriorly on the head and red nape. The red rings (orange-red in life) and black rings are about equal in width (4-6 scales); the creamy yellow rings are one or two scales in width. The rings on the head is pale yellow. Few scales in the red rings have black tips. 6 1035+ 110 mm. Occurrence. — Two were in primary forest and one in secondary forest. One was found in a trench in early evening, so the snake might have fallen in the trench by day; the other individuals were active on the forest floor by day. Life History.— No data are available. Micrurus spixii obscurus Jan ( Plate 4 ) Material. — Santa Cecilia, 7. Identification.— Oi the three species 262 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY of Micrurus having rings in triads (black- yellow -black -red -black -yellow -black ) , M. spixii is distinctive because the yellow rings are as wide as adjacent black rings (5-6 scales). Micrurus lemniscatus and M. surimimensis have yellow rings much narrower than the black rings. Further- more, in M. spixii the top of the head is black, whereas M. lemniscatus has a yel- low ring on the head, and M. surinamen- sis has black and red scales on the top of the head. Other snakes in the area hav- ing red, black, and yellow rings and dor- sal scales in 15 rows include Erythro- lamprus aesculapii and some Atroctus elaps. The foniier has a yellow snout and black and yellow rings on the head. Atratcus elaps has a single anal plate and one pair of chinshields, whereas M. spixii has a divided anal plate and two pairs of chinshields. The red rings are coral red in life, and the dorsal scales in the red and pale yellow rings are suffused with black, especially middorsally. A red mark pos- terior to the eye includes the posterior labials and temporals. S 1299+70; $ 778+43 mm. Occurrence. — One was on the ground at the edge of the forest by day. All others were brought in by natives who indicated that the snakes were found on the forest floor by day. Life History. — A female obtained in July contained small ovarian eggs. Micrurus surinamensis surinamensis (Cuvier) ( Plate 4 ) Material. — Santa Cecilia, 1. Identification. — This coral snake dif- fers from other Micrurus and coral snake mimics by having extremely small eyes directed anterolaterally, a small frontal, fourth upper labial barely entering the ■Orbit, and huge anterior and posterior temporals. The head is black with dull red on the anterior edges of the scales. A narrow creamy yellow ring separates the head and body patterns. The latter consists of six triads: black-yellow-black- red-black-yellow-black. The outermost black rings are much wider than the others (9-10 scales middorsally); the next widest are the red rings (6-7 scales ) , innennost black rings ( 4 scales ) , and the yellow rings (1-1^2 scales). The yellow rings are expanded ventrally at the expense of the outennost black rings to include three or four ventrals. The tips of the dorsal scales in the red rings are black. c5 1115+147+ mm (incom- plete ) . Occurrence. — The single specimen was brought in by an Indian, who re- ferred to it as culehra del agua (water snake). A snake, presumably the same species, was seen by day in a small stream in a clearing. Life History. — No data are available. Food. — According to Terry D. Schwaner (pers. com.), this specimen contained fish vertebrae in its stomach. Schmidt (1952:32) reported a small eel, Synbranchus marmoratus, in the stomach of an individual from Iquitos, Peru. These food items, together with the ob- servation at Santa Cecilia and the de- pressed head with modified cephalic scutellation, are indicative of aquatic habits. In fact, this species of Micrurus is like Hydrops triangularis in scutella- tion and habitus. Taxonomic Comments. — The present specimen (AUM 8636) is notably larger than the maximum of 1105 mm total length reported by Schmidt (1952:31). Crotalidae Bothrops atrox (Linnaeus) (Figure 163) Material. — Dureno, 2; Lago Agrio, 1; Puerto Libre, 3; Santa Cecilia, 30. Identification. — This is the most vari- able pit viper in the area. It differs from Lachesis muta in having 25, instead of 35, rows of dorsal scales at midbody and two, instead of four or five, rows of scales between the labials and the orbit. The EQUATORIAL HERPETOFAUNA 263 Fig. 163.— Bothrops atrox, juvenile, 470 mm SVL (KU 121935). dorsal color pattern consists of irregular dark brown blotches on a tan or brown ground color. The other two species of Bothrops in the area differ markedly in coloration; B. hilineatus is green with a pale ventrolateral stripe, and B. castel- naiidi is greenish yellow with chocolate brown blotches. In adults and juveniles, the dorsum is dull yellowish brown, grayish brown, or brown with darker brown or black ir- regular middorsal blotches and smaller lateral blotches. In juveniles the venter is white with black transverse blotches; the chin is mostly black with white dots. In larger individuals the white venter changes to creamy yellow, and the black ventral markings change to dark gray or brown without greatly increasing in size; thus, adults appear to have paler venters than do juveniles. In juveniles the la- bials are gray with two or three vertical black bars; in adults the labials are uni- formly yellowish tan. The tenninal half of the tail is dull yellow above and below in small juveniles ( < 455 mm total length) and yellow below in larger ju- veniles ( < 470 mm total length ) . In adults the venter of the tail is darker in males than in females. 6 1024-]- 181 mm; 9 1288+222 mm. Occurrence. — Bothrops atrox, locally called equis, because of the X-shaped dorsal markings, is the most abundant pit viper in the area. Only two were found in primary forest, whereas 11 were in secondary forest, one at forest edge, and four in clearings. These account for only 18 of the 36 specimens; others were captured by natives, usually around their homes or in their cultivated clearings. Eleven individuals were found on vegetation at night. Most of these were juveniles coiled on large leaves less than 1.5 m above the ground, but individuals having body lengths to 732 mm were 264 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY found in trees. One was crawling along a branch about 4 ni aboxe the ground, and another was coiled on a vine aljout 1 m above the ground. Individuals have been observed actively moving by day and night. Life History. — A female collected in July (1073 mm in body length) con- tained 18 embr\'Os, and one from Novem- ber ( 1197 mm in body length) contained 24 embryos. Juveniles having body lengths of 227-340 mm were found in February (1), May (2), June (1), July (4), and August ( 1 ). These limited data suggest that the reproductive season is lengthy or most likely extends through- out the year. Food. — Five juveniles contained food: two frogs, Leptodactylus mysta- ceiis, and two lizards, Amciva anieiva petersii and Leposoina paricfale. These items indicate that Botlirops atrox prob- ably feeds by night and day. Bothrops bilineatus smaragdinus Hoge (Figure 164) Material. — Santa Cecilia, 2. Identification. — This distinctive, green pit viper has a prehensile tail and there- by differs from all other pit vipers in the area. The only other heavy-bodied, ar- boreal green snake in the area is the boa, CoraUus caninus, which has smooth scales, sensory pits in the labials, and no sensory pit between nostril and eye. The dorsum is pale green with mi- nute black flecks; a pale yellow stripe occurs on the first row of dorsal scales. The edges of the ventrals are pale green, and the belly is white. The lips are yel- lowish green, and the iris is pale green. S 365+64 mm; ? 555+90 mm. Occurrence. — One male was crawling along a leafy vine 2 m ab()vc> the ground in primary forest at night. The other specimen was brought in by a native. Apparently this snake, locally called Joro (parrot), is more common than our few specimens indicate. At the time of our field work at Santa Cecilia in 196(S and Fig. 164. — Botlirops bilineatus smaragdiniis, $ , 429 mm SVL ( KU 112296). 1969, two trail cutters working for the oil company were bitten by Joroa. Life History. — A female obtained in February contained immature ovarian eggs. Bothrops castehiaudi Dumeril, Bibron and Dumeril Materia], — Santa Cecilia, 1. Identification. — This slender pit vi- per differs from other Bothrops in the area by its distinctive coloration. The dorsum is pale greenish gray with yellow flecks and square black blotches middor- sally and smaller black spots laterally; the ventral surfaces are dark brown with pale yellow flecks, especially anteriorly. Bothrops bilineatus differs by being green with a yellow ventrolateral stripe, and B. atrox has a blotched black and white or brown and cream venter and no yellow flecks dorsally. Lachesis niuta has dorsal scales in 35 rows (25 in B. castehiaudi), four or five rows of scales between the orbit and the labials (two rows in B. castehiaudi) and a tan dor- sum with dark brown marks. The single specimen from Santa Cecilia is a fe- male: 486+84 mm. Occurrence. — The single specimen, obtained by a native, has no ecological data. Life History. — No data are available. EQUATORIAL HERPETOFAUNA 265 Fig. 165. — Lachesis muta muta, Santa Cecilia ( Photo by Joseph T. CoHins ) . Lachesis muta muta (Linnaeus) (Figure 165) Material. — Santa Cecilia, 2. Identification. — The largest pit viper in the area differs from other pit vipers there by having knob-like keels on the middorsal scales, dorsal scales in 35 rows, and four or five rows of scales separating the orbit from the labials. Pit vipers of the genus Bothrops have straight, even keels on the dorsal scales, which are in 25 rows, and two rows of scales between the orbit and the labials. The body is a rich golden tan to pinkish tan with large dark brown or black, middorsal, diamond-shaped marks, usually narrowly bordered by cream. Vertical dark bars extend from the lat- eral corners of the blotches nearly to the ventrals. The top of the head is brown; a black postorbital stripe extends to the angle of the jaw. The lips are tan, and the venter is cream. 9 2030-(-180 mm. Occurrence. — The bushmaster ap- parently is uncommon in the area. One was on the ground in primary forest, and another was next to a log at the edge of the forest. Both were inactive when found by day. Life History. — One female having a body length of 2030 mm obtained in August contained 13 oviducal eggs, each about 29 mm in length. REPRODUCTIVE BIOLOGY The continued existence of a species in a community is dependent upon suc- cessful reproduction by the adults and survival of the eggs and young to sexual maturity. My purpose here is to review the accumulated data on reproduction in the herpetofauna at Santa Cecilia with respect to annual reproductive patterns, fecundity, and reproductive strategies. A cursory review of data summarized by Turner (1962), Tinkle, Wilbur, and Til- ley (1970), Fitch (1970), and Salthe and Mecham ( 1974 ) reveals the limited ex- tent of our knowledge of reproduction in tropical amphibians and reptiles. By far the most informative study is that by Crump (1974) on frogs at Santa Cecilia. The following analysis has four limi- tations: 1) The unevenness of sampling throughout the year precludes an accu- rate picture of annual reproductive pat- terns; thus, annual patterns are at least as broad as demonstrated by the data and in many cases probably much more so. 2) The data on caecilians, salaman- ders, tiu-tles and crocodilians are too few 266 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY to be useful; accordingly, these groups ( total of 15 species ) have been excluded from the analysis, which therefore in- cludes only frogs, lizards, and snakes. 3) Only the reproductive condition of females was studied; I have been con- cerned with fecundit\' and time of breed- ing, and I echo Fitch's (1970:6) state- ment: "When fecund females are pres- ent in the population there are usually males available to inseminate them." 4) The actual frequency of breeding of in- dividuals is unknown; therefore, the data pertain only to populations, and infer- ences to individual cycles must be ex- trapolated. ANNUAL REPRODUCTIVE PATTERNS Most studies of reproduction in am- phibians and reptiles have been con- ducted in seasonal environments, in which environmental cues, such as changes in temperature, photoperiod, or rainfall initiate breeding activity; in at least some species these changes can be correlated with hormonal activity and gonadal development, as well. In the essentially aseasonal environment of Santa Cecilia, we must search for other factors that influence patterns of repro- duction. These factors may he intrinsic, such as a cycle that was established in the evolutionary history of a species in a seasonal environment prior to its inva- sion of a more ecjuable environment, or differences in the time required for de- velopment of the ova. Extrinsic factors might include slight fluctuations in rain- fall, prolonged periods of heavy cloud cover or many consecutive days of in- tense sunlight; moonlight has a distinctly negative effect on the activity of noc- turnal species. In the following discussion anurans, lizards, and snakes are treated separately prior to a final synthesis. Anurans The amount of data on each of the 87 species of frogs and toads is highly variable, but the aggregate is much more extensive than that for the lizards and snakes. Crump (1974) presented an account of 74 species from Santa Ce- cilia based on data that she accumulated in 1971-73. I have augmented her data with additional information from pre- vious work at Santa Cecilia and the other study sites. Three criteria must be clarified be- fore discussing reproductive patterns in anurans. Mode of reproduction, as de- fined by Salthe and Duellman (1973), is a concept combining the site of de- velopment of the eggs with the mode of development; the latter is a complex of factors, including rate and duration of development, size and stage at hatching, and kind of parental care, if any. I rec- ognize 11 modes of reproduction amongst the anurans at Santa Cecilia. Crump (1974) recognized 10 modes; she combined my modes 4 and 5 into one category. The reproductive modes are defined in Table 10. The ovarian size factor (Duellman and Crump, 1974) is an index for comparing fecundity and egg size relative to adult body size by means of the formula CS(OD)/SVL, where CS=clutch size, OD=ovum di- ameter, and SVL^snout-vent length of females depositing clutches. In the pres- ent study, mature ovarian complement was substituted for clutch size. I have followed Crump (1974) in distinguish- ing different types of annual reproduc- tive patterns: Continuous. — Gravid females and/or juveniles found throughout the year. Opportunistic. — Species that breed regularly after heavy rains which occur throughout the year. Therefore, these species are somewhat more restricted in their breeding activity than are the continuous breeders. Sporadic Wet. — Species that breed sporadically after heavy rains; these spe- cies breed less frequently than the op- portunistic breeders. EQUATORIAL HERPETOFAUNA Table 10. — Distribution of Reproductive Modes among Species of Anurans. 267 Site of De%'elopment 0) OS n o 03 O H-1 oi 12 '2 c 3 OJ 12 O C 12 12 "2 0) U 0) '2 OS 0) o „ Eggs and larvae in water: A. Mode 1 — unconstrained body of water — . B. Mode 2 — tree cavity above ground — - C. Mode 3 — constructed nest — - Eggs out of water; larvae develop in water: D. Mode 4 — eggs on vegetation above pond —. E. Mode 5 — eggs on vegetation above stream F. Mode 6 — eggs in foam nest; larvae in water ... _ G. Mode 7 — eggs on land; larvae carried to water ..__ Neither eggs nor larvae unprotected in water: H. Mode 8 — eggs and larvae in terrestrial nest .... I. Mode 9 — terrestrial eggs with direct development .... J. Mode 10 — eggs undergoing direct development on back of aquatic female 1 K. Mode 11 — eggs undergoing direct development on back of terrestrial female 1 17 24 1 1 11 5 35 1 1 - 11 3 9 6 1 1 18 Sporadic Dry. — Species that breed sporadically during dry periods; these species breed least frequently of any of the anurans in the area. The relationships between annual re- productive patterns and mode of repro- duction are summarized in Table 11. When viewing the entire anuran fauna, it is evident that reproduction occurs throughout the year (Fig. 166); examination of all female frogs collected reveals that the percentage of gravid females varies from 54 percent in July to 86 percent in February. However, when the data are examined with re- spect to species, we find no more than 49 (56% of total species) species gravid in a given month (July), when a total of 64 species was found; thus, 77 percent of the species found in that month in- cluded gravid females. The extremes in the relative numbers of species to spe- cies gravid varies from 61 percent in August to 85 percent in February. It is interesting to note that in 1971-72, years when most of the data were accumu- lated, less rain fell in August and Febru- ary than any other months ( see Table 2) . When annual reproductive patterns are examined with respect to mode of reproduction (Table 11), we find only two unique combinations include more than one species. Thus, the centrolenids that deposit eggs on vegetation over streams (mode 5) breed sporadically after rains, and all species that carry 268 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Table 11. — Anuran Annual Reproductive Patterns in Relation to Mode of Reproduction. (Percentages in parentheses; see Table 10 for definitions of modes.) Mode o a o U t; o ft a O o •1—1 a u o ft T5 cii u o ft C/5 <: o 1 2 3 4 5 6 7 8 9 10 11 TOTAL 8 ( 9.2) 1 ( 1.1) 6 ( 6.9) 10 (11.5) 1 ( 1.1) 8 ( 9.2) 5 ( 5.7) iTTl) 26 (29.8) 14 (16.0) 19 (21.8) 1 ( 1.1) 6 ( 6.9) 3 ( 3.4) 4 ( 4.6) 1 8 1 43 1.1) 9.2) ( 1.1) (49.2) 1 (1.1) 3 (3.4) 4 (4.5) 35 1 1 11 3 9 6 1 18 1 1 87 (40.2) ( 1.1) ( 1.1) (12.2) ( 3.4) (10.3) ( 6.9) ( 1.1) (20.7) ( 1.1) ( 1.1) (100) ^80 o 70 oteo ^ 50 60 CO u o UJ Q. CO 50 40 u 30 □D 20 10 56 '64 . • ...261 ^-•^ 270 207 647*\283,/|24 All Females Gravid Females JFMAMJJASOND MONTHS Fig. 166 — Annual distribution of female frogs; upper graph is percent of gravid females (79 species); numbers above data points are total number of females. their tadpoles from terrestrial oviposi- tional sites to water (dcndrobatids) (mode 7) seem to be continuous breed- ers. Nearly half of the species at Santa Cecilia are classified as sporadic, wet breeders; I suspect that further study will reveal that some of these are more opportunistic or actually continuous breeders. Many of the 18 species having terrestrial eggs with direct development (mode 9) are represented by few speci- mens; thus, more data probably would reveal that several of the species with mode 9 are continuous, rather than spo- radic, wet breeders. Frogs having eight of the 11 recognized modes of reproduc- tion are sporadic, wet breeders; only five modes are represented among the con- tinuous breeders, accounting for 29.8 percent of the 87 species (Table 12). Although the data are woefully in- complete for many species, on the basis of the presence of gravid females of at least some species in each month and the climatic equitability, it is reasonable to conclude that anurans at Santa Ce- cilia are capable of breeding throughout the year. Rainfall is probably the single most important environmental factor controlling time of breeding, the breed- ing activity of most species is positively EQUATORIAL HERPETOFAUNA 269 o a o w bC c O CC ;-! OJ fc T3 O o w ^ 4) o 'q -H (D ft 0) ^ _ J2 c rt o E o :/3 U (D en C bO bo Z (U <-lH o bo > W o fl m _rt U( 3 > O (-■ o '> 4-' ^ c« ^ ^ (— ■ 4-1 C ■^ 0) t/D <4H . 2 < »\ -M w c ^ O "rt ^ g 1 0) 1. mh cm' <4-4 1—1 O H i-l PC ,13 < S H 2 c wi m c < H O H cr ^o 05^ inM cDoo 0000 mo looi i-nin o^I> t-— i 1— ICO 0-* CM05 ooo com I— 105 ■— 100 coco ooco c I— IrHi— (1— I I— li— li— (OlC-l OOCQ NO mo OO ■— lO (MO CIO OlO C5I> 1— !■* I— I o o o o CD n'tn Tfm (MO mo com mo ooo ^o coco <-ioo moj (>■! mi— lOi— II> CD O O COi— ll>COCD coo ost OO -^co -^m ■— loo coo i— lO oo i— i^ -ti— 100 O O(M00CJ5t^ mo 1— ICO 1— lo mo coo coo cmo (mo (3>t> ■— it> -^^^^ CD om 1-icomco coco cot-- 1— lO OO coco t^CD CD1> COO ^CO COCO ^COfMi-l CO 00 CD^t-'^COOCO mo t^t^ 00O5 'Tfit"- I>0 1— l-^ 00t> (MO t^t^ Gi -^ (MTf(MCO(M0O(M00 t-COOi— iCD^m^^^CO (M (MO ^co com coo OO mro i>o ^co co(m --ico ll^^T5<(M05i— lO (MOO Oi-lCO(Ma5CMl> i--(M coo t-co OO coco 005 (Mm co'f coco op ^00 m COi— lO CO(M00(MO5CMl>CMCO^l^ Tt^O TfO mo (MO (MO i-<00 '-lo o mo mo ooo —i^ SS2 ^iG fi^ oo O O Oi-IO 0'-OC0OOpm'H o m o o o o o oi— it>coa) 1—1 1—1 CO t> ro O 05 c a. V. c ■a o a. o o (a Co 5b c o o O t — , o 3 c CJ '^ o < (T iD O Of 60 50 40 30 20 10 0 20 CO UJ o LU a. en o en cr LlI m 15 - 10 5 - 94 \I59 .61 89 39 17 15 ALL FEMALES GRAVID FEMALES MJ JASOND MONTHS Fig. 167. — Annual distribution of female liz- ards; upper graph is percent of gra\id females (30 species); numbers above data points are total number of females. tive. The data on lizards at Santa Cecilia are incomplete. In fact, adequate data are available for only four species, three of which are Anolis, which evidently breed throughout the year (Fig. 168). The other species is Ameiva ameiva, fe- males of which contained oviducal eggs from May through December (93 speci- mens). Simmons' (1975) analysis of ova size and volume of fat l^odies demon- started the cyclic reproductive pattern in this species. By combining data from Santa Ce- cilia with those provided by Dixon and Soini (1975) for many of the same spe- cies at Iquitos, Peru (Fig. 169), it is possible to obtain a better idea of annual reproductive patterns in the lizards. Still, I am unable to demonstrate accu- rately the three patterns, as suggested by Sherljrooke (1975). With the present amount of data, the following 13 species EQUATORIAL HERPETOFAUNA 271 lOOh 75 Q 50 > ^ 25 LjJ ^100 Lu 75 -50 y 25h < A. chrysolepis ^y 0 0/ /4. fuscoauratus II II UJ oiOO LU "^ 75 50 25 0 I A. trachyderma 4 22 24 12 II. JFMAMJJASOND MONTHS Fig. 168. — Percent of female.s gravid through- out the year in three species of Anoih: num- bers above data points are total number of females. seem to be continuous breeders: Gona- todes concinnatus, Pseudogonatodes gukmensis, AnoJis (all six species), Plica umbra, Kentropyx elegans, Lcposoma parietale, Neusticums ecpleopus, and Prionodacti/Jus argidus. The following seven species probably are non-continu- ous breeders: Thecadactyliis rapicaiida, EnyaJioides cofanorum and laticeps, Alopoglossus atriventris and copei, Ameiva ameiva, and Mahuya mahowja. Data are either non-existent or too mea- ger for the other nine species to assign them to a category. Of the non-continu- ous breeders, the breeding season is longest in Ameiva ameiva ( May-Decem- ber), whereas females with oviducal eggs of the other species were found only in March-August. Generally those species that are con- tinuous breeders are small lizards; the .range in mean snout-vent length of gravid females is 26.5-100.7 (x=57.4) mm. The snout- vent lengths of the non- continuous breeders are 47.7-114.6 (xr= 89.1) mm. Snakes To one who has worked extensively in lowland tropical rainforests, it is soon obvious that snakes are not found in large numbers. The apparent scarcity of snakes becomes more obvious when the reproductive data are examined. Of 546 snakes from the Santa Cecilia area only 168 are adult females; only 26 (15.5%) of those were gravid (Fig. 170). Examination of the data on those species best represented by females re- veals no clear patterns (Table 13), and the paucity of data precludes any con- clusions. The largest collection of snakes from any one area in the Amazon Basin is the Bassler Collection in the American Museum of Natiual History; the snakes were collected over a period of several years in the region of Iquitos, Peru. Oliver ( 1947) reported on the Leptophis aJmetuUa in the collection, and Fitch (1970) summarized the reproductive data on Imantodes cenchoa, Leimadoph- is regimie, Leptodcira annidafa, Xerio- don severus, and Bothrops atrox. They concluded that these six species (the only ones represented by adequate sam- ples of gravid females and/or young) breed throughout the year. Perhaps the general trend in Amazonian snakes is to breed throughout most or all of the year, but I suspect that diverse patterns exist. FECUNDITY Although empirical data on frequency of reproduction are lacking for most of the species, it is possible to draw certain inferences from the quantitative assess- ments of ovarian complement, ova size, and female body size. This infonuation provides some insight into the quantita- tive aspects of reproduction in the herpe- tofauna. The available data are much more substantive for the anurans than for the lizards; data are meager for snakes. I have drawn upon material 272 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY J F M A M J J A s 0 N 0 J F M A M J J A S 0 N D Gonatodes concinnatus • • • ▲ A A A A Alopoglossus atriventris A A A Pseudogonatodes gutanensis O • o • o O O O o o Alopoglossus copii • • A Thecadactylus rapicauda • o • o Ameiva ameiva ▲ ▲ o o o o • A • O • O ■ ■ o Anolis chrysolepis ■ ■ ■ ■ ■ ■ ■ • Arthrosaura reticulata • A • Anolis fuscoauratus O ■ ■ A O ■ ■ o o ■ • A ■ o ■ o • o Bactiia trinasale O o • o o Anolis ortoni o o o • O o D n Iptiisa elegans A o o o Anolis punctatus o o o • • o ■ • o o Kentropyx pelviceps ■ o ■ ▲ O o o A O A o Anolis trachyderma • ■ • o ■ ■ A ■ o ■ ■ D • o Leposoma parietale O • • • o • o o D O Anolis transversdiis o o o o o Neusticurus ecpleopus ■ • o o o ▲ A D o o O Enyalioides cofanorum • ▲ A • Prionodactylus argulus ▲ D A ■ a A D O D A D O Enyalioides laticeps ■ ▲ o • Pnonodactylus manicatus • A A A Plica umbra • o o o • o o o Ptychoglossus brevifrontalis • o o Polychrus marmoratus o o o o o o Mabuya mabouya E E E E E E A -J F M A M J J A s 0 N D J F M A M J J A S o N D Fig. 169. — Reproductive data for 26 species of lizards from the Santa Cecilia region, Ecuador, compared with data from the Iquitos region, Peru (Dixon and Soini, 1975). Circles represent the presence of mature oviducal eggs; triangles represent hatchlings; squares represent eggs and hatchlings, and E's represent embryos. Solid symbols are data for Santa Cecilia; open symbols are data for Iquitos. from other localities in the Amazon Ba- sin and from the literature in order to have comparative data for some of the species of lizards and several species of snakes that are poorly represented in the collections from Santa Cecilia. As before, anurans, lizards, and snakes are discussed independently. Anurans Crump's (1974) thorough analysis of quantitative reproductive variables in 66 species at Santa Cecilia obviates any lengthy discussion of the data here. Her data base has been expanded to 76 spe- cies (Table 14). The additional data lend more support to Crump's conclu- sions regarding size-fecundity relation- ships; these conclusions can be sum- marized, as follows: 1. Species depositing eggs in water have larger ovarian complements ( 143- 8598, x=1225.3, N=31) than do those depositing eggs in aquatic foam nests (81-1589, x=457.0, N=6) or on vegeta- tion over water (61-1036, x=342.5, N= 11). 2. Species having terrestrial eggs un- dergoing direct development but no pa- rental care have larger ovarian comple- ments (5-62, x=19.3, N=17) than do those species in which the adults carry tadpoles (7-28, x=13.7, ]Vr=6). 3. Species depositing eggs in water have smaller eggs (0.8-2.1 mm, .x=1.35, N^31) than those depositing eggs in a foam nest (1.0-2.4 mm, x=1.85, N=6) or on vegetation over water ( 1.0-3.6 mm, x=1.91, N=ll). 4. Species having terrestrial eggs undergoing direct development have larger eggs ( 1.8-4.1 mm, x=2.72, N=17) than those having terrestrial eggs hatch- ing into tadpoles (1.5-2.8 mm, x=1.88, N=6). EQUATORIAL HERPETOFAUNA 273 9 40 ^ 30 o ot20 CM- 10 0 25 CO y 20 o LlI Q. cn LlI m 15 10 20 39 10 52 ALL FEMALES GRAVID FEMALES J FMAMJ JASOND MONTHS Fig. 170. — Annual distribution of female snakes; upper graph is percent gravid females (52 species); numbers above data points are total number of females. 5. The ovarian size factor is highest and most variable in species depositing eggs in water (4.9-110.2, .\=:24.7, N= 31), followed by species depositing eggs on vegetation over water (4.5-26.2, x=: 8.9, N=ll) and in foam nests (1.1-8.4, x=4.2, N=6). Among species deposit- ing eggs in water, the ovarian size factor is greater in larger species. 6. The ovarian size factor is lowest in those species having terrestrial eggs un- dergoing direct development (0.5-4.4 x=1.2, N=6). 7. In some species within a given reproductive mode there is a positive correlation between the number of eaes and the size of the female. 8. In some species within a given reproductive mode there is a positive correlation between ovum size and the size of the female. Lizards Size of females and reproductive mode (viviparous versus oviparous) are not highly correlated with the size of the ovarian complement in most lizards. Instead, the size of the ovarian comple- ment seems to be rather constant within plnlogenetic groups (Table 15). All gekkonids ha\'e onl\' one oviducal e^s at one time; anoles (Iguanidae) have one or two oviducal eggs, and microteiids normally have two eggs. Comparison of ovarian complement with size of female in the other lizards shows a positive cor- relation (Fig. 171). Generally, the smaller species of liz- ards have smaller eggs; the length of oviducal eggs ranges from 5.2 mm in Leposoma parietaJc to 33.7 mm in Tiipi- Table 13. — Monthly Distribution of Females of Six Species of Snakes with Mature Eggs. ( First line is number of females; second line is number of gra\id females with mature eggs. ) J F M A M J J O N D TOTAL Dipsas catesbiji 0 0 Imantodes cenchoa 1 0 Leimadophis reginae — - - 0 0 Leptophis ahiietulla 0 0 Oxijbelis argenteus 1 0 Bothrops atrox 0 0 0 4 2 1 3 0 1 0 0 0 11 0 0 2 0 1 0 0 0 0 0 0 3 0 2 0 2 1 1 1 0 0 1 0 9 0 1 0 0 1 1 0 0 0 1 0 4 1 0 1 1 0 2 0 0 0 0 0 5 1 0 1 0 0 0 0 0 0 0 0 2 0 0 1 1 1 1 0 0 0 0 0 4 0 0 1 0 0 1 0 0 0 0 0 2 1 2 1 1 1 4 3 1 C) 1 0 16 0 0 0 0 0 1 1 0 0 0 0 2 1 2 1 2 1 3 0 1 0 1 0 12 00000010001 274 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY u ^ a r\) o > o N 4-* o 03 (1) y 'c « ^ P s O Q 4-> c 0) be c r" rt s K a 1— c o U c rt ^ c n1 rt ^ 1) •5 C hJ •4-> c > I -t-J 3 C C C/} CH bC C CO cfl Q O V5 u O U on o c o in (MiC'-HOsoiiX''— ii;Dt^'^ir5'— |c<^o5'-^>-;^-■ o^-^l>T}^coc— ICOOlr-li— li— llO'~'C0CO 1— li— I i^oqa5T}i— iCO00CO(M000Om'*^CO(MCO^ t^i— li— iCNi— I ooqcDi-jin^i— 10000— i^iraoqcorHCOTf oo' t-' o o CO o 05 t^ o i> CO t--^ lo i-< -t^' -^ in cxicoinTfi^oococoinci'M'-^^'— 'cotMoo 00 1—1 1—1 -fOa5cocooinco C<|TticOCOCO"^I>"^Ccoi— ii>i— icooini— lOfM 1— I'^Csli— <>— iCNi— <(M "—((Ml— I OCMin oo oq o oq CM i in 00 TP ! CO CD i-( 00 O 05 Tt I> l^ — 1 1—1 1—1 ^H Tj< t^ CO 05 ; 05 1> O O CD CO in 1-1 in CM (M o a> CM CM O 1-H 1-J cm' CM 00 CO (M CO 1— ( O 1—1 1—1 1—1 o o CO o in in O 00 CD 1— ( . 1 , . o , in . . 1— * ? i 1— ( 1 1 ; 1 : ! i i::^? I'? 1 i"? 00 1 o : : ; rH o 1 1— ( 1 ! O CD ■ 00 ' ' CM ' in ' ' t- ^ I> 1— 1 CO T}< o 1-1 o o O O 00 : o O o o o —1 O 00 o o CM 00 Tf m CO in in 00 in ^ 1—1 : CO O O) t in in —1 in 00 CM '. CM O O 00 CO ' 1— 1 1— 1 in t^ i> (M CM CM t> t^ 1—1 I— 1 i 11 1 ! in ' ' CO ; ^ : : CO , , , (M 1 1 1 •— 1 , O ■7 i : -<5< 1 lei i ' in ' ' i i i^ iiA i ' CM ' in o 00 o o 1 O O CT> 1 CO O O CT3 05 05 CD CD CO rjH in l> CD ! CO 00 CD ' CM "^ 1-1 1 t^ CM -1^ oq cj in CD I— I CD 1—1 1—1 CD 1-1 i-i r-< XO CM q o CM CO Tf CD CO 1—1 1—1 CM 1—1 ^^^uuu^^u^u^^uo^u^o?:^oooooo^^u^^^^^^ CM 3 3 c S •= ? ? ? S 2 2 £ •2 I St o j- Ci) ,, 3 a o o ti) So b£-S a a o c C 3 ^"^ ^^H* ^ll^rf ^11^ ^ll^J ^' m^ ^ ^ f"^^ ^11^ ri I _| s s c 2 ? « a. ■ 35 3> a: a: 3> t: 50 3 05 3 3 "^ ■^ ~ -a r^r *- c: c- 3 3 o a ja 3 (Jj CD >~ Cj u r- '^ O O tjj ^ 3 C ~ &; 3i 3i 3 a ,^ ^ 3 ^ o o O 3 3 J- ^ 5J 3i 3 3 Co ■~-^ 3 "^ _co Co 2 2 3 3 do Co .3 3 3 3 j^^^co to to^,™ !i3.3 — .-^-.o a5a::a:;ooO(x,esooc/3K:oooa 3 •♦.;> 3 3 S i '^^ 3 5 » tx _co CO S 3 ^^ 3i O »3 3 S2. Co O 3 3 3 C 3 Co a la '5) a. 3 O 3 a -5 £ -G 3 3> EQUATORIAL HERPETOFAUNA 275 c CTi > o c75 c 0) S dj 'a S o U 'C > Ot So .2 Q c in be c 0; cc; 4-) c a> > •t-' 3 O C c C/D Q o en U U M O5i^coc]^i>oq-^i;(>iincococsi ; ^ ^ CE> ^H .— ( ^H >— ( CM i ^ 00 CNl oi I— i I— i I— 1 1— i >-< c-! — I I I I I 0 00 O 00 _ _ . 01 CO (M CO -H CO -n< o o in lo -< in CD -^ ^ CO (M CM I I I I O 05 CO 00 00 (N O) .-H -^-(Ot00 ^H CO 00 CM O 1-1 CO 'I* _ . in in 00 CO CO CO f CT) in CD (N o 05 05 05 o ■-^. CD in in CO c-i 00 in p cjj oi CO o o (m" CO 05 CO iri t--' oo' in CO 05C0(Mc<|rpTPC0C0rpoint>CNC-; M t^_ oi l> >-H "— i O oq t-. CD i-_ p '^ CO in r-- Ti< I— iOi— — i CO oi ci CO cq CO o in o 1-1 CO coc<, o in O 05 O rH t- TjH i:o 1— I I CO CO 03 I I CO T CO CO I— I in CM Tf r— I (M tM CD I I I I I 00 CD (M CM t- -—I ^H C} 00 o t- l-l I I CM CO 00 O CO — ^ p CO -t t-; 00 t-; -r (M_ p p : p 00 in 1 1 : p in o ^ P 00 oi CD i ' 00 CO 00 ' CM CM i ; 00 cm' CD ' ' 00 — 1 Tt< in CO CO Tf< »-( (M CM CM I I I I CD t 00 O I— I C] "—I CM CO 00 d) •-( CM in CO in CM Cvl CO CO ni CM Tji I 1 I I I I I Tt< in t^ i- o 00 o CM CM OJ CM -^ i— I T}< coinooqppinppt-;oq^_coco- in ■— i f>i CO in CD 00 00 cm' o cm COincO-— iCMCMCMi— (CM(MCMCMTfiCM't>t^I>t^t>00C7)O5C7)O5C75 ? s >/3 o l~ c t: !i) o ^ o •^ r^ i-Q a a a ;:r) rn ;ri Q to 'a "^ •2 a c 5=^ £ s: G ^ « "3 Q l~^ ^ "^j »5 Co c*-> ?^ 5 « = S 2 - ^ Q c: ts -^ "13 a -- c; a 1^ : Co CO 3 C •2 .„ S s; ^ SI 00 3 a 32 I — t3 CO 3 5 CO 3 o £= S »- a 2 "^ O .CO "c CD 5 •s i 2-2 rS rS -S -2 '^"51'X , 3. 35 3^ =5',.-',.^ ^, 5: a; u: &: Bs A. ft. '^I33i353>3^353i C.SOCJOCJCJCJ vTOOOOOOO .+^ '^— .i— H-* .^-J H-j H.^ S.- 3 Co *« i; c/5 C-^ 3i >>J Co 'Ij 2 a _ . o t; CO CO '*~- CU 1^ CO c c 3-, 1^ !D ~. a ^ cj o ^ c a a g 't: -t; 'Q t) g g g £ <^ S ^ o -c ^i: ri: 35 3-. 3i CJ o o o t: kj tij tq tq tq 276 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY .2 n> o o in i> ^ "^ ; in •-I o CD 00 i> i> ^ "^ I— 1 CM O "-H "-H : CD .-1 oi r-H CD CM o > CD 05 O ■d 'i 4-< in og o in CO o o ^ in o --; oo_ « 3 ^6 3 CM CO Csl (M C o 4-1 c 1) 3 O C CO Ot W) c be c OS c (U W D o en U U w C/2 CD CM CO O CO 1— I in "— I '— ' '"' I 11 II i> o in 05 in CM in ^ cxD CO p ai d i>- ai n CO ■— t (M CM --H in CD I I I CM in in o CO p o !>: --; '^. in i> (M CM CD in in CM CO CO cx) CD CO a CM xf CM ^, C>1 c:^ 4cr)<^ CO CM CO ■— I CM in t-; t-; p -t; CO 00 ■— ! oi CD (M CO CM CM (M 00 00 CO CT) CD ^ CM CM CM I 00 CD CM Tf (M CD 1—1 I I I CM t- CM CM '^ •— I p 00 p o -t; p '^'^ cjj o crJ CJ5 -^ CM CO I— I CM CM '■f CM in ■— I I— ( O] --H ^ 00 Tr< in ^ CO in CM o CD CM l^ CD uuoou^^^^^uu^o^ CDCJ5a3CJ5CX)C35aiCJ)CJ5a5a>a5050 3 05 3 CO 3 &3 03 3 '» - ^ =? -ti ' 22 ? a c^ a Cic 100 B, D • 1- 0 ^ 90 A. ^ E. UJ _i 5 80 UJ u_ / c. 2 1 1 1 1 1 2 2 4 6 8 10 ^ MEAN OVARIAN COMPLEMENT Fig. 171. — Mean size-fecundity relationships in eight species of lizards: A. Uracentron flavi- ceps, B. Emjalioides cofanorum, C. Flica umhra, D. Amciva ameiva, E. Mabutja mahouya, F. Kcntropyx pelviceps, G. Emjalioides laticeps, H. Pohjchrus nmnnoratus, r=0.78444; P<0.05. nambis tequixin. However, if all lizards are con.sidered there is no significant correlation between ovum length and female snout-vent length ( ri=: 0.58028; P=>0.05). Presumably the gekkonids (with the possible exception of Thecadactyhis), the anoles, and the microteiids (with the exception of the two species of Alopo- glos.sus) have multiple broods. This as- sumption is leased on the small clutch size (one or two eggs) and the presence of numerous gravid females throughout the year. Plica um])ra and Kcntropyx pelviceps appear to be continuous breed- ers and may have multiple clutches in a year. Ameiva ameiva is a cyclic breeder, and Simmons ( 1975 ) found no evidence for multiple broods. Limited data for other species suggest that single broods may be the rule in Emjalioides and Ma- huija. Snakes The paucity of data precludes any broad conclusions regarding fecundity; the meauer data from Santa Cecilia have EQUATORIAL HERPETOFAUNA 277 Table 15. — Size-Fecundity Relationships of Lizards and Amphisbaenians Based on Mature Ovarian Complement. 5 Snout-vent Length Species N Gonatodes concinnatiis 6 P.scudo^onatoclcs giiianensis 2 Thecadacttjhts rapicauda 4 Anolis chnjsolcpis 25 Anolis fuscoauratus 89 Anolis ortonii 4 Anolis punctatus -.. 5 Anolis tiachyderma 68 Anolis transversalis 1 Enyalioides confanorum 2 Emjalioides laticeps — . 5 Plica umbra 4 Polychrus marmoratus"^ 7 Uraccntron ftaviceps^ 3 Mahuya mahouya 6 Alopoglossus atriventris^ 7 Alopoglossus copii 4 Ameiva ameiva — 12 Arthrosaura reticulata 3 Bachia trinasale 2 Dracaena guianensis .— .. Iphisa elcgans _ 1 Kentropyx pelviccps 3 Leposoma parictale 12 Neusticurus ccplcopus 2 Prionodactylus argulus 4 Prionodactylus manicatus 1 Ptychoglossus hrcvifrontalis 2 Tupinamhis tequixin 1 Amphishaena fuliginosa .— Mean Range Ovarian Mean Complement Ovum Mean Range Lengtli 41.6 26.5 107.8 73.7 43.8 42.5 70.6 50.9 72.0 99.5 114.6 83.5 128.0 95.0 93.7 47.7 58.3 102.2 66.6 74.5 52.0 100.7 36.2 61.5 42.5 64.0 58.5 307.0 35-48 26-27 97-116 66-86 38-50 42-43 67-72 45-56 92-107 107-125 80-90 109-147 79-117 89-100 41-51 53-62 90-118 64-70 70-79 95-106 34-39 56-67 39-44 58-59 1.0 1.0 1.0 1.3 1.3 1.5 1.4 1.1 1.0 2.0 6.3 2.5 9.3 2.0 5.2 2.0 2.0 3.7 2.0 2.0 2.0 5.0 2.0 2.0 1.8 2.0 2.0 3.0 1-2 1-2 1-2 1-2 1-2 5-7 2-4 7-11 4-6 2-6 4-6 1-2 8.9 6.0 19.0 16.3 9.1 9.9 16.3 13.1 14.4 27.4 15.8 22.6 13.1 23.7 10.7 10.5 21.0 13.3 11.0 7.6 17.5 5.2 11.0 9.1 15.0 8.3 33.7 "■ Data from Dixon and Soini ( 1975). ^ Viviparous. been supplemented by additional data from specimens from other localities and from the literatiu'e (Table 16). Vivi- parous species, which also are among the larger snakes, have larger broods than do oviparous species. Mean ovar- ian complements of viviparous species are 10-42 (x=25.2, N=4) as compared to 1.8-22 (x=7.6, N=28) in oviparous species. Among the oviparous species, the largest ovarian complements were found in the larger heavy-bodied snakes, such as Clelia, Oxi/rhopus formosus, Xenodon, and Lachesis. Moderately large complements (11-12) were found in the two aquatic Helicops, which may be facultatively viviparous (Rossman, 1973). The slender arboreal snakes (Dipsas, Imantodes, Leptodeira, Lepto- phis, and Oxybelis) have no more than six eggs. There is no evidence from the data from Santa Cecilia that any of the snakes have multiple broods during the year. In fact, the presence of so few gravid females suggests that reproduction with- in individuals may be cyclic - perhaps annual or bi-annual. On the other hand, some of the more abundant species, such as OxijheJis and Leptodcira, have smaller clutches than do less common species, such as Boa, Clclia, and Xenodon. If the limited data on clutch sizes and rela- tive abundance have any significance. 278 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY So C c OJ C/3 0+ •^ t; t^ CO "-* CO — ' t! 1— ( 05 P O) <^ fc CO (U 05 05 a> 3 .-1 ^ i-H O O -C 0) C/2 aii a o ^ o PM fe P^ ■5 to c c OJ in W) (— r^ CD rt 'a P5 S o U c •S C a > O 0) Q O CO 0 u I© (M O o 3 o W 3 cf CO •l-H to J rt rt O pq m CO CJ § s" s" O s ^ ^ s '3 — I oq CO CO oi ^ oi in d -i CO CO 00 1> ci T pppinpqpppcD cdccddcdoq cq CO CO t^ CQ cq i i M i i^ ! i^ I I 1 i i^ ; 1 1 1 1 : CD : i » ; 1 ! 1 i 00 o CO 00 lO CO CO 05O05O0010OOO10 COIO^Cnc0005t^05CO lOCOOJCOCOtMCNOCOTr I— ( I— I oq Clr-li-((Mi— I"— li— li— |i— ICO 13 u w >-l lO in 00 > CO : CO I- (N 1 : : 1— 1 CO CT) i oi oi in i ' oq oq CM ' i i in d CTJ ' ' eg c CO t> M in O CO t^ CT) ro t-- "^ 00 CD o in ^ CO t^ CO '^ >— I f— I 1— I ■<* I— I C>>>>0000000000000 000\\0000 00 0 0000 O O Co 00 C 3 ■V. ;j c S..C 3 ►a CO « 2 o o CQ O (^ ^ a c u a Cj c o CJ <5j s o Co Co Co Cj ?j Co Co ^ 3 Ho HO e Ho Ho Ho c •^ ^ 'O O O O Q ? a e ^^ C/^ Co c/^ p tq tq ^ ^ -< U O O G U Q Q Q Co c •2 ^ 1^ -i; ^*- *'— H— ""^ ^ o . :o ^'^ Co • ^ Co 00 ol ~^ IT:. E^ = o Co '~ . 1^ ir Ho ~ C ^ -2i o, (i) C a J^ ~ c ;^ ~ >--: 'o 'o ' '^ "^ '■— o ^ s. S 5 2 2 i o o Q Q Q Qt5 i^:^ c ' a ^ ^ " S 5 f§ •? •2 -2 ^ '^ ^ I 't^ 2 '^ S'^ ~2 -2 "S .5 ^ ^ 2 2 O Q-^ e ■S .£ U a. ~ ■= S 5 S" S" -2 :~ Kol hj sj Kol kJ ;2; c fcjO o s o 5; Ho c a *-. ro« s »o o 1^) tx H-, (^ Jo a Co CO .^ sa. 5i. ^o O 0 ^ f«a -CJ '^ '^ s» 2:5 :^ >■: !-; H ooo EQUATORIAL HERPETOFAUNA 279 0) o u o 1=1 S "^ -< O J -^ c o CD I— ( n c (U QJ M C 'a OS E o U c _rt C a > o C I -t-J 3 o c bO c OS Q o CO in J3 fN : CO ; ; ; : c:> 1 1 1 ; o : : , o CO i CO i i i i CO i i ■ i en 00 q o q q c4 CO q q 1—1 N o CO CO o 1 "—I ; 05 1 ; ; ; CO CD ; : i t- ; csi ; ; o t- CD i-H O CO ID i ; CO CO i-H ■—I o r-H CM OQ oooooooooooooo u o o « CO O ft. 2 § 2 cv '-^ 2 ^ CD s e Co = ^ ™ ^ -S.S ^:§ Bh k; CT) h« •~ r: *r a S Ci "^ *^ ^ O c^ CO c a to ^ CO *>? O C2 "' '^ ~ ■~ IT = c s :5 "E i a S 5. ? 7- '^™ ^— Co _CJ en Co Co Co S c c: c c S ^ ~ o = = a o o i: s~ -^ i- ?j o cj c; c; •^ 5 ? s: ^^ C ^ "3 Q^ § S c' seek shelter. Studies on reproduction in a variety of tropical lizards by Marshall and Hook (1960), Inger and Greenberg (1966), Sexton and Turner (1971), Sexton et ah (1971), Leon and Ruiz (1971), and An- drews and Rand ( 1974 ) demonstrate that in seasonal environments, oviposi- tion occurs during the rainy season, whereas in continuously wet tropical environments, oviposition is continu- ous, or nearly so, throughout the year. As noted by Tinkle, Wilbur, and Til- ley (1970), lizards inhabiting the wet tropics generally are early matur- ing with multiple broods annually. One strategy of the early maturing lizards is to place a large hatchling in the environment. The ratio of snout- vent length of hatchlings to that of gravid females in ten species of lizards at Santa Cecilia is 37.4-43.2 (x=39.9) percent. The ratios vary only slightly in different adaptive groups: 40.9 in Gona- todes, 38.9-43.2 "(x=40.4) in three anoles, 37.5 in Enyalioides laticeps, 37.4- 42.9 (.x=40.3) in three microteiids, and 39.1-39.2 in two macroteiids. Thus, the number of eggs in a clutch shows no significant correlation with the relative size of the hatchling, although those species having larger clutches have pro- portionately smaller hatchlings than do some of the species having smaller clutches. Seemingly, reproductive strategies have evolved in order to provide the most effective input of young into the population. The production of young must be viewed over the reproductive life span of the female. Here, the aspect of risk becomes important (Tinkle, 1969). Obviously, the burden of ovi- ducal eggs is risky to the gravid female; a small hatchling runs a greater preda- tion risk than a larger one, and a late- maturing lizard runs a greater risk of death prior to reproduction than does an early-maturing species. All of the re- strictions must be balanced against an effort towards high fecundity. A reduc- tion in clutch size is offset by increasing the number of clutches. A reduction in clutch size theoretically also can be off- set by producing larger hatchlings that reach sexual maturity in a shorter period of time. The reproductive strategies exhibited by the species comprising the commu- nity at Santa Cecilia can be summarized, as follows: 1. Early-maturing with multiple clutches A. Small clutches inherent phy- logenetically ( possibly also due to load-bearing of adhe- sive lamellae) - gekkonids. B. Small clutches primarily due to load-bearing of adhesive lamellae and need for agility - anoles. C. Small clutches due to neces- sity of rapid escape - Enya- lioides cofanorum. Plica um- bra. D. Small clutches because of re- striction of body size - mi- croteiids. E. Large clutches possibly be- cause of large size and slow movements - Enyalioides lati- ceps, Polychnis marmoratus, and Kentropyx pelviceps. 2. Late-maturing with single clutches A. Smaller clutches due to nec- essity of rapid movements - Ameiva ameiva. B. Larger clutches, viviparous - Malmya mabouya. C. Presumably in this group - Dracaena, Tupinambis, and Uracentron. Snakes Within the serpents there is a basic dichotomy of reproductive modes - ovi- parity versus viviparity. Neill (1964) 288 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY and Fitch (1970) addressed the two modes. In contrast to oviparons species, viviparous snakes generally have smaller broods and reproduce less frequently. Advantages to viviparity are the lack of need to find a suitable nest site and pro- tection of the embryos from predators and unfavorable environmental condi- tions. A definite disadvantage to vivi- parity is the length of time that the fe- male must carry the young and her susceptibility to predation while gravid. Neill ( 1964 ) emphasized the protection afforded the developing young of veno- mous species, and Fitch (1970) further noted that viviparity was most prevalent in heavy-bodied, slow-moving or aquatic snakes. At Santa Cecilia viviparity occurs in the five species of boids and three spe- cies of Bofhrops, all relatively heavy- bodied, sluggish snakes. The more slen- der Lachesis muta is the only species of crotalid known to be oviparous. Appar- ently the water snakes of the genus HeU- cops are facultatively oviparous or vivi- parous (Rossman, 1973.) Viviparity in these aquatic snakes eliminates the ne- cessity of their leaving the water to de- posit eggs. All other snakes at Santa Cecilia are oviparous. Clutches are larger in the viviparous species than in the oviparous ones; this apparent contradiction is obviously due to the fact that the largest species in the community are viviparous. Among the oviparous species the heavy-bodied ter- restrial snakes have larger clutches than the slender-bodied arboreal species. The latter certainly would be hampered in their movements by the burden of a large ovarian complement. Som(> slender ter- restrial snakes (Chironius and Dendro- phidion) may be similarly restricted. DISCUSSION Certain aspects of reproduction in the fauna have not been covc^red in the foregoing sections, (-ither becaus(> they were not pertinent or because they in- volved arguments or concepts not rele- vant to any one of the organized cate- gories. Reproduction in Other Groups In order to complete the presentation of the data on groups not covered in the preceding accounts, the information is summarized here. Caecdiam. — Presumably all of the caecilians at Santa Cecilia are viviparous or at least oviparous with terrestrial eggs and no aquatic larvae. No information is available on annual reproductive pat- terns or fecundity. Salamanders. — The two species of BoUtoglossa presumably lay terrestrial eggs like other members of this wide- spread Neotropical genus. BoUtoglossa peruviana has 6-13 ova maturing at one time, and the seasonal incidence of grav- id females and young suggests that re- production is acyclic. Turtles. — According to local resi- dents, at least some of the aquatic pleu- rodire turtles (Cheliis, Mesoclemmys, Phnjnops, and Platemys) deposit eggs on sand banks at times of low water in the Rio Aguarico - usually in December; presumably only one clutch is laid per year. Geochelone also probably deposits only one clutch annually, but Kinoster- non may have multiple clutches. Crocoddians. — As a group, the Neo- tropical crocodilians apparently have single large clutches of eggs each year. Reproductive Potential The data on, and discussion of, fe- cundity are based almost exclusively on ovarian complements. Although these data are basic to certain aspects of our knowledge of the reproductive biology of the species involved and of the com- munity as a whole, they neither provide a comprehensive picture of fecundity nor of reproductive potential. Also, in order to lune a true picture of reproduc- tive potential, it is necessary to know the reproductive life span of the females. Andrews and Rand (1974) reported EQUATORIAL HERPETOFAUNA 289 that Anolis Jimifrons in Panama lays one egg every 8.1 days; females reach sexual maturity in four months. Thus, based on survival data, each female has the po- tential for depositing about IS eggs dur- ing her life. At Santa Cecilia AnoJis fuscoauratus (a close relative of A. limi- from) and A. trachyderma are like A. Umifrom in size and habitat. I suspect that the reproductive potential of these species is comparable with that of A. Jimifrons. Crump (1974) reported one female of Hijh rhodopepla from Santa Cecilia deposited clutches of 390 and 260 eggs 31 days apart. If a given female repro- duced at this rate for an entire year, she would produce only about 3900 eggs. This is a relativelv small number of ecics, for some species, such as Bufo marinus have clutches in excess of 10,000 esiss. Obviously survivorship is extremely low in most high fecund lizards and frogs. '-' O For example, if all the eggs of a given clutch deposited by two pairs of Biifo marinus matured to adults, the resulting toads would completely cover one square kilometer. General Conclusions In a final consideration of the repro- ductive modes and strategies operant in a herpetological community in an asea- sonal tropical rainforest, it must be borne in mind that, with possibly few minor exceptions, the mode of reproduction and strategy utilized by any given spe- cies evolved in that species prior to its inclusion in the present community. Thus, it is not reasonable to infer that a species evolved a certain reproductive mode or strategy in response to the pres- ence of other species in the community as we know it today. Instead, each re- productive strategy is the result of selec- tion for an effective means of producing a succeeding generation. The selective forces include, but are not necessarily restricted to adaptations for survival of eggs and young (including larvae) in unpredictable (sporadically fluctuating) environments and the survival of gravid females against predation. Although re- productive potential may be high in many species, survivorship, especially of eggs, larvae, and young, is low, whereas survivorship is higher in species with lower fecundity. Although there are some exceptions, lower fecundity and higher survivorship prolxibly are char- acteristic of species at higher trophic levels, species inhal)iting the more pre- dictal^le environments, and those with parental care. In the aseasonal climatic regime at Santa Cecilia, reproduction in most spe- cies apparently is continuous, or nearly so. There is a continuous infusion of young into the population and overlap- ping generations. Continuity of success- ful reproduction tends to decrease popu- lation fluctuation. Increase in population fluctuation increases the rate of extinc- tion (MacArthur and Wilson, 1963). Thus, as suggested by Inger and Green- berg ( 1966 ) , the continuous breeding of the species in the community may be one of the reasons for high species rich- ness in humid tropical rainforests. Seasonal Versus Aseasonal Environments Subsequent to Dobzhanky's (1950) suggestion that natural selection in the tropics favors lower fecundity and slow- er development, as opposed to higher fecundity and faster development in the temperate regions, the literary frenzy of evolutionary biologists has resulted in a monumental pile of publications offering multitudinous theories and comparative- ly few data (for a realistic appraisal of the situation, see MacArthur, 1972:216). Because seasonality is characteristic of temperate and some tropical environ- ments, I confine my comparisons between the aseasonal tropical rainforest and sea- sonal environments, both tropical and temperate. Although the climatic re- gimes of temperate regions and extreme- ly seasonal tropical regions, such as those having a monsoon climate are dissimilar, the effect on the biota is much the same. 290 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Periods of activity are restricted, either directly or indirectly, due to cold or drought, and adverse periods (with the exception of non-migratory endother- mal vertebrates) necessitate inactivity through hiberation or aestivation of adults, larvae, pupae, or zygotes. In the case of aniphil)ians and rep- tiles, a single clutch of eggs is character- istic of species inhabiting extreme sea- sonal environments, although the same species may produce more clutches in more equable parts of its range. Fur- thermore, reproductive maturity may not be attained for two or more years. Tim- ing of breeding is usually dependent up- on the presence of available moisture - rainfall in dry areas and thawing as well as temperature in cold areas. The timing of the production of eggs or young is not only critical with respect to the avail- ability of suitable oviposition sites, but also to the availability of food for the young as well as the adults. Also, it is necessary that the young have sufficient time to build up an adequate fat supply to carry them through the forthcoming period of inactivity. Amphibians and reptiles have evolved diverse adaptations for those adverse climatic regimes, in- cluding autumnal fertilization, overwin- tering of tadpoles and hatchlings, and viviparity- in squamates in temperate climatic regimes, and rapid development of ova and larvae and cunning ways to conserve bodily fluids in monsoon tropi- cal areas. These kinds of environmental stresses are minimized or non-existent in equable humid tropical rainforests. Aperiodic fluctuations in rainfall may result in the desiccation of eggs or larvae, but aperi- odic fluctuations are even more frequent in the other climatic regimes. Conse- quently, in populations where all sex- ually mature adults breed simultane- ously, the elimination of one generation is not an uncommon phenomenon in extremely seasonal environments. By comparison, in the tropical rainforest where onlv a fraction of the adults are breeding at any one time, the elimination of their offspring will not be nearly so catastrophic for the population. The advent of aperiodic climatic fluc- tuations will have a more profound ef- fect on population fluctuation in species in seasonal environments than in the aseasonal tropical rainforests. Popula- tion size is more unifonn in the tropical rainforests than in the seasonal environ- ments. The continuous activity of non- fluctuating populations lends greater sta- bility to the biotic environment in an aseasonal tropical rainforest than is pos- sible in areas with highly seasonal cli- matic regimes. Because of greater varieties of struc- tural habits and more equable environ- mental conditions, amphibians and rep- tiles in aseasonal tropical rainforests have exploited a wider array of reproductive strategies than they have in temperate and seasonal tropical areas. This is es- pecially true among anurans, which at Santa Cecilia are represented by 11 modes of reproduction; only the primi- tive Mode 1 is found in frogs inhabiting extremes of seasonal environments. Salthe and Duellman (1973) noted the diversity of reproductive modes in tropi- cal anurans and suggested that small frogs, into which general category most tropical frogs can be placed, are pre- adaptcd for e\'olutionary exploration of new reproductive modes. Although clutch or brood size usu- ally is larger in species in seasonal en- vironments than in comparable species in aseasonal tropical environments, breeding is more frequent in the species in the aseasonal areas, or increased pa- rental care in specialized reproductive modes in the tropics presumably results in greater survivorship. Thus, it is un- likely that any generalities about higher fecundity in the temperate regions have any validity-. Likewise, developmental time in ectothermal tropical species is likely to be shorter in the warm tropical regime than in the cooler temperate en- vironments. Of course, this comparison EQUATORIAL HERPETOFAUNA 291 must be made within a given reproduc- tive mode. In summary, amphibians and reptiles in ascasonal tropical rainforests employ a wider array of reproductive strategies, reproduce more frequently, mature ear- lier, and have more stable populations, but probably have lower survi\'orship and shorter life spans than do their coun- terparts in seasonal en\'ironments. The apparently complete absence of pair bonds combine with frequent mating and reproduction results in a greater genetic mixing, which, in addition to the diversity of structural habitat and vast array of other available resources pro- vides the potential for rapid differentia- tion and subsequent speciation. COMMUNITY STRUCTURE Although MacArthur (1971) defined a community as "any set of related or- ganisms near each other and about which it is interesting to talk," in reality a com- munity consists of all of the organisms that live together in a particular place. Usually only portions of communities are studied, and most frequently the division is taxonomic. Thus, the present analysis of the herpetological communit\^ is un- usual only in the species richness of the community. In order to study a community it is necessary to measure various parameters. A major weakness inherent in this ap- proach is that we measure those featiues that we think are important, but our in- ability to view the environment from the eyes of the organism imparts a bias and certainly a gross incompleteness to result- ing analyses. For example, the absence of temporal, spatial, and dietary differ- ences in the available data on the three species of Dipsas at Santa Cecilia can only be interpreted reasonably as an ab- sence of measurements of the ecological factors that are different in these snakes. The point here is that most analyses are not refined; certainly many more differ- ences exist than are normally measured, and the differences generally can be in- terpreted as being greater than those in- dicated in the analyses. Furthennore, unequal sampling in different habitats at different times re- sults in biased data (see p. 13). None- the less, given these inadequacies of sampling and measurement, it is possible to provide a basic analysis of the herpe- to faunal community. This analysis is more extensive than any heretofore at- tempted for a tropical rainforest hcrpeto- community. Temis used herein are defined, and where appropriate, analytical techniques are discussed below. Coefficient of Communitij. — This is Whittaker's (1970) mathematical meas- ure of the relative similarity of samples from two communities: ^^=^J(ih/ (So-|-S;, — Sab), where S,,,, is the number of species shared by samples A and B, S„ is the total number of species present in sam- ple A, and Si, is the total number of spe- cies in sample B. Ecological AUopafry. — Two or more species living in the same habitat but separated by differences in microhabitat or diel activity. Ecological Synipatry. — Two or more species living in the same microhabitat and overlapping in their did activity. Habitat. — That major portion of the physical environment in which an or- ganism carries out its life processes. As used here, habitats are major structural aspects of the area: forest, clearing, river, etc. Habitat Generalists and Specialists. — These terms are used in the sense of Crump (1971). A habitat generalist is a species utilizing a broad spectrum of the environment; this can be detennined from relative niche breadth scores. Hab- itat specialists are species living in a narrow range of the environmental spec- 292 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY trum; species that are not at the extremes of the continuum of habitat iitihzation are termed habitat intermediates. Microhobifat. — This is a subdivision of the habitat. For example, within the primary forest habitat, the following microhabitats can be defined: fossorial, leaf litter, ground, bush, tree trunk, and canopy. Niche. — This often maligned term is used in the sense of Hutchinson (1957) - an abstract concept of a multidimen- sional hypervolume. The dimensions are physical factors and biotic relationships necessary for the survival of the species. Niche Breadth. — This temi describes the spectrum of any given dimension of the niche hypervolume. The niche breadth of a species in a resource matrix can be measured by the methods sug- gested by Levins (1968) and modified by Colwell and Futuyma (1971) and Pianka(1972): j where p,j is the proportion of individuals of species i associated with resource /'. Standardized niche breadth values (Bst) can be obtained by dividing the niche breadth value (B) by the number or resource states. Niche Overlap. — This is a measure- ment of the association of two or more species with respect to some dimension of the niche hypervolume. The niche overlap of two or more species in a re- source matrix can be measured by the formula proposed by Colwell and Fu- tuyma (1971): Cin=l—}2%\pij — phj\, j where C is the amount of shared re- source states between species i and h, pij is the proportion of individuals of species i associated with resource /, and h is the second species in the matrix. In this measure, C has a value of 0 when species i and h share no resource states and a maximum value of 1 when the pro- portional distributions of the two species are the same. Combining overlap values from two or more independent analyses of difi^erent resources provides an over- all niche overlap value. The overall values were obtained by multiplying for each pair of species the total of the indi- vidual niche overlap values times 100. This is the "product alpha" of Cody (1974). Resource Partitioning. — The differen- tial utilization of the physical and/or biotic environment by different species. Species Diversity. — The concept of species diversity consists of two com- ponents - species richness and equitabil- ity. Species richness is solely the num- ber of species, whereas equitability is a measurement of the evenness of distribu- tion of individuals among species. In- dices of species diversity were calculated by means of the information theory of Shannon (1948): D=—%Pilog^Pi^ i C/N (NlogioN— VnilogioUi), i where p, is the probability of encounter- ing species /, C is the constant 3.321928 when using Base 2, N is the total num- ber of individuals, and n, is the number of individuals of species /. The fonnula describes an infinitely large population and results in the average diversity per species. The ecological data recorded in the field, plus food and size recorded in the laboratory, were coded in the following categories. A. Habitat: 1. Primary forest 2. Secondary forest 3. Forest-edge 4. Aquatic B. Microhabitat: 1. Fossorial 2. Ground 3. Leaf-litter 4. Bush (<1.5m) 5. Tree 6. Water EQUATORIAL HERPETOFAUNA 293 C. Activity Period: 1. Nocturnal 2. Nocturnal and Diurnal 3. Diurnal (shade) 4. Diurnal (sun) D. Activity: 1. Sleeping 2. Resting 3. Moving 4. Feeding 5. Calling 6. Mating E. Food: 1. Number of prey items in each taxonomic category 2. Size of largest prey item F. Size: 1. Snout-vent length For purposes of the computed similarity matrices and resulting distance pheno- grams, the modal code was used for each species, unless more than 25 per- cent of the individuals were found out- side of the mode; in those cases combi- natorial codes were used. Snout-vent lengths were coded by size classes: 1) ± 25 mm, 2) 26-50 mm, 3) 51-100 mm, etc. Prey items were coded by taxo- nomic category, except that arthropods were coded as "small" or "large"; ants and termites were coded separately in those species that feed exclusively on those items. For the anurans, five addi- tional parameters were analyzed: 1) mode of reproduction (see Table 10); 2) calling site of males coded to modal site; and 3) three parameters of the mat- ing call - note repetition rate, pulse rate, and dominant frequency. The pheno- grams were constructed from un- weighted arithmetic means. A general view of the community structure can be obtained by analyzing the three major taxonomic groups (anu- rans, snakes, lizards) independently (Figs. 172-174). Species of like size, habitat, diel activity, and food are grouped in these phenograms. Thus, amongst the lizards (Fig. 172), the small forest-floor inhabitants are grouped at the top of the phenogram; arboreal spe- cies are in the middle, and larger ter- restrial species are at the bottom. In the analysis of the snakes (Fig. 173), we find that most of those species in the upper half of the phenogram are diurnal terrestrial species, those just below the middle are primarily noctinnal arboreal species, and those at the bottom are aquatic. The introduction of reproduc- tive parameters into the anuran analysis provides additional refinement (Fig. 174), so that most taxonomic groups fall out together in the phenogram - thus, all of the microhylids, dendrobatids, and centrolenids are grouped with other members of their families. On the other hand, larger and more diverse families have representatives in different parts of the phenogram. For example, large hy- lids are separated from the small pond- breeding species, which are adjacent to the phyllomedusines, and all are widely separated from the carnivorous Hemi- phractus. Likewise, among the lepto- dactylids, the groups that have aquatic larvae are separate from the small bush- dwelling Eleutherodactyhis, which are disjunct from the larger terrestrial spe- cies. These phenograms show the ecologi- cal relationships of any given species with other members of the same taxo- nomic category, but their infonnation content is limited in that similarities are exaggerated. If similar data from Africa or Papua were interjected into the anu- ran phenogram, individual clusters would contain species from widely dis- parate regions, so that we would be analyzing the ecological similarities of allopatric species. To a lesser extent, the same thing is true of the analysis of the fauna at Santa Cecilia. Due to differ- ences in habitat use there is ecological allopatry among the species in the com- munity. Thus, it is necessary to examine the differences among ecologically sym- patric species. 294 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 1 r 1 \ \ r - — l- -c BACHIA TRINASALE NEUSTICURUS ECPLEOPUS PRIONODACTYLUS MANICATUS ALOPOGLOSSUS COPII ALOPOGLOSSUS ATRIVENTRIS PTYCHOGLOSSUS BREVIFRONTALIS ARTHROSAURA RETICULATA IPHISA ELEGANS PRIONODACTYLUS ARGULUS PSEUDOGONATODES GUIANENSIS ANOLIS FUSCOAURATUS ANOLIS TRACHYDERMA LEPOSOMA PARIETALE ANOLIS ORTONI GONATODES CONCINNATUS ANOLIS CHRYSOLEPIS ANOLIS PUNCTATUS THECADACTYLUS RAPICAUDA ENYALIOSAURUS LATICEPS URACENTRON FLAVICEPS ANOLIS TRANSVERSALIS PLICA UMBRA TUPINAMBIS TEQUIXIN DRACAENA GUINANENSIS POLYCHRUS MARMORATUS ENYALIOSAURUS COFANORUM KENTROPYX PELVICEPS AMEIVA AMEIVA MABUYA MABOUYA ,4 .6 SIMILARITY 1.0 Fig. 172. — Modal resource utilization by lizards based on five parameters: habitat, vertical distribution, diel activity, food and size. HABITAT ANALYSIS Five major habitats can be discerned at Santa Cecilia - primary forest, secon- dary forest, forest edge, clearings, and aquatic sites. The latter includes all types of pennanent and temporary ponds, streams, lakes, and rivers, all of which are interspersed within and be- tween the terrestrial habitats and, thus, are not comparable with the terrestrial habitats. Only exclusively aquatic spe- cies are considered to be aquatic inhabi- tants; these include one amphibian and 11 reptiles: one anuran (Pipa pipa), four snakes {FAinectes imirinus, Ilelicops angulatus, II. petersi, and Micrurus siiri- namen.sis), the two crocodilians, and five turtles (all except Geochelone) . Among the terrestrial habitats we find the greatest number of species in the primary forest. Thirty-one species ap- parently are restricted to that habitat, whereas 12 are restricted to secondary forests, four to forest edges, and 13 to clearings. Comparisons of the inhabi- tants of the four terrestrial habitats were made by using Whittaker's (1970) meas- ure of the coefficient of community (Ta- ble 18). As would be expected, the greatest similarity exists between primary and secondary forests (CC=0.6S0). Gener- ally, the secondary forest contains a slightly depauperate primary forest fauna. The apparent presence of cer- tain species only in secondary forest may be an artifact of sampling; some of these species are not well represented in the collection. Others, such as Eleuthero- dactijhis variahilis, Ilyhi Inevifrons, EQUATORIAL HERPETOFAUNA 295 1 1 1 1 1 1 1 1 1 — - 1 I - L 1 — 1 1 1 1 1 1 1 1 1 OXYRHOPUS PSEUDOBOA RHADINAEA DRYMOBIUS MELANOGENYS CORONATA BREVIROSTRiS RHOMBIFER DRYMOLUBER DICHROUS LEIMADOPHIS REGINAE DENDROPHIDION DENDROPHIS CHIRONIUS MULTIVENTRIS CHIRONIUS FUSCUS CHIRONIUS SCURRULUS CHIRONIUS CARINATUS ERYTHROLAMPRUS AESCULAPII MICRURUS SPIXll MICRURUS LEMNISCATUS PSEUSTES SULPHUREUS LEPTOPHIS AHUETULLA LEIMADOPHIS SP. MICRURUS LANGSDORFFI BOTHROPS CASTELNAUDI LACHESIS MUTA XENODON SEVERUS BOA CONSTRICTOR SIPHLOPHIS CERVINUS OXYRHOPUS FORMOSUS OXYRHOPUS PETOLA CLELIA CLELIA DREPANOIDES ANOMALUS LIOPHIS COBELLA LEPTODEIRA ANNULATA TRIPANURGOS COMPRESSUS IMANTODES CENCHOA DIPSAS CATESBYI DIPSAS PAVONINA DIPSAS INDICA OXYBELiS ARGENTEUS BOTHROPS BILINEATUS CORALLUS CANINUS CORALLUS ENYDRIS IMANTODES LENTIFERUS TANTILLA MELANOCEPHALA LEPTOMICRURUS NARDUCCll AMPHISBAENA FULIGINOSA ANILIUS SCYTALE NINIA HUDSON! XENOPHOLIS SCALARIS ATRACTUS MAJOR ATRACTUS OCCIPITOALBUS ATRACTUS FLAPS BOTHROPS ATROX EPICRATES CENCHRIA MICRURUS SURINAMENSIS HELICOPS ANGULATUS HELICOPS PETERSI EUNECTES MURINUS 1.0 Fig. 173. 2 .4 6 .8 SIMILARITY -Modal resource utilization by snakes and aniphisl:)aenians, based on four parameters: habitat, vertical distribution, diel activity, and food. 296 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY J L L 1 — 1— , I I '-- 1 1 L 1 1 PI PA PI PA BUFO MARINUS BUFO GLABERRIMUS HYLA BOANS NYCTIMANTIS RUGICEPS PHRYNOHYAS VENULOSA PHRYNOHYAS CORIACEA OSTEOCEPHALUS TAURINUS HYLA GEOGRAPHICA RANA PALM! PES CERATOPHRYS CORNUTA CHIASMOCLEIS ANATIPES CHIASMOCLEIS VENTRIMACULATA CHIASMOCLEIS BASSLERI HAMPTOPHRYNE BOLIVIANA CTENOPHRYNE GEAYI PHYSALAEMUS PETERSI LEPTODACTYLUS STENODEMA Y, y rC r LEPTODACTYLUS LEPTODACTYLUS LEPTODACTYLUS LEPTODACTYLUS LEPTODACTYLUS LITHODYTES ADENOMERA ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS ELEUTHERODACTYLUS CENTROLENELLA CENTROLENELLA J I I \ I I L .6 ILARITY 8 .0 PENTADACTYLUS RHODOMYSTAX WAGNERI MYSTACEUS DISCODACTYLUS LINEATUS ANDREAE ACUMINATUS ORPHNOLAIMUS LACRIMOSUS QUAQUAVERSUS ALTAMAZONICUS MARTIAE CROCEOINGUINIS VARIABILIS PAULULUS PSEUDOACUMINATUS LANTHANITES CONSPICILLATUS OCKENDENI DIADEMATUS MIDAS MUNOZORUM CENTROLENELLA RESPLENDENS Fig. 174. — Modal resource utilization by anurans, based on five parameters: habitat, vertical distribution, diel activity, food and size ( taxa indicated by dashed lines ) . Those taxa indicated by solid lines are arranged on ten parameters, in addition to the five above: mode of repro- duction, calling site, note repetition rate, pulse rate, and dominant frequency of mating call. EQUATORIAL HERPETOFAUNA 297 A A ^ -c HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA HYLA LANCIFORMIS CRUENTOMMA GARBEI TRIANGULUM SARAYACUENSIS BIFURCA ROSSALLENI RIVEROl PUNCTATA BOKERMANNI MINUTA RHODOPEPLA BREVIFRONS MARMORATA SPHAENORHYNCHUS CARNEUS SPHAENORHYNCHUS EURHOSTUS HYLA GRANOSA ALBOGUTTATA FASCIATA CALCARATA PARVICEPS RUBRA FUNEREA FAVOSA LEUCOPHYLLATA OSTEOCEPHALUS LEPRIEURII OSTEOCEPHALUS BUCKLEYI PHYLLOMEDUSA TOMOPTERNA PHYLLOMEDUSA TARSIUS PHYLLOMEDUSA VAILLANTl PHYLLOMEDUSA PALLIATA SYNCOPE ANTENORI EDALORHINA PEREZI ELEUTHERODACTYLUS NIGROVITTATUS ELEUTHERODACTYLUS SULCATUS ISCHNOCNEMA QUIXENSIS US PICTUS FEMORALIS MARCHESIANUS SAULI QUINQUEVITTATUS PARVULUS DENDROPHRYNISCUS MINUTUS HEMIPHRACTUS PROBOSCIDEUS BUFO TYPHONI DENDROBATES PHYLLOBATES COLOSTETHUS COLOSTETHUS DENDROBATES DENDROBATES .4 6 SIMILARITY 8 1.0 298 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY o o (D 05 o 0) 0) ,o c« tn u 102 8 8 118 8 11 065 12 8 083 .285 24 Table 18. — Comparison of the Herpetofauna in the Four Terrestrial Habitats at Santa Cecilia. (Boldface numbers are the numbers of species occurring in a given habitat; Roman numbers are the numbers of species shared between two habitats, and italicized numbers are the com- munity coefficients. See text for explanation.) in > a s •l-H PL, Primary Forest 134 Secondary Forest .. .680 Forest Edge .058 Clearings .053 Hijhi criienf omnia, Ilijla rliodopcpki, and Imantodes lentifcnis are well represent- ed in samples from secondary forest and absent in samples from primary forest and clearings. Eight species have been found in pri- mary and secondary forests and in clear- ings; these include five frogs (Lepfochic- tijlus mystaceus, L. ivagneri, Ilijhi hi- fiirca, H. garbei, H. loncifonnis) and three snakes (Chironius scurndtis, Oxy- rlwpus petola, Rhadinaea hrevirostris) . One lizard (Tupinand)is tcquixin) and two snakes {Boa constrictor, BotJirops atrox) occur in secondary forest and clearings. These eleven species are the habitat generalists in the community. Primary Forest The primary forest is the most stable habitat at Santa Cecilia, and it can be viewed as the climax vegetation type in the area. Even so, it is a patchy envi- ronment in that differences exist in the amount of moisture and in the amount of shade. Slight variations in topography result in higher, better drained terrain with interspersed depressions that may contain water after heavy rains. Fur- thermore, the forest floor is dissected by numerous small permanent or intermit- tent streams. Tree falls create temporary areas receiving full solar radiation. As noted previously, the primary for- est is the richest habitat in numbers of species of amphibians and reptiles. The large fauna of 134 species can be divided conveniently into two assemblages - spe- cies that are terrestrial and those that are arboreal. Each of these assemblages can be subdivided into noctinnal and diurnal species. The following analysis excludes the fossorial caecilians and the aquatic turtles and crocodilians. Only one ter- restrial turtle (Geochelone dentictdata) occurs in the primary forest; it is diurnal and herbivorous. Each species is cate- gorized by place and time of activity. Some species that occur on the ground and on bushes are included in two analyses; likewise, species that are active by day and night are included as mem- bers of both diurnal and nocturnal as- semblages. On the other hand, species that are active on the ground but sleep in bushes and trees are treated in the terrestrial category. Looking first at the terrestrial sub- community in the primary forest, we find 43 diurnal and IS noctiu-nal species (Figs. 175-176). The principal groups comprising the diurnal assemblage are microteiid lizards (9 species), dendro- batid frogs (5 species), leptodactylid frogs (6 species), racer-like snakes (8 species), and litter-dwelling snakes (8 species ) . Although the phenogram ( Fig. 175) presents a general picture of the ecological relationships of the diurnal, terrestrial subcommunity in the primary forest, it is instructive to examine more closely the relationships of some groups within the subcommunity. All of the microteiid lizards at Santa Cecilia are diurnal and inhabit primary forest. One species is fossorial (BacJiia trinasale ) ; the others are active on the forest floor (Fig. 177). Although these nine species are all similar in size and general habitat, they do exhibit differ- ences in microhabitats and in food. Neu- sticurus occurs only along the margins of streams, and Alopo'^lossus copii prefers depressions that are more moist than the EQUATORIAL HERPETOFAUNA 299 -c: £Z ± ALOPOGLOSSUS ATRIVENTRIS ALOPOGLOSSUS COPII ARTHROSAURA RETICULATA IPHISA ELEGANS PTYCHOGLOSSUS BREVIROSTRIS NEUSTICURUS ECPLEOPUS ANOLIS FUSCOAURATUS PRIONODACTYLUS ARGULUS PSEUDOGONATODES GUIANENSIS EDALORHINA PEREZI ELEUTHERODACTYLUS SULCATUS LEPOSOMA PARIETALE PRIONODACTYLUS MANICATUS ATRACTUS OCCIPITOALBUS TANTILLA MELANOCEPHALA XENOPHOLIS SCALARIS ANOLIS CHRYSOLEPIS BUFO TYPHONIUS ELEUTHERODACTYLUS NIGROVITTATUS COLOSTETHUS MARCHESIANUS PHYLLOBATES FEMORALIS DENDROBATES PICTUS DENDROBATES QUINQUEVITTATUS DENDROPHRYNISCUS MINUTUS DENDROBATES PARVULUS ADENOMERA ANDREAE LEPTODACTYLUS DISCODACTYLUS LITHODYTES LINEATUS ENYALIOIDES COFANORUM KENTROPYX PELVICEPS CHIRONIUS MULTIVENTRIS CHIRONIUS FUSCUS DENDROPHIDION DENDROPHIS LEIMADOPHIS REGINAE DRYMOBIUS RHOMBIFER DRYMOLUBER DICHROUS MICRURUS LEMNISCATUS ERYTHROLAMPRUS AESCULAPII MICRURUS SPIXII PSEUSTES SULPHUREUS LEIMADOPHIS SP. ATRACTUS FLAPS ATRACTUS MAJOR .0 Fig. 175. 1,0 2 4 .6 .8 SIMILARITY -Modal resource utilization by diurnal terrestrial species in primary forest, based on substrate, shade versus sun, size, and food. higher areas characteristically inhabited by the other microteiids. Among the latter, Leposoma parietale and Priono- dactylus tnaniculatiis are heliophiles, whereas the other five species usually are active only in deep shade. There are some obvious diff^erences in the prey eaten by the microteiids (Table 19). Taking into account various physical en- vironmental parameters, plus food, niche overlap values were calculated by means of the formula proposed by Colwell and Futuyma (Table 20). In the microteiid lizards the extreme mean overlap values of 35.69 (AIopo- glossus atriventris) and 0.03 (Ptijcho- 300 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY -cz -d ANILIUS SCYTALE BOTHROPS CASTELNAUDI OXYRHOPUS FORMOSUS LACHESIS MUTA XENODON SEVERUS ISCHNOCNEMA QUIXENSIS LEPTODACTYLUS RHODOMYSTAX LEPTODACTYLUS PENTADACTYLUS CHIASMOCLEIS ANATIPES CHIASMOCLEIS BASSLERI CHIASMOCLEIS VENTRIMACULATA HAMPTOPHRYNE BOLIVIANA LEPTODACTYLUS DISCODACTYLUS LITHODYTES LINEATUS CTENOPHRYNE GEAYI PHYSALAEMUS PETERSI NINIA HUDSON! ADENOMERA ANDREAE .0 Fig. 176.— Modal 1.0 4 .6 8 SIMILARITY resource utilization by nocturnal terrestrial species in primary forest, based on substrate, size, and food. glossus hrevifrontalis) are of limited meaning due to the absence of data on food. Among the other species, the fos- sorial Bachia trinasale has the lowest overlap value; with the other species, all of which are terrestrial. The other three terrestrial species active in deep shade have overlap values of 8.96-19.74 (x=12.69), and the two heliophilous species have an overlap value of 17.73. Excluding from consideration the two species with incomplete data we find that all pair-wise comparisons have val- ues of less than 20. Thus, there is less than 20 percent overlap in total resource utilization by any pair of species of microteiid lizards. Mean overlap values for other terrestrial forest lizards with the microteiids are: Pseudogonatodes gidanensis (S.32, Anolis chrysolepis 2.23, Enyalioides cofanorum 6.69, and Kentro- pyx pelviceps 2.09. Thirteen species of frogs are active on the forest floor by day. These include five dendrobatids ranging in snout- vent length from 14.0 to 24.5 mm (mean lengths of males), six leptodactylids hav- ing snout-vent lengths of 18.5-39.3 mm, and two bufonids - Dendrophryniscus minutus (15.1 mm) and Bufo typhoniiis (58.7 mm). All of these are found most frequently amidst leaf litter; Leptodac- Table 19.- —Comparative Major Dietary Items (by volume) in Microteiid Lizards. Food Species N Primary Secondary Alopoglossus atriventris 0 Alopoglossus copii 10 Arthwsaura reticulata 6 Bachia trinasale 6 Iphi.sa elegans 1 Leposoma parietale 38 Neusiicurus ccpleopus 25 Prionodacti/his ara,uJus 20 Prionodactyhts inanicutus 11 Ftrjchofjjossus hrevifrontalis () Arachnida (47%) Orthoptera (48%) Coleoptera (55%) Ordioptcra (100%) Formicidae (21%) Orthoptera (31%) Arachnida (46%) Arachnida (60%) Ordioptera (30%) Isopoda (33%) Oligochaeta (36%) Hemiptera (21%) Arachnida (20%) Orthoptera (27%) Orthoptera (22%) EQUATORIAL HERPETOFAUNA 301 Fig. 177. — Microhahitats of ten species of microteiids at Santa Cecilia. A. Alopoglossus atriventris, B. Alopoglossus copii, C. Aiihrosauia reticulata, D. Bachia tiinasale, E. Iphisa elegans, F. Leposoma parietale, G. Neusticunis ecpleopus, H. Prionodacttjlus argulus, I. Priono- dactijlus manicatus, J. Pttjchoglossus brevifrontalis. Table 20. — Niche Overlap Values for Microteiid Lizards. (Product values X 100 of four overlap values — habitat, substrate, vertical distribution, food; see text for e.xplanation.) i; a (3 5i •«2 ^ S a O o t; !» « u c C^ CO ^ o^ ^ 5S a a Q Co 00 -^ tXl CO J 5 Cud tJO o p -2 a a. ^ r^ ti o 1r r^ PQ a a o CO a o a o o c o o c o v. to O O Alopoglossus atriventris'^ .... Alopoglossus copii 9.86 Arthrosaura reticulata 100.00 2.66 Bachia trinasale 10.98 0.00 0.00 Iphisa elegans 64.00 3.84 8.96 0.00 Leposoma parietale 42.53 8.36 11.90 0.74 3.05 Neusticurus ecpleopus 4.97 4.30 1.09 0.21 1.69 4.15 Prionodactijlus argulus 44.82 19.74 15.05 0.51 9.38 31.62 5.74 Prionodacttjlus manicatus .... 45.06 14.00 11.72 0.07 6.11 17.73 2.20 43.02 Pttjchoglossus brevifrontalis^ 0.00 0.00 0.00 0.02 0.00 0.13 0.00 0.09 0.00 Mean Overlap 35.69 6.97 16.82 1.39 10.78 13.28 2.71 18.89 15.55 0.03 " Exclusive of food. 302 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Table 21. — Comparative Major Dietary Items in Diurnal Terrestrial Frogs in Primary Forest. Species N Bufo ttjphonius 25 Lithodytes lineatiis 9 ElcuthcrodactyJus sulcatus 15 EdaJorhina perezi 4 Leptodactijlus discodactyJus 8 Adenomera andrcae 11 Phyllohates femoralis 21 Dcndrohates pictus 26 Dendiobates paivulus 23 Eleutlicrodacfyhis nigrovittatus 4 Colostcthtis marchcsianus 25 Dciidrophryniscus inimitus 25 Dendiobates quinqucvittatus __ 1 Size Ratio Food Primary Secondary 1.49 1.16 1.00 1.30 1.04 1.02 1.21 1.04 1.05 1.07 1.15 1.08 Formicidae (88%) Formicidae (42%) Orthoptera (48%) Diptera (50%) Coleoptera (41%) Formicidae (35%) Arachnida (15%) Fomiicidae (70%) Formicidae ( 62% ) Coleoptera (39%) Formicidae (33%) Fonnicidae (87%) Formicidae (100%) Coleoptera Hemiptera Chilopoda Orthoptera Orthoptera Ortlioptera Coleoptera Coleoptera Coleoptera Formicidae Coleoptera Coleoptera (10%) (12%) (20%) (25%) (12%) (19%) (14%) (10%) (18%) (22%) (28%) (3%) tijlus discodactyJus inhabits swampy areas, but the other species character- istically occur on higher ground. Al- though ants are the most common item in the diets of these 13 species, compris- ing 41 percent of tlie volume of food (Table 21), niche overlap values for prey items in the 13 species vary from 0.050 to 0.875 (x=0.4S2) (Table 22). Analysis of only the five species of den- drobatids provides mean niche overlap values of 0.43S to 0.650; these values are exceeded by the overlap of Dcndro- plirt/nisciis minutus with the five dendro- Ixitids - 0.535-0.825 (.x=0.661). Food data are available for 11 of the 16 species of snakes, each of which is narrow in its dietary preferences: in- sects 1, earthworms 3, frogs 3, lizards 2, snakes 1, mammals 1. The nocturnal terrestrial assemblage in the primary forest consists of 12 frogs and six snakes (Fig. 176). The frogs comprise four groups: 1) the large beetle-eating microhylid Ctenophnjne geaiji, 2) the four small primarily ant- eating microhylids, 3) the small termite- eating leptodact\'lid Physcdaetnus pe- tersi, and 4 ) the six larger leptodactylids that eat a wide range of arthropods. The four small microhylids primarily eat ants, but they also ingest some Coleoptera, Dermaptera, and Isoptera. Niche over- lap values for food in these four species are 0.450-1.000 (.x=0.699). Hampto- pliryne and the three species of Chias- mocleis have been found in the same small area (± 1 hectar) on the same night. The six species of leptodactylids range from the small Adenomera andreae (average snout- vent length, S S ^25.1 mm) to Leptodactyhis pentadactylus (x= 154.5). Niche overlap values for food in these six species are 0.125-0.720 (x=0.393). Of the nocturnal terrestrial snakes, one feeds on mammals, one on lizards, one on caecilians, and two on frogs. The analysis of the large arboreal herpetofauna in the primary forest is most readily accomplished by distin- guishing "bush" inhabitants from "tree" inhabitants and each category separated into diurnal and nocturnal groups. Bush inhabitants are those species that char- acteristically are active in the lower strata of woody and herbaceous vegeta- tion within 1.5 m of the ground. Tree inhabitants are those that occupy the higher woody vegetation and tree trunks. Some species occur on trees and bushes and are included in both analyses. The higher strata in the primary forest is extremely difficult to sample; the data on the amphibians and reptiles from there probably are incomplete. Diurnal bush inhabitants include five lizards and three snakes (Fig. 178). Three of the lizards (Anolis fuscoaura- tiis, A. trachyderma, and Flica timhra) EQUATORIAL HERPETOFAUNA 303 o o O ■M 1=1 o be o Q G <5 >^X3 rO C8 c s 11 4-* a C« X W 0) k4 >^ o (U <-M l-l ■4-1 X! (1) 4-* S 0) OJ C/5 Ul a > D. Cl IH 0) > O (D J3 C) ^ 1 in o o CO 0 CD 0 in CD in 0 0 CO CO 0 in in in CD in o CD_ in CO in in CD 0 CO in CO 0 in in o in CO in CD in 05 in in CO in in CO in 00 p 1—1 1—1 in CO in CD CO o CD CO o O) o in Ol 0 CO 0 in in eg 0 in in r-( CO o o in in CO o in O O o CM 0 0 0 00 in 00 (M 0 in p CD CM CO in CO in in CO in CO CO o CM in in 1— ( CO in in in CO in in tM in in 01 in CM CM 00 in o oo t> o 00 in CO in -r CD o in o 1— ( o 0:1 0 01 in 0 00 CD in 00 0 CO in CO 3 a CO s O 3 O Co 3 CJ 5a CO C a 3 a Co 3 i3 o c 3 a 3 "■■ a ^ ;= ^ iT r- Ci c„ "^ CO 3 Cr C 32 a ■S t3 GO Co CO CO ^ 35 to "S to to to CI to to "C tti »-j <; e ~a -a -■a o o Q »- >~ te '^ "^ S to to s: Q Q p ^ -^ t- CO 3 O ^^- to P4 < £ 3 ri *- - « > 3 O to <; p o c c to to 304 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY I ANOLIS FUSCOAURATUS ANOLIS TRACHYDERMA ANOLIS PUNCTATUS PLICA UMBRA POLYCHRUS MARMORATUS EPICRATUS CENCHRIA OXYBELIS ARGENTEUS LEPTOPHIS AHUETULLA .0 1.0 Fig. .2468 SIMILARITY 178. — Modal resource utilization by diurnal bush inhabitants in primary forest, based on substrate, shade versus sun, size, and food. also forage on the ground, and one snake (Epicrates cenchria) probably forages more on the ground than in bushes. Uti- lizing mean snout-vent lengths of males, we find two small lizards (AnoJis fus- coauratus, 43 A mm; A. trachy derma, 48.9 mm), two medium lizards (Anolis punctatus, 79.0 mm; Plica umbra, S3.1 mm), and one large lizard (Pohjchnis marmaratus, 115.0 mm). The two small Anolis feed on a variety of small arthro- pods and have a niche overlap value of 0.65 for food resources. AnoJis puncta- tus also preys on a variety of arthropods, usually larger insects, and has niche overlap values for food resources of 0.54 and 0.28 with A. fuscoauratus and A. trachyderma, respectively. Although all three species are preying on similar kinds of food, their actual overlap is probably much less than that indicated by the foregoing values, due to terres- trial foraging activity of the two smaller species. Plica umbra feeds exclusively on ants and has niche overlap values for food resources of 0.00-0.28 (highest with A. punctatus) with the other bush-in- habiting lizards. Polychrus marmoratus primarily feeds on large orthopterans and has niche overlap values for food resources of 0.00-0.43 (highest with A. punctatus) with the other lizards. The Anolis utilize small perches, whereas Plica and Polychrus usually are on branches. Epicrates cenchria differs from the other snakes by eating mam- mals; this species is also active at night. when it probably is feeding. Oxybelis argenteus feeds on lizards and frogs, and Leptophis ahuetulla apparently eats only frogs. The presence of nocturnal frogs in the diets of these diurnal snakes sug- gests that the snakes are capable of find- ing frogs in their diurnal retreats, or perhaps they are crepuscular in their feeding schedules. In the diurnal tree-inhabiting cate- gory, Oxybelis and Leptophis are the only snakes (Fig. 179). The small spe- cies of Anolis from the bush-inhabiting group are absent, but two larger lizards are present. Anolis transversalis (71.5 mm) feeding exclusively on orthopte- rans, and Uracentron flaviceps ( 123.0 mm) feeding only on coleopterans, oc- cur in the canopy. The other tree in- habiting lizards are restricted to trunks - saplings for Enyalioides laticeps and bases of large trees for Gonatodes con- cinnatus and Anolis ortonii. The mean product values of niche overlaps of four parameters for the ar- boreal lizards are small (Table 23). The only values exceeding 6.0 are for the overlap of the three small Anolis. Thus, with the possible exception of these three species, the diurnal arboreal liz- ards are well segregated from one anoth- er ecologically. By comparison with the diurnal ar- boreal assemblage, the noctmnal arbo- real one is much larger and consists prin- cipally of anurans rather than lizards (Fig. ISO-lSl). Many of the anurans EQUATORIAL HERPETOFAUNA 305 .0 .2 .4 .6 .8 SIMILARITY 1.0 ANOLIS TRANSVERSALIS ANGUS PUNCTATUS PLICA UMBRA POLYCHRUS MARMORATUS ENYALIOIDES LATICEPS ANOLIS ORTONII GONATODES CONCINNATUS URACENTRON FLAVICEPS OXYBELiS ARGENTEUS LEPTOPHIS AHUETULLA Fig. 179. — Modal resource utilization by diurnal tree inhabitants in primary forest, based on substrate, size, and food. found on bushes at night retreat to cover in the trees by day; this is especially true for the hylids, most of which move from trees to low vegetation around ponds after rains. Frogs of the genus EJeiitherodactyhis probably more accurately reflect the uti- lization of the bush habitat at night. Some of these species seek diurnal shel- ter amidst leaf litter on the ground, and others have diurnal retreats in the trees. However, they do not congregate at ponds for breeding; instead they remain in rather circumscribed areas. iMost in- dividuals of the 12 species of Eleuthero- ckictylus perch on leaves 0.5-1.5 m above the ground. The mean snout-vent lengths of adult males is 15.0-30.4 mm and for adult females, 18.5-44.6 mm. The size ratios fall below the scale of 1.2-1.5 proposed bv Hutchinson (1958) and Schoener (1970) (Fig. 182). All of the Eleutherodactyhis feed on small arthro- pods, and with few exceptions they are catholic in their diets, resulting in high niche overlap values for food resources (Table 24). Eleutherodactyhis ociimi- natus is the most distinctive in diet; 90 Table 23. — Niche Overlap Values for Arboreal Lizards. (Product values X 100 of four overlap values — habitat, substrate, vertical distribution, food; see text for explanation. ) c •S c o o c o U Gonatodes concinnatus Anolis fuscoaiiratus 5.67 Anolis ortonii 1.64 Anolis punctatus 0.00 Anolis irachydcrma 5.65 Anolis transvcrsalis 2.27 Enyalioidcs laticeps 3.85 Plica umbra 0.04 Uracentron -flaviccps 0.00 Mean Overlap 2.39 Co cc i CO a. •« 3 35 (a p ^ o c o rJ2 Co Co 3 o o s U ^ cS > O o U n <: sn^.ioavnbDiib -gr stifDuiumoDopudsci -^ snpipwd '[J jUJpUJ>fJO "2/ SD^UDipuvi "3 snsoiuijjDi ■7[ SUJDlUDpDjp --J simn'riuiojjo.io '^ sn^uiijjulsuoj '3 snojuozDuiv^D '3 SIl^DUlUlllJV '3 00 o o in 1— 1 1— 1 in in o o CO t— 1 o CO o o o in CO in CO CO in CO 00 CO )n 00 o 00 CO in o in o in in 05 T -f O Ol 05 Tf^ i> CM "^ in "^ o in o o o in '-I I— ( CO -f CO O l^ Cv| in in in "^ i> in in oinoinoinom CDOOCO^COTt'fM'-H cDTticocococoinin oininooinoinco t^cooonicooocjicM^ inininincooiincom inoooinooooco c^icJicocooinfO'— iM< inco-toort-) a w > O < w h-ltqtqtqijjiiltiltqfjJtqtqKl EQUATORIAL HERPETOFAUNA 309 45 CO LJ a 40 cn a: UJ X h- UJ 35 C/) u. 30 o 25 m 20 I I- Z) o ^ '5 • MALES OFEMALES 15 20 25 30 35 40 45 SNOUT-VENT LENGTH OF NEXT LARGEST SPECIES Fig. 182. — Ratios of body lengtlis of succeeding larger nocturnal arboreal species of Eleuthcio- dactijlus in primary forest. The lines delimit die ratios of 1.2-1.5 suggested by Hutchinson (1958). pling. Among the 26 species in this as- semblage (IS frogs, 7 snakes, 1 hzard), three species were not in the bush-in- habiting subcommunity; these are: 1) the only nocturnal lizard, Thecadactyhis rapicaiida, which feeds on large arthro- pods, mostly roaches, on tiiniks and limbs of large trees; 2) the hylid frog Nyctimantis rugiceps, feeding entirely on orthopterans and inhabiting cavities in trees and large bamboo; and 3) the boid snake CoraUus caninus, which feeds on small arboreal mammals. Most taxonomic groups inhabiting the primary forest are either terrestrial or arboreal. Some Eleuflicrodoctyhis are active on the ground, but most are ar- boreal. The six species of Anolis pro- vide a notable exception and occur from the ground to the canopy (Fig. 183). Anolis transversaJis occurs exclusively in the trees, whereas A. punctatiis occurs in trees and bushes. Examples of both species have been obtained from the same tree. Anolis fuscoauratus and A. trachydernia forage in low bushes and on the ground, and A. ortonii inhabits the bases of trees and forages on the sur- rounding ground. Anolis chrysolepis is strictly terrestrial. The most common 310 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY transversalis trachyderma Fig. 183. — Microliabitats of six species of Anolis in primary forest. prey items of the six species include four groups of arthropods (Table 25). These data and the niche overlap values ( Table 23) show that among the species of AnoJis, fuscoauratus and tracliy derma are more nearly alike than are the others. Comparisons of sites of captiue and rela- tive numbers of individuals ( see account of A. trachyderma for data) suggest that A. trachyderma is more abundant in secondary forest than is A. fuscoauratus; perhaps the latter is less tolerant of sun- light and more susceptible to desiccation than is A. trachyderma. Thus, their pri- mary foraging zones might be different but probably overlap broadly; both spe- cies have been observed in the same bush and foraging within a meter of one another on the ground. Secondary Forest As noted previously, most of the 118 species known to occur in secondary forest also inhabit primary forest. Of those species that have been collected only in secondary forest, the following are represented by no more than three specimens each: BoUtoglossa equatori- Table 25. — Comparative Major Dietary Items (by volume) in Anolis. Food Species N A. chnjsolepis .^ _ _,.. 25 A. fuscoauratus ___ 25 A. ortonii .._. 7 A. punctatus ___. 14 A. trachyderma _ _ 27 A. transversalis ____ 3 Primary Secondary Orthoptera (28%) Orthoptera (25%) Formicidae (54%) Orthoptera (36%) Arachnida (42%) Coleoptera (100%) Coleoptera (16%) Arachnida (24%) Orthoptera (21%) Formicidae (29%) Orthoptera (10%) EQUATORIAL HERPETOFAUNA 311 .0 .4 6 8 SIMILARITY 10 AMEIVA AMEIVA MABUYA MABOUYA TUPINAMBIS TEQUIXIN MICRURUS LANGSDORFFI OXYRHOPUS MELANOGENYS PSEUDOBOA CORONATA RHADINAEA BREVIROSTRIS CHIRONIUS SCURRULUS BOTHROPS ATROX Fig. 184. — Modal resource utilization by diumal reptiles in clearings based on size, substrate, and food. ana, Ceratophrijs cormita, Leptoclacty- his stenodcma, and CoraUus enijdris. The scant amount of data on these species does not preclude their existence in pri- mary forest. Of the remaining six species appar- ently confined to secondary forests, four are small bush-dwelling species of Hijla (brevifrons\ cruenfomma, rhoclopepki, and riveroi) that breed in ephemeral ponds. The very abundant, small EIcu- tJierodactyhis variabilis also is a bush inhabitant. The last species is a bush- dwelling snake, Imantodes lentiferus. All of the species apparently confined to secondary forest are nocturnal. Only Ceratophrijs cormita and Leptodacfi/Jus stenodema are terrestrial. Ceratophrijs is a large predatory species, consuming large arthropods, frogs, and small mam- mals; the salamander and the other frogs eat small arthropods. CoraUus enijdris and Imantodes lentiferus eat frogs. Clearings By comparison with the fauna in the forests, the subcommunity inhabiting clearings is small (24 species) and rela- tively simple to analyze. Although some of the noctiunal frogs do perch on Ixishes and herbs, all species are consid- ered to be terrestrial. They are readily identified into two assemblages - diurnal (9 species) and nocturnal (16 species); Bothrops atrox is a member of both as- semblages (Figs. 184-185). Four am- phibians (Bufo pjaherrimus, B. marinus, HijIa rubra, and H. triangiduni), two lizards (Ameiva anieiva and Mabuija inaboiiya) , and seven snakes CleJia cle- lia, Drepanoides anomahis, Liophis co- bella, Micrurus langsdorffi Oxijrhopus inelanogenys, Fseudoboa coronata, and Sipldophis cervinus) apparently are re- stricted to clearings. Of all the assemblages analyzed, the diurnal group in clearings is unique in the absence of anurans (Fig. 184). This assemblage consists of three lizards and six snakes. All of the lizards are helio- philous; Ameiva and Mabuya prey on a variety of arthropods and Tupimnnbis is omnivorous. The diet of Micrurus langs- dorffi is unknown, but if it is like other terrestrial Micrurus it feeds on snakes and lizards. Chironius fuscus ferrets frogs out of their diurnal retreats, and the other snakes feed on lizards; Both- rops atrox also eats frogs and mammals, both of which it probably captures at night. In the nocturnal assemblage, there are nine frogs and seven snakes (Fig. 185). The frogs contain three ubiquitous species (Bufo marinus, Hyla lanciformis and Hyla rubra) that usually can be observed on any night of the year. The five species of Hyla and Leptodactylus mystaceus congregate at ephemeral ponds, whereas the Bufo and Leptodac- tylus ivagneri are scattered throughout the clearings. All of the anurans feed on 312 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ± .0 .4 6 8 SIMILARITY 1,0 HYLA GARBEI HYLA BIFURCA HYLA TRIANGULUM HYLA LANCIFORMIS HYLA RUBRA LEPTODACTYLUS MYSTACEUS LEPTODACTYLUS WAGNERI BUFO GLABERRIMUS BUFO MARINUS SIPHLOPHIS CERVINUS OXYRHOPUS PETOLA LIOPHIS COBELLA DREPANOIDES ANOMALUS CLELIA CLELIA BOA CONSTRICTOR BOTHROPS ATROX Fig. 185. — Modal resource utilization by nocturnal amphibians and reptiles in clearings, based on size, substrate, and food. a wide variety of arthropods. Of the snakes, only Liophis is restricted to frogs, Bothrops, Boa, and CleJia feed on small mammals and lizards; the fomier also eats frogs and the latter snakes. Oxy- rhopus petola and Sipljlopliis apparently seek out lizards in their nocturnal re- treats. One Clelia was eating an Ameiva when captured at night. According to Bailey (1966:68), Drepanoides eats rep- tile eggs. DIEL ACTIVITY The herpetological community at Santa Cecilia contains more nocturnal than diurnal species (Table 26); most of the anurans are nocturnal, hut only one (Thecadactyhis rapicauda) of 30 lizards is nocturnal. The strictly diurnal anurans occur in the forest and consist of all six species of dendrobatids, two bufonids (Bufo typhonius and Dendrophryniscus minutus), and three leptodactylids (Eda- lorhina perezi, Elcutherodactyhis nigro- vittatiis and E. sulcatus). Eight other anurans are active both at night ( ground or low vegetation) and by day (ground); these include the hylid Hemiphractus prohoscideufi and seven leptodactylids {ElcuthcrodactyJiis conspicdkitus, E. croceoingiiinis, E. lanthanitcs, E. ocken- deni, Leptodactyhis discodactylus, Lith- odyte.s lineatus, and Adenomera an- dreae). The three snakes that are active by day and night include the terrestrial Bothrops atrox, the forest-dwelling Epi- crafes cenchria, and the aquatic Helicops angidatus. Analysis of habitat and vertical dis- tribution with respect to diel activity re- veals that most terrestrial species are di- urnal, whereas most arboreal species are nocturnal (Table 27). The iguanid liz- ards are a major exception to this gen- erality; all iguanids are diurnal, and nine Table 26. — Diel Activity of Taxonomic Groups in the Herpetofauna. Group Diurnal Nocturnal Both Salamanders 0 (0%) 2 (100%) 0 (0%) Anurans 11 (13%) 68 (78%) 8 (9%) Lizards 29 (97%) 1 (3%) 0 (0%) Snakes 26(49%) 24 (45%) 3(6%) TOTALS 66 (39%) 95 (55%) 11 (6%) EQUATORIAL HERPETOFAUNA Table 27. — Diel Activity in Relation to Habitat and Vertical Distribution. ( Numbers are amphibians/reptiles. ) Diurnal Nocturnal Habitat Arboreal Terrestrial Arboreal Terrestrial Secondary Forest ..— _ 4/12 16/40 49/11 16/8 Primary Forest _ 3/8 12/32 53/12 15/8 Clearings 0/0 0/9 5/0 4/7 313 of the 11 species are at least partly ar- boreal. Some diurnal terrestrial species sleep at night on herbs or bushes. The anurans Dendrobates (3 species), Dendrophry- niscus, and Biifo typhonius usually sleep on leaves of herbs < 0.5 m above the ground, but the Bufo sometimes utilize perches > 1 m. Many of the racer-like snakes (Cliironins, Dendrophidion, Dry- moluher) sleep on bushes and low trees 1-3 ni above the ground. All diurnal amphibians shun sun- light; thus, they are present only in the forest. Eighteen of the 29 diurnal liz- ards live in deep shade. This group in- cludes all of the gekkonnids, and Anolis, and most of the microteiids. Other ter- restrial species in the forest are active in patches of sunlight; these include Kentropyx, Lcposoma, Prionodactylus manicatus, and Enycdioides cofanonim, plus FUca umbra, which also basks on tree trunks. Enyalioides Jaticeps, and FoJychrus marmoratus are arboreal helio- philcs. The large teiids Dracaena and Tupinambis are active in direct sunlight, but they also seek shade at warmer times of the day. The most conspicuously he- liophilous species are Ameiva ameiva and Mabuya mabotiya; both are active only in sunlight. No individuals of either of these common species were observed during one period of 30 days of heavy cloud cover. Fitch (1968) reported on body tem- peratures of some lizards at Santa Ce- cilia. Mean body temperatures for four heliophilous species (Ameiva, Kentro- pyx, Mabuya, and Prionodactyhis mani- catus) were 29.7-37.0 (x=34.31)°C; temperatures for five sciophilous species (Neusticurus, Gonafodes, Iphisa, Priono- dactylus argidus, and AnoJis chrysolepis) were 26.8-30.1 (x=28.3)°C. Although these data are limited to a small sample of species, it is obvious that the species that are active in the shade have lower optimal temperature requirements than do the heliophilous species. In contrast to the lizards, most di- urnal snakes are heliophiles; at least most individuals have been found in sunny situations. The optimal time for finding diurnal snakes is mid-morning on a sunny day. Mid-day temperatures (sunny days) apparently are excessive for most species. Only four diurnal snakes are known to be sciophilous; these are Atractus occipitoaJbus, Lepto- micrurus narduccii, TantiUa melano- cephala, and XenophoUs scalaris. Activity of nocturnal species is nega- ti\'ely correlated with moonlight. On nights with three-quarter to full moon there is relatively little anuran and snake activity, even though it may be raining and have a moderate amount of cloud cover. Abundant species of frogs, such as Hyla lanciformis and H. rubra, which begin calling nightly at dusk, sometimes cease calling in about half an hour on moonlit nights, whereas on dark nights they may call throughout the night. DIET AND FEEDING With a few exceptions, amphibians and lizards have generalized diets con- sisting of a wide array of arthropods. These insectivorous animals are all es- sentially secondary consumers (tertiary when the diet consists of spiders). They prey on any insect (except distasteful ones) that they can capture. Prey size is a limiting factor in that man\' tropical 314 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY insects, especially oithopterans and cole- opterans are too large to be captured, let alone ingested, by many small species of amphibians and lizards. On the other hand, larger species frequently consume many kinds of small prey. Thus, in these generalists, size of the predator is posi- tively correlated with the maximum, but not the minimum, size of the prey. Among the frogs obligate feeding on ants occurs in two species of microhylid frogs (Chiasmocleis anafipes and C. ventrimaculata), two hylids (Sphaeno- rhynchus carneus and S. curhostus) and in Dendrobates quinquevittatus. Ants also form the majority of the diet in Chiasmocleis bassleri, Hamptophryne holiviatia, and Eleutherodactylus acumi- natus. All of these frogs are small ( maxi- mum snout-vent lengths of 14.0-39.0 in males and 16.0-48.0 in females) and, with the exception of Dendrobates quin- quevittatus, have pointed snouts. The small Vhysalaemus petersi feeds only on termites. Two frogs are .specialized for con- summing large prey; Ceratophrys cor- nuta and Hemiphractus proboscideus have relatively huge heads and large gapes. These "hopping stomachs" are capable of ingesting prey nearly equal to their own bulk; both eat frogs and large orthopterans, and Ceratophrys also eats small mammals. Only two lizards are obligate in their diets. PUca umbra feeds exclusively on ants, and Dracaena guianensis eats snails. The large teiid, Tupinam])is tequixin, is omnivorous, feeding on insects, small mammals, bird eggs, and vegetation. No other species of lizard at Santa Cecilia is herbivorous. Partial analyses of major dietary items and niche overlap in food for some eco- logical groups (guilds) of amphibians and reptiles are presented in tables 19- 25. The lizards were chosen as an ex- ample of niche breadth analysis using Simpson's index of diversity for niche breadth (Colwell and Futuyma, 1971: 568; Pianka, 1972:59). Prey items from 526 lizards of 27 species were identified in 19 categories. Niche overlap values range from 0.00 to 0.86, whereas niche breadth scores are 1.00 to 18.91 (stand- ardized scores = 0.053-0.995) (Table 28). Two hundred seventy-seven (85%) of the 325 potential overlaps are realized (Cii, > 0.0) along the food dimension. Thirteen of the 27 species have over- lap values of 30-40 percent; none is more than 40 percent, and only four species are below 10 percent. Only two species have standardized niche breadth scores of more than 0.5 {Gonatodes concinna- tus 0.516, Ameiva ameiva 0.995). In contrast to anurans and lizards, snakes are more specialized in their feed- ing, although juveniles may feed on dif- ferent kinds of prey than the adults. Dentitional modifications can be corre- lated with specialized feeding in diverse taxa of snakes ( Anthony, 1955; Edmund, 1969). Most snakes feed on vertebrates, but three feed exclusively on earth- worms, three on arboreal snails, and one on centipedes. Frogs are eaten by 16 species, lizards by 14, and mammals by six. Another six food categories are dis- tributed as follows: birds 1, snakes, 4, reptile eggs 1, tadpoles 1, caecilians 1, and fish 2. These data are more meaningful when viewed with respect to structural habitat and diel activity of the snakes (Table 29). Thus, there is differential utilization of food resources at difl^erent times and in different structural habitats. SIZE RATIOS Hutchinson ( 1959 ) and Schoener ( 1965 ) noted progressive increases in size of trophic appendage ratios for mammals and birds. The sizes of the trophic appendages are positively corre- lated with body size in lizards of the genus Anolis (Schoener, 1967) and in hylid frogs (Caldwell, 1973); thus, body size ratios can be used in these groups to determine if a regular progression of body size or "biological constant" EQUATORIAL HERPETOFAUNA 315 (Schoener, 1970) exists among sympatric associations of these groups. Schoener (1965, 1969, 1970) discov- ered that body size ratios in insectivor- ous birds and anohne Hzards on West Indian islands sliow a regular progres- sion of size within a community. Gen- erally, larger ratios occurred among larger species and smaller ratios among among smaller species. Schoener (1965) suggested that this pattern may be due to the uneven distribution of the bio- mass of insects upon which the birds were feeding. Caldwell ( 1973 ) found similar patterns in communities of tree frogs and concluded that food probably is the most important factor influencing body size in tree frog communities. Size ratios of anurans and lizards at Santa Cecilia were calculated from the mean snout-vent lengths of the 10 larg- est individuals of each sex (fewer if there were less than 10 adults). The studies by Caldwell and Schoener (su- pra cit. ) each dealt with one congeneric assemblage of species. The ratios gen- erally ranged from 1.2 between smaller species to 1.5 between larger species. The ratios among the six species of Anoli.s at Santa Cecilia are 1.02-1.46 in males and 1.01-1.30 in females, but the species are not in the same size order in males and females (Table 30). Simi- lar discrepancies are found in ratios of body size in frogs inhabiting the forest floor by day (Table 21) and in nocturnal arboreal species of Eleutlierodacfijlus (Fig. 182). In fact no grouping, either taxonomic or ecological, yields the re- sults anticipated from conclusions of Hutchinson, Schoener, and Caldwell. TADPOLES Larval amphibians have a completely different suite of ecological requirements than do the adults; consequently they are subject to entirely different selective pressures. No detailed ecological studies of tadpoles were undertaken, but some general observations indicate tadpoles occur into different ecological situations. Wilbur (1972) demonstrated experi- mentally several factors significant to the coexistence of small communities. Structural differences (principally in the caudal fins and mouth) among tadpoles are adaptations for locomotion and feed- ing in different situations. For example, high dorsal fins and anteroventral mouths are characteristic of midwater grazers, and moderate dorsal fins and anterior mouths are characteristic of pelagic filter feeders, whereas low caudal fins and ventral mouths are characteristic of bot- tom grazers. The pond-dwelling tadpoles at Santa Cecilia have been grouped into three categories: vegetation-choked areas, pe- lagic, and bottom-dwellers. No more than 10 species of tadpoles were found in any one pond at one time. At a forest pond, the numbers of species of tadpoles inhabiting vegetation-choked areas, pe- lagic situations, and bottom were 2-7-1. At a forest-edge pond the numbers were 3-4-1, and at a pond in a clearing they were 7-2-1. I suspect that analysis of gut contents of tadpoles combined with more refined ecological analysis will demonstrate that the numerous species of tadpoles in- habiting forest utilize different resources in their coexistence in restricted habitats. Heyer ( 1976 ) suggested that interspe- cific competition is non-existent among tadpoles. There are few species of stream-in- habiting tadpoles at Santa Cecilia; tad- poles of CentroleneUa live under cover on the bottoms of streams, whereas tad- poles of dendrobatids graze on algae on rocks. Tadpoles of three species are unique in their habits. The tadpoles of Hijla geographica occur in large schools in lakes. Those of Hyla hoans develop in shallow nests at the edges of rivers. The carnivorous tadpoles of Ceratophrys cor- nuta inhabit shallow ponds where they prey on tadpoles. Tadpoles of Nyctiman- tis riigiceps and of the species of Den- drobates develop in constrained bodies of water (tree holes, bases of palm 316 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Table 28. — Niche Overlap Values and Niche Bieadtli Scores in Lizards and Amphisbaenians for prey eaten. P3 — ' ^ U CD 03 13 to o « o ^ 5 Q H ^ !;:; fc 0 2 Co s CO Co g 5 C/5 ^ ^ s i^ ^ 'O 0 0 hJ "^^ r::^ 0 c ^ -c C/^ C/^ Co c e .»»„ ^ 0 J-* 0 i--I ^ 5ij 0 "^ H^ 0 s 2 J3 0 "is '5 2 c c^ C/^ 0 On c^ Co ^--. ^3 ^— . ■o r-^ r^ (^ r— J e Q ?^ ^ ■o 0 0 0 0 0 0 ii a> S5 .^ ^ ^ ^ ^ ^ c c c 0 c~< <: < <: -< -^ <, lij Kl s: o. A 10 B 11 .50 C 7 .36 .30 D 26 .62 .34 .32 E 25 .53 .47 .36 .68 F 7 .37 .30 .21 .34 .41 G 14 .43 .29 .26 .54 .54 .47 H 27 .61 .31 .30 .57 .65 .31 .28 I 3 .07 .15 .18 .33 .24 .11 .24 J 6 .36 .43 .27 .36 .46 .22 .37 K 24 .50 .30 .39 .35 .43 .22 .34 L 11 .07 .08 .00 .04 .14 .56 .28 M 6 .15 .19 .18 .29 .31 .11 .33 N 1 .07 .00 .00 .07 .07 .00 .22 O 29 .56 .41 .48 .52 .55 .36 .57 P 10 .56 .42 .38 .26 .31 .25 .22 Q 106 .60 .29 .32 .63 .51 .30 .52 R 6 .43 .15 .14 .26 .21 .27 .22 S 11 .08 .28 .09 .07 .03 .00 .00 T 1 .14 .00 .45 .04 .00 .00 .06 U 84 .51 .33 .54 .54 .66 .29 .44 V 38 .61 .38 .18 .45 .47 .42 .43 W -- 25 .58 .58 .49 .50 .66 .38 .40 X 20 .68 .37 .42 .43 .52 .41 .25 Y 11 .48 .31 .49 .45 .58 .33 .35 Z 1 .20 .07 .18 .29 .24 .11 .39 AA -_.... 3 .07 .08 .10 .05 .16 .36 .28 xCi^ .393 .282 .284 .344 .391 .278 .333 B 9.80 5.46 2.69 5.32 5.50 2.80 4.44 BgT .516 .287 .142 .280 .289 .147 .234 .09 .42 .14 .62 .04 .28 .00 .00 .00 .00 .15 .86 .28 .10 .00 .06 .00 .29 .07 .00 .14 .43 .57 .39 .47 .06 .49 .52 .07 .28 .54 .00 .07 .55 .20 .32 .52 .02 .27 .21 .00 .28 .16 .00 .14 .06 .00 .14 .23 .00 .00 .03 .00 .00 .07 .00 .00 .51 .40 .31 .36 .00 .40 .36 .07 .21 .30 .23 .07 .55 .13 .45 .52 .13 .19 .68 .07 .25 .58 .04 .07 .61 .20 .29 .48 .00 .20 .15 .50 .14 .23 .00 .14 .00 .00 .14 .08 .33 .00 .347 .180 .291 .317 .076 .198 .36 1.00 4.00 4.56 1.00 1.32 .282 .053 .211 .240 .053 .070 EQUATORIAL HERPETOFAUNA 317 >* < ^^ ec ^^^ ^ X Co < 5 CO 5S. lis a O 3 O C f — . •5 •*- 00 On > e Co J Co £ CO -S 3 i .8 5b "^ rfi CO a •S "S s ^ c CJ e ,^ t— a 3 e a a r-^ a c/^ 3> 35 CO c c o g s 3 2 ■4^ .1 a. p 5 Co O a. •2 3 CD So 0 0 0 .5 "S. 3 a .CO S ;::) < < ^ eq 1"^ ;^ K-1 ;^ Ci^ ftn t^ < .06 .00 .45 .07 .54 .33 .14 .25 .27 .27 .00 .04 .00 .04 .00 .00 .10 .13 .05 .14 .00 .01 .70 .42 .46 .28 .00 .16 .00 .31 .26 .44 .28 .10 .06 .36 .06 .57 .55 .40 .22 .18 .16 .53 .47 .00 .45 .66 .42 .36 .06 .18 .59 .51 .58 .00 .49 .66 .42 .26 .01 .13 .68 .32 .48 .72 .00 .49 .14 .39 .00 .00 .00 .40 .06 .12 .06 .20 .00 .11 .01 .03 .01 .42 .01 .00 .23 .16 .04 .01 .01 .050 .397 .300 .340 .190 .070 .073 .380 .290 .400 .360 .352 .158 .010 1.00 4.23 3.69 18.91 4.45 2.28 1.00 3.77 6.33 7.01 4.40 2.78 2.00 3.00 .053 .223 .194 .995 .234 .120 .053 .198 .333 .369 .237 .146 .105 .158 318 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Table 29. — Prey of Snakes in Relation to Habitat. (Numbers are diurnal/nocturnal species of snakes.) Prey Terrestrial Arboreal Aquatic Total Eartliwomis — Snails Centipedes Fish Caecilians Tadpoles Frogs Lizards ___ _. Snakes Reptile Eggs - Birds Mammals TOTAL 3/0 0/0 1/0 0/0 1/0 1/0 8/5 5/8 3/1 0/1 0/0 2/2 24/17 0/0 0/3 0/0 0/0 0/0 0/0 3/8 2/3 0/0 0/0 0/1 1/2 6/17 0/0 0/0 0/0 1/1 0/0 0/1 0/0 0/0 0/0 0/0 0/0 1/1 2/3 3/0 0/3 1/0 1/1 1/0 1/1 11/13 7/11 3/1 0/1 0/1 4/5 32/37 Table 30. — Size Ratios in Sympatric Species of Anolis. Males xSVL Ratio Females X SVL Ratio A. punctatiis 79.0 A. chnjsolepis _.. 75.8 A. transversalis 71.5 A. trachyderma 48.9 A. ortonii 44.4 A. fuscoauratus 43.4 1.04 1.06 1.46 1,10 1.02 A. chnjsolepis 78.8 A. transversalis 72.0 A. punctatus 71.5 A. trachyderma 54.8 A. fuscoauratus 46.8 A. ortonii 42.5 1.09 1.01 1.30 1.17 1.10 fronds, bromeliads ) ; those of at least some species of Dendrobates are canni- balistic. ANURAN MATING CALLS In addition to the differential utiliza- tion of the physical and biotic resources in the environment, most anurans have another dimension in which they exhibit differences. Each species has a characer- istic mating call that differs from the calls of other syntopic species. Fouquette (1960) demonstrated the importance of the mating call as a prc-mating isolating mechanism, and Duellman (1967) elab- orated on the mating call and other courtship isolating mechanisms in ten sympatric hylids in Costa Rica. Presumably differences in mating calls are most essential in multi-species breeding congregations, such as common to many species of hylids. Accordingly, data obtained from recordings of 1-10 individuals of 20 species of pond-breed- ing hylids at Santa Cecilia were sub- jected to stepwise discriminant analvsis. Seven variables were used: 1) number of notes in call group, 2) note repetition rate, 3) number of secondary notes, 4) duration of note, 5) pulse rate, 6) funda- mental frequency, 7) dominant fre- quency (see Duellman, 1970b for defini- tions ) . In the first analysis two questions were posed: Can the 20 species be dis- tinguished on the basis of the call param- eters used? Which character(s) are most important in discriminating between spe- cies? Examination of the correlation matrix reveals that of the seven param- eters, only numbers 6 (fundamental fre- quency) and 7 (dominant frequency) are highly correlated (r=0.903). Vari- able six is the best discriminator, fol- lowed by variables seven, five, and three. At this point the model discriminates 96 percent of the individuals. The multi- variate means of the 20 species are shown in a three dimensional plot (Fig. 186), in which canonical axis I subsumes 62 percent of the dispersion in the multi- variate data cloud; axis II subsumes EQUATORIAL HERPETOFAUNA 319 Fig. 186. — Three dimensional plot of the multivariate means of call parameters of 20 species of pond-breeding hylid frogs (see text for explanation of the canonical axes). A. Hyla hifurca, B. H. hokermanni, C. H. brevifrons, D. //. edemata, E. H. cmentomma, F. H. garhci, G. H. geog.raphica, H. H. granoso, I. H. lanciformis, J. H. Icucophyllata, K. H. marmorata, L. H. paiviccps, M. H. pimctata, N. H. rhodopeph, O. H. rubra, P. H. .sarayacucnsis, Q. H. tri- angulum, R. Phyllomediisa palliata, S. P. tarsius, T. Sphacnorhynchus carneus. about 20 percent, and axis III about 8 percent. In general, the species are well sorted, but some are quite similar to one another. If only sympatric and synchronously breeding species are analysed together, similar broad sorting occurs but the spe- cies that closely approximated one anoth- er in figure 1S6 are in separate breeding congregations (Figs. 187-190). Thus, within given breeding congregations each species tends to have an individual acoustical niche. NICHE DIMENSIONS Ideally, the realized niche for each of the species in the community could be assessed by measuring numerous di- mensions (resources utilized); compari- sons of the dimensions for each species would provide a picture of the ecological relationships of the members of the com- munity. The ideal approach is not feasi- ble for the entire community; too few data are available on snakes to provide a meaningful analysis. An overall analy- sis of the anurans likewise has little merit, because data on many of the spe- cies only reflect certain parameters when the animals are in breeding congrega- tions. The data on lizards are the most suitable for multidimensional analysis. Ecological data on 1506 specimens of 28 lizards and one amphisbaenian were analyzed with respect to four dimensions of the niche: habitat (13 categories), substrate (9), microhabitat (5), and prey (19). 8 Diel activity was not in- cluded in the analysis; all of the lizards, except Thecadactyhis rapicauda, are di- 320 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY (J) (D (5) ® f Fig. 187. — Tliree dimensional plot of tlie multi\'aiiate means of call parameters of eight species of hylid frogs in a breeding aggregation at a pond in primary forest (Letters refer to species listed in legend to figure 186 ). n Q ® (z) © (e) ^ 1 (s, Fig. 188. — Three dimensional plot of tlie multivariate means of call parameters of ten species of hylid frogs in a breeding aggregation at a pond in secondary forest ( Letters refer to species listed in legend to figure 186 ) . EQUATORIAL HERPETOFAUNA 321 n (D (5) ^ ■Q ! ® Fig. 189. — Three dimensional plot of the multivariate means of call parameters of eight species of liylid frogs in a breeding aggregation at a forest-edge pond (Letters refer to species listed in legend for figure 186 ) . n (Q) ^ ■(K). (£) (0) Q) Fig. 190. — Three dimensional plot of the multivariate means of call parameters of six species of hylid frogs in a breeding aggregation at a pond in a clearing (Letters refer to species listed in legend to figure 186 ) . 322 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY iirnal. No Pohjcliriis marmoratus were found while active (all were sleeping at night), so that species was omitted from the analysis. The proportions of individuals of each species associated with each cate- gory provided the basis for quantif\'ing the niche breadth (B) along each di- mension using Simpson's index of diver- sity (Colwell and Futuyma, 1971). In order to compare niche breadths along different dimensions, each niche value was standardized (Bst) by dividing the value by the ninnber of categories in that particular dimension. The overall niche breadth (4-dimensional) was cal- culated as the mean of the four stand- ardized niche breadths (^Bst/4). Niche overlaps values (C,,, ) were computed using the formula proposed by Colwell and Futuyma, 1971). Overall niche over- lap was computed by multiplying for each pair the four overlap values and the total X 100. In this method overall niche overlap is 0 if overlap along any one dimension is 0. Frequency distributions of niche overlaps for each dimension are given in figures 191-192; frequency distributions of standardized niche breadths, numbers of overlaps in four dimensions, and mean niche overlap values are shown in figure 193. Various niche overlap values are given in Table 31. From these analyses certain features are evident. Mean niche Ijreadth is a function of sample size (r=0.42, 0.05 > P > 0.01). This correlation could be a reflection of sampling bias in the esti- mates of niche breadth, or it could mean that the more common species have broader niches (which is why they are common), or it could be a combination of the two. There is no correlation be- tween mean overlap values and mean standardized niche breadth (r= — 0.016). There is a highly significant cor- relation (r=0.62, P < 0.01) between mean niche lireadth and the number of overlaps in 4-space. Thus, lizards with broad niches tend to overlap with more 80-? 60 40 ^ 20 < CL ^200H Ijj > o 180 a: QQ 80 VERTICAL i ; i SUBSTRATE 60 40 20- 0.0 02 0.4 0.6 0.8 1.00 Fig. 191. — Number of o\erlap pairs among 29 species of lizards with respect to vertical distri- bution and substrate. species than do species with narrow niches, but they do not necessarily share more overlap space. Of the six possible comparisons among four niche dimen- sions, only habitat-substrate is significant (r=0.698, P < 0.01); thus, lizards with broad habitats also have broad substrate utilization. Relative positions of all species along a standardized niche breadth continuum are shown in Figure 194. Forest inhabi- tants ( those species for which 50 percent or more of the specimens were taken in primary forests) tend to have narrower niches (X Bst=0.232) than on-forest species (X Bst=0.190), but the differ- ences are not statistically significant. The EQUATORIAL HERPETOFAUNA 323 if) < d. < _l Ql LU > O 80 60 40 20- 0 120 100- ttr ^ 80 60 40 20 0 FOOD SITE "wm 00 02 0.4 06 08 1.00 Fig. 192. — Number of overlap pairs among 29 species of lizards with respect to food and site. mean number of overlaps in 4-.space for forest species and non-forest species does not differ significantly (14.1 and 13.0, respectively, deleting species represent- ed by only a single specimen ) . Also, the average amount of niche overlap in 4- space for forest and non-forest species (5.10 and 1.93, respectively) does not differ significantly. SPECIES DIVERSITY Quantitative comparison of commu- nities involving the numbers of species (richness) and relative numbers of indi- viduals of each species (equitability) has given rise to the concept of species diver- sity (see Margalef, 195S; MacArthur, 1965; and Pianka, 1966, for reviews). STANDARDIZED NICHE BREADTHS (x=l.92) 8- 6- 4- 2- iii n - iiii ■;■:■:■: 1' 1 > 1.0 2.0 3.0 4.0 ,fi8- NUMBERS OF OVERLAPS IN 4-SPACE Z5 o (x=l2.82) uj 6- 4 - WWl^ 2- n - WmM 1 4 7 10 13 16 19 22 25 8- MEAN OVERLAP VALUES im (x = 3.39) 6- m\ 4- 11 -:•:■:•: ■ m^ 2- 0- "+" "" , , , PI, 10 12 18 Fig. 193. — Standardized niche breadths, num- bers of overlaps, and mean overlap values among 29 species of lizards. MacArthur's (1957) "broken-stick mod- ed" was questioned by Hairston (1969), and Lloyd and Ghelardi (1964) pro- posed an equitability factor for the meas- ure of fit of observed relative abundances to those predicted by MacArthur's model. Hurlbert ( 1971 ) criticized se- mantic, conceptual, and technical as- pects of species diversity. Despite Hurlbert's (1971) critique of diversity measurements, it is evident from the work of MacArthur (1965), Pianka (1967), Rosenweig (1975), and Cody (1975) that latitudinal gradients do exist in species richness and that there are differences in equitability in different communities. A quantitative expression of differences in diversity and equitabil- ity perhaps will be a useful method of 324 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Table 31. — -Niche Overlap Values ( 4-Dimcnsiona] ) in Lizards and Amphisbaenians. C ^ 50% or more individuals in mature forest. ) No. of Mean Total Species Overlaps Overlap Overlap Gonatodes concinnatus^ 21 0.825 23.09 Fseudogonatodcs guianensis'^ .._ 12 6.320 176.90 Thecadactijlus rapicauda 2 0.001 0.04 Anolis chnjsolepis'^ 14 4.440 124.26 AnoJis fuscoauratus'' 18 5.200 145.51 Anolis ortonii" 15 3.280 91.85 Anolis punctatus 1 0.020 0.27 Anolis trachtjderma*' 11 4.960 138.87 Anolis transversalis'* 11 0.200 5.72 Enyalioides cofanorum*' 12 6.910 193.35 Enijalioides laticeps" 14 0.560 15.68 Plica umbra" 12 0.110 3.03 Uraccntron jlaviceps" 1 0.280 3.95 Mabmja mabouija 22 2.370 66.57 Alopoglossiis atriventris'' 13 17.230 482.48 Alopoglossus copii 20 3.790 106.11 Ameiva ameiva 16 1.300 36.43 Arthrosaura reticulata'* 11 6.750 188.94 Bachia trinasale" 11 0.600 16.67 Dracaena guianensis 10 2.490 69.73 Iphisa elegans" 11 3.500 98.10 Kentropyx pelviceps 17 4.190 117.78 Leposoma parietale'* ..__ 24 7.000 196.00 Neusticurus ecpleopus 16 1.290 36.08 Prionodactylus argulus'* 23 8.390 234.93 Prionodactylus manicatus'* 19 15.500 217.00 Ptijchoglosstis brevifrontalis 5 0.030 0.77 Tupinarnbis tequixin 8 0.890 24.99 Amphisbaena fuliginosa'^ 2 0.002 0.06 I c: <; lo ^ ^ ^ (i. lO lo" c^ C; "2 C ^ 1 N^ n: ^^ 4 — i^H-^ ■$ I 5 3 ^ 5 I f-ii >i) !> supply) for eff^ective competi- tion to occur. Most anurans and lizards in the com- munity at Santa Cecilia are secondary consumers feeding mostly on a wide variety of insects. With the exception of the large mollusk-eating lizard, Dracaena guianensis, the only food specialists feed on ants or termites, both of which are extremely abundant. Niche-overlap val- ues of 0.5 for prey are common among lizards and anurans (see Tables 22 and 24). Thus, due to the abundance of food throughout the vear and the wide selection of prey taken by most anurans and lizards, it seems as though food is not a limited resource for these animals. Anurans and lizards have minimal re- quirements for shelter in tropical rain- forests. Diurnal species either retreat to subterranean cavities or perch on vege- tation at night. Noctiual species, all of which apparently are more suspectible to desiccation than most diurnal species, find abundant shelter amidst debris on the ground, beneath bark of trees, in crevices in logs and trees, on the under- sides of leaves of trees, and in brome- liads. The super-abundance of such shel- ters and the wide dispersal of animals in the available shelters seem to preclude shelter as being a limited resource. A third important resource is oviposi- tion sites. For the lizards and those anu- rans that have terrestrial eggs, most of which are laid amidst debris on the ground, oviposition sites are nearly om- nipresc>nt. Although obviously more re- stricted, calling and oviposition sites for those anurans that deposit their eggs in water or in vegetation above water also are present in greater abundance than utilized by the community. Therefore, EQUATORIAL HERPETOFAUNA 329 oviposition sites do not seem to he a lim- ited resource. Most of the statements in the preced- ing three paragraphs probahly also apply to the snakes, which are higher in the trophic structure, feeding primarily on small lizards and anurans. Occasionally food may be a limited resource for the few species that feed on small mammals or on other snakes. What then are limited resources, if any, in this herpetological community? The primary forest can be viewed as an open system in that there are no (or only few) limitations of the resources needed for the populations of amphib- ians and reptiles living there. However, there may be distinct limitations of the resources available in the ponds and swamps where tadpoles develop (see p. 315). Nonetheless, the competition pres- sure, if indeed it does exist, does not explain the persistence of the richest anuran fauna known for any locality in the world. It can be argued that the large num- bers of species of amphibians and rep- tiles at Santa Cecilia can coexist due to the absence of, or very little pressure from, interspecific competition. The ab- sence of interspecific competition is due to 1) the abundance of available re- sources, 2) structural heterogeneity of the environment, 3) climatic equability of the environment, and 4) differential utilization of resources spatially and tem- porally by the herpetofauna. One might question the species pack- ing of the herpetofauna. Are all of the available niches filled? Possibly not. Ob- vious absentees are herbivorous arbo- real lizards, such as Iguana, and brome- liad-breeding and stream-breeding hylid frogs. There are no nocturnal snakes feeding on centipedes, nor are there noc- turnal snakes feeding on tadpoles in ponds. Certainly there must be many other less-obvious absentees. It can be inferred from the discus- sion above that competition is not in- tense in the herpetofauna and therefore populations are not at or near the carry- ing capacity of the environment. If com- petition does not play a significant role in the exclusion or dimunition of num- bers of individuals, what factors are operant on population control? Two fac- tors, one abiotic and the other biotic, apparently are significant in regulating populations. The primary abiotic factor is aperi- odic environmental fluctuations, the most striking of which are periods of drought or periods of excessive rainfall. The former has an adverse effect on ephemeral aquatic breeding sites uti- lized by many anurans. Some small ponds and marshes frequently dry up resulting in the deaths of many tadpoles and eggs. Those species most severely effected by short droughts are the repro- ductive r-strategists (see discussion of fecundity in section on Reproductive Biology). Excessive rainfall results in flooding of areas that are usually dry. This results in some animals being driven from their normal shelters and the drowning of terrestrial eggs. The primary biotic factor regulating population sizes in anurans and lizards is predation. In a lucid discussion of competition and predation in natural communities Connell (1975:475) stated: "Predation should be regarded as being of primary importance, either directly determining the species composition or in preventing competitive exclusion, ex- cept where the effect of predation is re- duced for some reason." Furthermore, Connell (1975:484) concluded: "Since predation seems to be more intense in more benign physical conditions, com- petition should be prevented regularly in more benign regimes." The evidence from the herpetofauna at Santa Cecilia strongly supports Con- nell's contentions. In tropical rainforest communities most anurans and lizards are abundant prey for diverse predators. Anuran eggs and tadpoles in ponds are preyed upon by birds, fish, and aquatic insects. Terrestrial anuran eggs and rep- 330 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY tile eggs are eaten by insects, snakes, and mammals; also the eggs are subject to attacks by mold. Snakes are important predators on frogs and lizards, these be- ing preyed upon both night and day. Small mammals, especially opossums (Marmosa) and night monkeys (Ae- tous), seem to relish small frogs, mainly Eleutherodactylus, and sleeping Anolis. A variety of birds are known to prey on frogs and lizards. Reproductive strategies of the com- ponent species are highly diverse (see section on Reproductive Biology). The population growth of the anurans and lizards is dependent primarily upon three density independent factors: 1) fecun- dity, 2) environmental fluctuations, and 3) predation. The latter is, or course, generally more significant in a prey spe- cies that has dense populations rather than one that is at a low density. Only if a species or group of species having considerable overlap in their utilization of available resources exceed the carry- ing capacity of these resources will com- petition become effective in limiting population growth or excluding species in the community. Theoretically, inter- specific competition might occur most frequently among r-strategists, but one must remember that the r-strategists live in the least stable environments and also are subject to heavy predation pres- sure. Therefore, only after lengthy pe- riods of favorable environmental condi- tions will their abundance negate the pressure from predation and will they reach the carrying capacity of the en- vironment, which also will be reached more precipitously by a sudden lowering of the canying capacity due to environ- mental fluctuations. Thus, in answering the two original questions: Herpetological communities in aseasonal tropical forests are not struc- tured in the same way as bird communi- ties. The models generated on evidence from bird communities do not generally apply to herpetological communities. Seasonal Versus Aseasonal Environments Although it is common to compare tropical and temperate environments, for the reasons given on p. 289 I prefer to make the more meaningful comparison between the aseasonal tropical rainforest and seasonal environments, both tropical and temperate. Abiotically, the asea- sonal tropical rainforest can be charac- terized by its equability in temperature, rainfall, and light. These equable con- ditions, plus the diversity of structural habitat and abundance of food provide a wide array of potential niches for a vast number of species, which are active throughout the year. By contrast, in seasonal environments, amphibians and many, if not all, reptiles are active only during part of the year. In temperate regions low temperatiu'es necessitate hi- bernation during part of the year, where- as in the dry tropics aestivation is normal for amphibians and not uncommon for reptiles. In the seasonal environments there is less diversity of structural habi- tat and less diversity of and seasonality of food; thus, the number of potential niches is less numerous than in the trop- ical rainforest. In his discussion of bird species rich- ness in the tropics, Mac-Arthur (1972: 216) stated: ". . . either the spectrum of resources, K, is greater in the tropics; the utilization per species, U, is less in the tropics; the overlap between species, O/H, is greater in the tropics; or the dimensionality of the environment, C, is greater in the tropics. Any of these can cause coexistence of the increased num- ber of species that has been observed in the tropics. The embarrassment is that all are likely to be true, at k-ast in some places!" In comparison with herpetofaunal communities in seasonal environments, communities in aseasonal tropical forests certainly enjoy a greater spectrum of resources and greater dimensionality. I believe that these two factors alone can explain most of the differences that exist EQUATORIAL HERPETOFAUNA 331 in relative species richness. At least among the anurans and lizards, the vast majority of which are habitat generalists and feeding generalists, resource utiliza- tion does not seem to be any less in the tropics; in fact, in many cases utilization may be broader. If, as has been sug- gested previously, the abundance of prey for the secondary consumers in the rain- forest precludes competition for food, then niche overlap might be significantly higher in the tropics without being detri- mental to any of the predators. Unfortunately, few studies on herpe- to faunal communities have been under- taken. Pianka ( 1973, 1975, and references cited therein) compared lizard commu- nities in desert regions in Africa, Aus- tralia, and South America. Fucntes ( 1976 ) compared Chilean and Califor- nian lizard communities. Crump (1971) analyzed ecological distribution of liz- ards and amphibians at Belem, Brasil. Rand and Humphrey (1968) inferred interspecific competition in habitat selec- tion among lizards at Belem, Brasil. Al- though the results of these studies, and others, show that species diversity and richness vary between communities and that some ecological counterparts are absent in certain communities, they have not provided conclusive evidence for the substantiation of predicted difi^erences between communities in aseasonal tropi- cal environments and those in seasonal environments. On the basis of the much larger num- bers of species in aseasonal tropical com- munities than in seasonal communities, it is reasonable to postulate that poten- tially, interspecific competition would be greater in the aseasonal tropical commu- nities. However, because of the struc- trn-al heterogeneity in the aseasonal trop- ical environment and the abundance and deversity of food, in comparison with that in the seasonal environments, actual levels of competition are minimized far below the potential levels. Relative equability of temperature and moisture are the most important factors regulating the richness of herpetological communi- ties. (Of course, the historical factor of the richness of the source material must be taken into consideration.) BIOGEOGRAPHY Now that the herpetological commu- nity at Santa Cecilia has been defined and the various aspects of the commu- nity interpreted, it is desirable to view the community in an historical context. In order to interpret certain historical aspects of the herpetofauna it is neces- saiy to determine the distribution of the components of the fauna. Then we can attempt to answer the following ques- tions: From where did the components of the herpetofauna come? How did they arrive at Santa Cecilia? How did the present community evolve? DISTRIBUTION OF THE HERPETOFAUNA The plotting of distributions of the 185 species in the herpetofauna at Santa Cecilia resulted in the definition of eight general patterns of distribution ( Fig. 197). The immediately obvious dichot- omy is that four patterns involve the en- tire Amazon Basin (Fig. 197-A-D). Twenty-eight amphibians and 56 reptiles (=84 species or 45% of the fauna) are widespread in the basin. The other di- chotomy involves species that are essen- tially restricted to the upper part of the basin ( Fig. 197E-H ) ; 66 amphibians and 35 reptiles (=101 species or 55% of the fauna) are so restricted. The most widespread pattern (Fig. 197A) includes the humid lowland trop- ics of the Amazon Basin, the Guianas, lower Central America, and the forested Pacific lowlands of Colombia and Ecua- dor. Some .species having this pattern also occur on the Caribbean lowlands of 332 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fig. 197. — Distribution patterns of species in the herpetofauna at Santa Cecilia. Numbers arc aniiDhibians/reptiles; the dot is Santa Cecilia. EQUATORIAL HERPETOFAUNA 333 Colombia and Venezuela and/or the low- lands of southeastern Brasil. Examples of these wide-ranging species are Bufo 77iarinus, Thecadactijhis rapicauda, and Boa constrictor. Most of the species in this group seem to have wide habitat tolerances and are not confined to wet forests. The second pattern (Fig. 197B) is like the fonner except for the exclusion of the Pacific lowlands of Colombia and Ecuador. Examples include Phrijnolnjas venuJosa, Kinosternon scorpioides, Amei- va omeiva, and Leptodeira anmdato. The most common pattern among species that range through the entire Amazon Basin also includes the Guianas but does not include northwestern South America or lower Central America ( Fig. 197C). Examples of this wide-spread component include Fipa pipa, Hi/hi p,eo- graphica, Anolis chrysolepis, Dracaena guianensis, and Micrurus lemniscatus. A much smaller number of species occur throughout the basin but do not extend into the Guianas ( Fig. 197D ) . Examples of these include Leptodactylus rhodo- mijstax, Drymoluher dichrous, and Mi- crurus spixii. Among those distributional patterns that include only the upper part of the Amazon Basin, the largest group of spe- cies (57=31%) inhabit most of the upper part of the basin ( Fig. 197E ) . Examples of this pattern include Dendrophryniscus minutus, Phyllomedusa farsius, Paleosu- chus trigonatus, Anolis trachyderma, Atractus chips, and Bothrops casteJnaudi. A small group of amphibians, including Oscaecdia hassJeri, BoJitogJossa peruvi- ana, Eleutherodactylus ockendeni, Ilyla hifurca, Hyla rhodopepJa, Chiasnwclcis hassJeri, and Hamptophryne holiviana, occurs only along the western edge of the basin from Colombia to southern Peru or Bolivia (Fig. 197G). Seventeen species have a similar pattern except that they also occur in the Guianas ( Fig. 197F). Examples of this group are Lith- odytes Jineatus, Hyla fasciata, Phryno- hyas coriacea, Dipsas indica and Ninia hudsoni. The most restricted pattern is that encompassing only the upper edge of the basin in Ecuador and adjacent Colombia (Fig. 197H). The IS amphibians and five reptiles in this group are: Micro- caecilia aJhiceps, Bolitoglossa ecuador- ensis, Eleutherodactylus diadematus, E. rnartiae, E. nigrovittatus, E. orphnohii- mus, E. paiduhis, E. pseudoacuminatus, E. quaquaversus, Bufo glaherrinius, Col- ostethus sauli, Uemiphractus prohosci- deus, Hyla hokermanni, Nyctimantis ru- giceps, Centrolenella midas, C. resplen- dens, Chiasmocleis anatipes, Syncope antenori, Alopoglossus copii, Atractus occipitoalbus, Helicops petersi, Leima- dophis sp., and Oxyrhopus formosus. The proximity of Santa Cecilia to the base of the Andes ( ± 40 km ) necessi- tates an examination of the altitudinal distribution of the species in the Santa Cecilia herpetofauna on the eastern face of the Andes in Ecuador. Many of the species (37%) occurring at Santa Cecilia do not ascend the slopes at all; their distributions terminate by 400 m eleva- tion (Fig. 198). At elevations above 600 m, 56 percent of the species are no longer present, and only 28 percent ex- tend above 1000 m. Six frogs (Eleuthero- dactylus acuminatus, E conspicillatus, E. quaquaversus, Osteocephalus huck- leyi, O. taurinus, and Bufo fyphonius) and one reptile {Lcinmdophis reginae) occur above 1500 m. A zone of luxuriant cloud forest occurs at elevations between 1100 and 1800 m on the eastern face of the Andes in Ecuador. This cool, moist forest apparently presents environmental conditions be>ond the tolerances of most lowland reptiles and many amphibians. Pond-breeding frogs are limited further by the near absence of ponds on the An- dean slopes. Although the distributions of species of amphibians and reptiles in the Ama- zon Basin are not known in great detail, sufficientlv thorough sampling has been 334 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 10 25 AMPHIBIANS T ■ CO UJ o LJ Q- CO '^ METERS: a: UJ CD z> 10 - 15 500 I,,,,., 1000 1500 rT-rl77 ■ ^ 12 2000 i 42 REPTILES Fig. 198. — Upper elevational limits on the east- em slopes of the Andes of species in the herpe- tofauna at Santa Cecilia. done at Iquitos, Peru (Dixon and Soini, 1975, 1976) and Belem, Brasil (Crump, 1971) to allow meaningful comparisons of these with Santa Cecilia and with the well-known fauna on Barro Colorado Is- land, Panama (Myers and Rand, 1969). Faunal comparisons were made by the method developed by Duellman (1965, 1966) of determining a faunal resem- blance factor by means of the formula FRF=2C/Ni+No, where Ni and No are the numbers of species in any two given faunas, and C is the number of species common to both faunas. As would be expected the greatest amount of faunal resemblance is between Santa Cecilia and Iquitos and the least between Santa Cecilia and Barro Colorado Island (Ta- ble 33). There are numerous species of rep- Table 33. — Comparison of Heipetofaunas at Four Tropical Localities. ( Boldface numbers are actual numbers of spe- cies at a locality; Roman numbers are the number of species in common between locali- ties; italicized numbers are tlie faunal resem- blance factors ( see text for explanation ) . In each group, the first row is amphibians, second row is reptiles, and third row is the total of amphibians and reptiles.) -a U 1 en p-( O 2 pa 1 m Barro Colorado Island, Panau Santa Cecilia, Ecuador . 94 53 22 4 91 83 42 13 185 136 64 17 Iquitos, Peru . .654 68 23 3 .715 141 60 19 .690 209 83 22 Belem, Brasil . .379 426 40 2 .494 .545 79 15 .421 .506 119 17 Barro Colorado Island, Panama . ^ .032 .030 .028 32 .163 .182 .204 68 .119 .142 .155 100 tiles that are widespread in the Amazon Basin that do not occur at Santa Cecilia. Many of these are aquatic species ( Heli- cops leopardimts, H. poIyJepis, Hydrops martii, H. triangularis, Pseudoenjx plica- tilis, Phrynops nasutus, Podocnemis ex- pansa, P. sextuberculata, P. iinifilis, Me- lanosuchtis niger, and Poleosuchus pal- pehrosus) that characteristically inhabit quiet rivers or swamps associated with flood plains. The gradients of the streams at Santa Cecilia and the near absence of swamps associated with rivers apparent- ly preclude the existence of these aquatic species at Santa Cecilia. Some other spe- cies (Iguana iguana. Plica plica, Ken- tropyx calcaratus, Amphishaena alba, Oxyhelis aeneus, and O. fidgidus) are usually associated with more open forest than that occurring naturally at Santa Cecilia. Certain species at Santa Cecilia EQUATORIAL HERPETOFAUNA 335 (e.g. Ameiva ameiva, Mahmja mohoinja, Hyla rubra and Bufo morimis) also be- long in this category. These arc "weed" species that disperse rapidly along roads and other man-made clearings. Perhaps the four species mentioned for Santa Ce- cilia are more vagile than the others, which in time might be expected to reach Santa Cecilia. ORIGIN OF THE HERPETOFAUNA The many recent studies on plate tec- tonics provide an overwhelming amount of evidence that South America was an isolated land mass throughout most of the Tertiary. The breakup of Western Gondwanaland commenced in the early Cretaceous (±125 million years BP) (Larson and Ladd, 1973), and the sepa- ration of Africa from South America occurred in the late Cretaceous ( ± 90 million years BP) (Grant, 1971). Raven and Axelrod (1972) summarized evidence for direct biotic migration between South America and Australia via Antarc- tica in the Eocene and probably well into the Oligocene ( ± 38 million years BP). After the initial breakup of Pan- gaea in the Triassic there was no direct land connection between North America and South America until the late Ter- tiary; their positions converged begin- ning in the mid-Cretaceous (Dietz and Holden, 1970). Holden and Dietz ( 1972) and Mafait and Dinkelman (1972) pro- vided evidence for an island arc between nuclear Central America and South America in the Cretaceous and early Tertiary; this arc moved northeastward (in relation to South America) through most of the Tertiary and fonned the present Lesser Antilles. The region of lower Central America formed as a vol- canic archipelago in the Oligocene with additional land emerging and coalescing with nuclear Central America about 10- 12 million years BP and finally with South America about 5.7 million years BP (Dengo, 1969; Emiliani, Gaertner, and Lidz, 1972; Haffer, 1970; Whitmore and Stewart, 1965). Further biological evidence for faunal interchange between Central America and South America was presented by Rosen (1975), Savage (1974), and Tedford (1974). Raven and Axelrod (1974:549) sum- marized the isolation of South America: "Judging from the geology of the region, and the relative motions of the plates South America was more accessible to immigration from Africa than from North America until after the Early Eocene. Subsequently, more and more insular connections with North America were established, culminating with a direct land connection only 5.7 my BP. Clearly, the history of South American biota has been one of evolution in isolation of an initial West Gondwanaland stock shared with Africa. To South America have come many cool temperate Australasian plants and animals, essentially overland until the Eo-Oligocene, and by overseas long-distance dispersal subsequently. South America contributed increasingly to the flora of tropical and subtropical North America during the Tertiary, it received immigrants from temperate North America only as the Cordillera rose in the late Neogene, at which time it also contributed montane tropical taxa to Central America." Summaries of the paleogeography of South American amphibians and reptiles were presented by Reig ( 1968 ) and Cra- craft (1974). The history of specific groups has been treated by various au- thors - caecilians (Estes and Wake, 1972), salamanders (Wake, 1966), anu- rans (Cei, 1968; Estes, 1975; Estes and Reig, 1973; Savage, 1973), crocodilians (Sill, 1968), turtles (Simpson, 1950), lizards (Estes and Price, 1973); Vanzo- lini, 1968). Laurent (1973) briefly com- pared the herpetofauna of Africa and South America. Of the 25 family groups of amphib- ians and reptiles comprising the fauna at Santa Cecilia, 17 are of Gondwanian ori- gin and eight are Laurasian in origin (Table 34). The primary sources of the 336 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Table 34. — Paleogeography of family groups in Santa Cecilia Herpetofauna. Center of Family Group Origin Caeciliidae Gondwanaland Plethodontidae N. Anier. Pipidae Afr.-S. Amer. Leptodactylidae S. Amer. Bufonidae .A.fr.-S. Amer. Dendrobatidae S. Amer. Hylidae S. Amer. Centrolenidae S. Amer. Microhylidae Gondwanaland Ranidae Africa Cheliidae Aust.-S. Amer. Kinostemidae N. Amer. Testudinidae Laurasia Crocodylidae - Afr.-S. Amer. Gekkonidae Gondwanaland Iguanidae S. Amer. Scincidae Laurasia Teiidae S. Amer. Amphisbaenidae Laurasia Aniliidae -— ? Laurasia Boidae Gondwanaland Colubrinae Laurasia Xenodontinae S. Amer. Elapidae . Gondwanaland Crotalidae Laurasia Source of S. Amer. Group Appearance IN S. Amer. S. Amer. C. Amer. S. Amer. S. Amer. S. Amer. S. Amer. S. Ajner. S. Amer. S. Amer. N. Amer. S. Amer. N. Amer. N. Amer. S. Amer. S. Amer. S. Ajiier. N. Amer. S. Amer. N. Amer. N. Amer. S. Amer. N. Amer. S. Amer. S. Amer. N. Amer. Paleocene Post-Miocene Paleocene Eocene Paleocene ? ? Eocene ? ? Post-Miocene Cretaceous Post-Miocene ? Miocene Triassic Paleocene Cretaceous Post-Miocene Cretaceous ? Miocene ? Cretaceous Post-Miocene Cretaceous Cretaceous familial component.s are South America (16), Central America (1); and North America (8). Thu.s, the members of most of the familial groups evolved in South America. Some groups, such as the Testudinidae, Amphisbaenidae, Ani- liidae, and Crotalidae apparently entered South America from North America prior to the closure of the Panamanian Portal in the early Pliocene. Other North and Central American groups probably did not enter South America until the Plio- cene; these include bolitoglossine pletho- dontids, Rana, Kinosternon, scincids, and colubrine colubrids. The xenodontine colubrids underwent their adaptive radi- ation in South Amcnica, but some stocks evidently migrated northward into nu- clear Central America prior to continu- ous land connections in the early Plio- cene. As xenodontine isolates, genera such as LeptocJeira (Dnellman, 1958) and RlunUnaea (Myers, 1974) evolved and speciated in Mexico and Central America. Later, probably in the Plio- cene, some species in these genera dis- persed into South America. It is safe to assume that by the end of the Pliocene ( ± 1.8 million years BP) all of the component families and most, if not all, of the genera of amphibians and reptiles existed in South America in or peripheral to the Amazon Basin. TERTIARY AND QUATERNARY HISTORY OF THE AMAZON BIOTA The interpretation of the liistorical biogeography of tropical regions has been obfuscated by the dogmatic beliefs that the tropical rainforests of the world are ancient environments that have changed very little in millions of years, and that they were unaffected by cli- matic fluctuations in the Pleistocene (see Schwabe, 1969, as a recent example). Moreau (1963, 1969) summarized evi- dence for climatic changes in the Pleisto- cene in tropical Africa and used this EQUATORIAL HERPETOFAUNA 337 evidence for explaining certain patterns of distribution of the biota. Haffer (1969), Vanzolini and Williams (1970), and Vuilleumier ( 1971 ) introduced the same concepts in South American bio- geographic analyses. The geological palynological, and climatic evidence for fluctuating environments in the equa- torial tropics in South America were doc- umented thoroughly by Miiller (1973) and Haffer (1974), and the palynologi- cal evidence was further summarized by Van der Hammen (1975). Haffer (1969, 1974, 1975) provided evidence from avian distributions and patterns of differentation for six per- sistent forest refugia in the Amazon Basin. Vanzolini and Williams (1910) proposed four core areas for the differ- entiation of Anolis chrysolepis: they pro- posed an evolutionary model of three expansions and two contractions of for- est environments. Mi.iller (1973) recog- nized four major dispersal centers in the Amazon Basin; the basis for his centers are coincidental distributional limits of vertebrates. On the basis of plant distri- butions, Prance ( 1973 ) provided evi- dence for forest refugia that are remark- ably consistent with those proposed by Haffer but more extensive than Vanzo- lini and W^illiams' core areas. Studies on heliconian butterflies by Brown ( 1972 ) and Brown and Mielki (1972), on hylid frogs by Duellman (1972b) and Duell- man and Crump ( 1974 ) , and on lepto- dactylid frogs by Heyer (1973) provide additional support for the hypothesis of distributional discontinuities and pat- terns of differentiation being correlated with the existence of forest refugia dur- ing arid climatic phases in the Quater- nary. Although the details of the Tertiary geologic history of the Amazon Basin are not known, Haffer (1974:128) stated: "Probably the sediments were deposited in large swamps and marshes, and on vast flood plains crossed by nu- merous meandering streams flowing into a huge inland sea, which connected to the sub-Andean basin to the west and with the Atlantic Ocean to the east. . . . During the Late Pliocene (end of Ter- tiary time) the Amazon valley probably was covered by a large lake with a nar- row eastern outlet. . . . The final uplift of the Andes and of the upper Amazon- ian lowlands (previously covered by a huge inland sea) led to the establish- ment of the eastwardly directed drainage pattern in Amazonia during early Qua- ternary time. The rivers shaped their present beds cutting into the Tertiary formations during the Pleistocene peri- ods of lowered world sea level and de- positing sediments in the broad Amazon valley during periods of raised sea level (interglacials), when a large portion of the Amazon valley was flooded." The uplift of the central and northern Andes began in the Miocene, but geo- logical (Petersen, 195S) and palynologi- cal (Van der Hammen, 1966; Van der Hammen and Gonzalez, 1964) evidence indicates that the Andes underwent their greatest uplift at the end of the Pliocene and at the beginning of the Pleistocene (± 1.8 million years BP). Haffer (1974:135) summarized the existing evidence on Pleistocene climates in the American tropics and concluded: "Two major events detemrined environ- mental conditions in tropical South America during this time interval. First, the main uplift of the Andes during the Late Pliocene and Early Pleistocene led to the vast expansion of dense forest onto the fully emerging upper Amazon basin. This forest also reached far to the north to the slopes and lowlands around the mountains along the Colombian Andes and south on the eastern slope to Bolivia. Second, the alterations of humid and dry climatic periods in the tropics, caused by the Pleistocene periodicity of world cli- mate, resulted in periodic shrinkage and expansion of the lowland forests and non- forest vegetation, respectively. At the same time, changing world sea levels led to the exposure of the continental shelves in glacial times, which increased the 338 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY width of tlie coastal lowlands, and to the flooding of part of the present coastal plains and of the Amazon Valley during interglacial periods." Post-Pleistocene changes in the Ama- zon Basin have consisted on alternating humid and arid climatic phases. Geo- morphological, pah'nological, and paleo- botanical evidence places two arid phases at about 11,000 and 2500-4000 years ago (Danmuth and Fairbridge, i970; Haff^er, 1974; Vanzolini and Ab'- Saber, 1968). During these arid phases the vast Amazonian rainforest became fragmented, and the nonforest environ- ments (savanna and cerrados) expand- ed. At the time of the last arid phase (2500-4000 years BP) the Amazon forest was probably completly separated into lower and upper Amazonian forest by a continuous dry belt extending from southern Venezuela to northeastern Bra- sil; this area today receives less than 2500 mm of rainfall annually. During the most severe arid phases tropical rainfor- est persisted in certain areas due to fa- vorable local edaphic, topographic, and climatic conditions (Haffer, 1969, 1974). These post-Pleistocene climatic changes and concomitant restriction and expansion of the Amazonian rainforest are the environmental bases for patterns of distribution and speciation seen in northern South America today (Haffer, 1969, 1974; Miiller, 1973; Vanzolini, 1973; Vanzolini and Williams, 1970). The upper part of the Amazon Basin apparently emerged from the mid-conti- nental lake and became covered with rainforest at about the beginning of the Pleistocene (± l.S million years BP). This is approximately 3.9 million years after the establishment of the continuous land connection between Central Amer- ica and South America. During this time the northern Andes were low rolling hills covered with tropical or subtropical for- est (Haffer, 1975); thus, dispersal routes were available for Central American groups to enter the Amazon Basin. EVOLUTION OF THE HERPETOFAUNAL COMMUNITY Santa Cecilia is located in one of the largest of the hypothesized forest re- fugia, the Napo Refugium; the Ucayali Refugium lies to the south in Peru ( Haf- fer, 1969). Coalescence of the forests in these refugia must have been more nearly continuous throughout the Qua- ternary than the meeting of either of these forests with others eastward in the Amazon Basin. Lying close to the base of the Andes, Santa Cecilia also must have received fauna from the Andean slopes when climates were depressed at times of Andean glaciation. Thus, throughout the Quaternary the Santa Ce- cilia area was a humid tropical forest and at various times was in contact with other forested areas to the south and east and at other times was isolated from these forests. Furthermore, the Santa Cecilia area was in continuous contact with the forested Andean slopes and in more or less continuous contact with peripheral Amazon Basin forests extend- ing south into Peru and perhaps Bolivia and north into Colombia and at times eastward from there to the Guiana Shield. These various periods of contact and isolation resulted in restricted dis- tribution patterns ( Fig. 197-E-H ) . The fluctuating environmental condi- tions in the Amazon Basin provided a scenario for the processes of speciation. Populations of species were isolated in ecological islands and differentiated; coalescence" of formerly isolated environ- ments resulted in the reunification of populations in these environments. If the isolated populations had not differ- entiated to the point of reproductive iso- lation, introgression would occur when and where the populations were re- united. If the isolated populations had differentiated to the point of genetic in- compatil^ility, their reunification could have any one of three results: 1) Two populations not dependant on a limited resource can coexist in geographic sym- EQUATORIAL HERPETOFAUNA 339 patiy. 2) Two populations equally adapted for utilization of a limited re- source will exist in geographic parapa- try. 3) If one of two populations was more effective in utilizing a limited re- source than the other, the former popu- lation will expand through the range of the latter population. Thus, the latter is an example of competitive dominance that theoretically will result in the ex- tinction of one population. The second situation is an example of competitive exclusion and may explain the restricted ranges of many species in the Amazon Basin. The first alternative provides us with an explanation of species richness in one communitv. Because of aseasonal reproductive cycles, complete absence of pair bonds, and frequent mating and reproduction, most populations of tropical amphibians and reptiles probably are highly heterog- enous genetically and therefore are likely to develop genetic differences during very brief periods in isolation. Introgres- sion of genetically compatible isolates may explain some of the bewildering mosaics of variation seen in some species, whereas differentiation to the point of reproductive isolation allows the coex- istence of several closely related species. The latter example was used by Duellman and Crump (1974) to explain the coexistence of three species in the Hijla parviceps group at Santa Cecilia, and the same explanation can be applied to the Hyla leucopliyllata group, five species of which coexist at Santa Cecilia. The species diff^er in mating calls, ovi- position sites, and kinds of ponds used for reproduction. The same explanation can be appled to the three species of Chiosmocleis, the Leptodactyhis penta- doctylus group, the Hyla geographica group, Osteocephahis, the two species of Enyalioides, the two species of Alopo- glossus, and perhaps several others. The herpetofauna at Santa Cecilia is composed mainly of forest-adapted spe- cies that either are restricted to the up- per Amazon Basin or widespread in the Amazon I3asin. Probably the majority of the fonuer group and some of the latter group evolved in the Napo and Ucayali refugia. Other components of the fauna are inhabitants of the Andean slopes that barely invade the Amazon Basin, whereas a few non-forest species (Biifo moritms, Hyla rubra, Ameiva ameiva, and Mabitya mahouya) are re- cent immigrants along the river playas. EPILOGUE During the ten years that I have been studying Amazonian herpetofaunas, I have come to realize several shortcom- ings of the study of the community at Santa Cecilia. How many times must investigators wish upon the completion of a study that they could begin again and do it right? On the basis of experi- ence at Santa Cecilia, I offer the follow- ing suggestions for the study of herpeto- logical communities in tropical rainfor- ests: 1. Data needed for thorough commu- nity analyses are derived in part from observations of the animals and in part from examination of preserved speci- mens. These two kinds of data cannot be collected at the same time and place without the collection of specimens in- terfering with the natural populations. Because both kinds of data are needed from throughout the year, a large study area could be divided into two parts - one for observations and one for collect- ing. A long term study in a smaller area could be divided temporally with inten- sive collecting following a period of ob- servation. 2. In order to obtain meaningful data on reproductive cycles it is necessary to have large series of specimens from throughout the year. It is doubtful if adequate samples of some lizards and most snakes could be obtained in fewer 340 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY than five years. Data on reproductive cycles of individuals need to be obtained by mark and recapture studies. In spe- cies-rich tropical communities it is un- feasible to attempt mark and recapture studies on all species. Instead selected species differing in reproductive modes and habits should be investigated. 3. Analyses of food should be based not only on taxonomic content but also on volume; analyses also should take into account size of predator and prey and time of year. Thus, series of speci- mens of all size classes are needed from throughout the year. 4. Concomitant with mark and recap- ture studies designed to obtain data on reproductive cycles it is possible to ob- tain data on growth, longevity, and pop- ulation turnover. Such data are practi- cally non-existent for tropical species. 5. The most accurate way I know of obtaining data on the numbers of indi- viduals and species is to clear the forest. A study area one kilometer square should be bordered by a 50 m wide cleared area. Systematic paring of the edge of the study by bulldozers and felling trees into the cleared area provides a method of sampling the fauna from the shallow subterranean level to the canopy. A thorough sampling of a systematically cleared forest will provide a much more accurate assessment of actual numbers of individuals of species than now avail- able. Care must be exercised in applying ecological and population models. At the present time our understanding of a min- imal data base on tropical communities is so limited that only the most general conclusions can be inferred. Models based on data from one group of organ- isms or from one environmental regime do not necessarily apply to other groups of organisms or to other environmental regimes. Before we have an adequate understanding of tropical communities much greater amounts of research funds must be designated for field studies. There is a great sense of urgency in the collection and study of the Ama- zonian biota. Within the past few years man in his frenetic pace of exploitation has devastated millions of hectares of the Amazonian forest, once the most vast area of tropical wilderness on earth. Data on the taxonomy and distribution of species can be accumulated by ex- tensive collecting throughout the basin. However, the data so accumulated in such salvage operations will provide only limited biological information on the fauna. The studies of communities nec- essary to acquire an understanding of tropical faunas and their ecology reciuire the preservation of natural regions. In- telligently selected areas of sufficient size (see Terborgh, 1975) will result in the conservation of biotic communities and provide areas for the study of commu- nities. The Republic of Ecuador, which has set a conservation precedent in the Galapagos Islands, currently is evaluat- ing the Proyecto Cayambe-Coca pro- posed by the FAO of the United Nations. The Proyecto Cayambe-Coca calls for the establishment of a national park and biological preserve extending from Vol- can Cayambe in the Andes to the Rio Coca in the Amazon Basin. This area includes parts of the cloud forest on the eastern face of the Andes and rainforest in the upper reaches of the Amazon Ba- sin. If this projected preserve becomes a reality, a significant part of the fauna associated with the Napo Refugium will be conserved for future study. The community at Santa Cecilia re- ported upon here is gone. The expanse of primary forest along the banks of the Rio Aguarico has been replaced by mil- pas, yuca, and pkitanos, most of which are abandoned in a few years to be taken over by dense bushes and vines or badly eroded. Man through his various cul- tural levels of utilization, exploitation and eventual destruction has destroyed this biological community in a few short EQUATORIAL HERPETOFAUNA 341 years. We can only hope that enhght- ened government agencies will set aside substantial natural preserves in an effort to prevent the "Green Hell" from becom- ing the "Red Desert" (Goodland and Irwin, 1975). RESUMEN Este estudio esta basado en observa- ciones y especimenes obtenidos duran- tes 48 hombres-meses de trabajo de cam- po en la alta cuenca del Amazonas en Ecuador. El sitio principal de estudio fue un area de cerca de 3 kilometros cuadrados en Santa Cecilia, en el Rio Aguarico, a 340 m de altitud, con 00^03' N, 76°59' W como coordenadas. Cuatro otras localidades del Rio Aguarico (Du- reno, Lago Agrio, Puerto Libre, y Puerto Ore) fueron estudiadas con menor in- tensidad. El material reunido en las cinco localidades consiste de 9035 es- pecimenes de 185 especies, mientras que de solo Santa Cecilia 7765 especimenes de 173 especies fueron colectados. Taxo- nomicamente, esta fauna esta compuesta por 5 especies de Gymnophiona (3 pre- sentes en Santa Cecilia), 2 dc Caudata (2), 87 de Anura (81), 6 de Testudines (6), 2 de Crocodilia (2), 29 de Sauria ( 27 ) , 1 de Amphisbaenia ( 1 ) , y 53 de Serpentes (51). La region alrededor de Santa Cecilia tiene un bajo relieve ( < 20 m ) con varios arroyuelos y depresiones. El Rio Aguarico es un rio de aguas blancas, y el Rio Conejo un arroyo sinuoso de aguas lentas. Existen alii dos lagos per- manentes y hay numerosos pantanos en las depresiones. El clima es tipico Af (del sistema de Koppen) con fluctua- ciones aperiodicas pero sin estaciones anuales distintas. Las fluctuaciones dia- rias de la temperatura son de cerca de 10° C, y la diferencia en promedios men- suales de cerca de 2° C. Las tempera- turas maxima y minima registradas han sido 37° y 17° C. La precipitacion anual para el periodo Julio 1971-Junio 1972 fue de 4289 mm, siendo Marzo el mes mas lluvioso (con 619 mm) y Agosto el mas seco ( con 220 mm ) . El mas extenso periodo sin lluvia duro cinco dias, mien- tras que la mas abundante lluvia en 24 horas fue de contuvo 390 mm. La vegetacion natural de Santa Ce- cilia es selva humeda tropical de tipo primaria, la cual esta mejor desarrollada en suelos pianos bien drenados. Los mas altos arboles forman una techumbre a 30-35 m de altura. Un segundo estrato se forma a 15-20 m de altura, y un tercer estrato a 8-12 m. El suelo esta cubierto de hierbas y helechos ( 10-25 cm de al- tura), donde una capa continua de ho- jarasca esta presente. Selvas secundarias, resultantes de in- tervenciones humanas o de caidas natu- rales de arboles, estan caracterizadas por una sucesion forestal sin estratificacion, dominada frecuentemente por Cecropia de alturas de 10 a 12 metros. En estas selvas hay densos matorrales y poca hojarasca; grupos de bambues de 15 a 18 m de altrua se encuentran repartidos a lo largo de este tipo de selva. Depresiones del terreno en ambas clases de selvas contienen agua al menos intermitentemente. Estos pantanos di- fieren de las pequeiias y poco profundas depresiones que fuertes lluvias se con- vierten en grandes pantanos semiperma- nentes solo despues de la fuertes lluvias. Los pantanos en la selva primaria suelen tener varias palmeras espinosas; aquellos en selvas secundarias suelen estar den- samente poblados con Heliconia. Cie- negas en claros hechos por el hombre tienen pastos, varias araceas, y pequeiias Heliconia. Para los propositos de analisis ecolo- gicos, las siguientes categorias terrestres fueron reconocidas: selva primaria, selva secundaria, borde de selva, bambu, claro, y terrenos cultivados. Los ambientes acuaticos fueron designados como pan- tano, cienega, lago, rio, y arroyo. Uno de los mas dificiles problemas 342 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY encontrados en este estiidio de la herpe- tofauna fue la identificacion de las espe- cies. Como una ayuda a futures traba- |os, claves ilustiadas de la herpetofauna son provistas. En el recuento de las especies, al numero de especimenes de cada locali- dad, lo sigue una descripcion diagnostica de la especie en cuestion. En las sec- ciones sobre piesencia e historia natinal, se incluyen datos sobre el habitat, activi- dad, reproduccion, y alimento. En los recuentos pertinentes hay discusiones sobre los renacuajos, cantos nupciales, variacion, status taxonomico, y nomcn- clatura. Un analisis de los modos reproduc- tivos anuales revela que los anuros son capaces de reproducirse a lo largo del aiio. La lluvia parece ser el mas impor- tante factor con influencia sobre el pe- riodo de reproduccion; la mayoria de las especies responden con un incremento de la actividad reproductiva a los incre- mentos de lluvia. Algunas especies se reproducen solo luego de grandes y es- poradicas lluvias, mientras otras se re- producen solo durante periodos inter- mitentes de pequeiias caidas de agua. Entre los anuros, aquellas especies que depositan los huevos en agua tienen un complemento ovarico mas grande (x:= 1225) que aquellas que depositan sus huevos en nidos de espuma en agua (x:= 457) o en vegetacion por encima del agua (x=342). Especies que tienen huevos terrestres con desarrollo directo tieben huevos mas grandes (x=2.72 mm) que aquellas con huevos terrestres y con renacuajos (x=1.88 mm). El fac- tor tamano del ovario (numero de hue- vos X diametro del huevo/longitud cor- poral de la hembra) es mas grande en especies que depositan huevos en agua (x=24.7) seguido por especies que de- positan huevos en vegetacion por encima del agua (x=8.9) y en nidos de espuma (x=4.2), y menor en acjuellas especies que tienen huevos terrestres con desar- rollo directo (x = 1.2). TTay una correla- cion positiva entre el nnmero de huevos y el tamafio de la hembra, dentro de un determinado modo reproductivo. En al- gunas especies hay una correlacion posi- tiva entre el tamaiio de los huevos y el tamafio de las hembras, dentro de un determinado modo reproductivo. Once modos reproductivos pueden separarse entre los anuros de Santa Ce- cilia: 1. Huevos depositados en amplios cuerpos de agua, con desarrollo de rena- cuajos en el agua. Las agrupaciones de huevos son grandes; huevos y recien nacidos son pequefios (35 especies). 2. Huevos y larvas con desarrollo en huecos de arboles llenos de agua. Las agrupaciones de huevos son moderadas en tamafio; huevos y recien nacidos son pequefios (Nyctimantis rugiceps). 3. Huevos depositados en nidos con forma de tazon. Agrupaciones de huevos grandes; los huevos y recien nacidos son pequefios (IlyJa ])oans). 4. Huevos depositados en la vegeta- cion por encima de aguas lenticas. Agru- paciones de huevos son moderadamente pequenas; huevos son solo ligeramente mayores que aquellos depositados en agua, y los pequefios recien nacidos caen al agua para completar su desarrollo (11 hylidos). 5. Huevos depositados en la vegeta- cion por encima de aguas loticas. Agru- paciones son pequenas; huevos y recien nacidos son moderadmente grandes, y los recien nacidos caen al arroyo (3 Cen- troJcneUa) . 6. Huevos suspendidos en un nido de espuma sobre el agua. Agrupaciones de huevos son moderadamente grandes; huevos y recien nacidos son pequefios ( 9 leptodactylinos ) . 7. Huevos depositados sobre el suelo; renacuajos recien nacidos llevados al agua en la espalda de adultos. Agrupa- ciones de huevos son pequenas; huevos y recien nacidos son relativamente grandes (6 dendrobatidos). 8. Huevos depositados en nidos de espuma sobre tierra. Las posturas son moderadamente pequenas y los huevos EQUATORIAL HERPETOFAUNA 343 relativamente grandes; renacuajos com- pletan su desarrollo dentro del nido de espuma (Adenomera andreae). 9. Huevos depositados en tierra y con desarrollo directo en pequefios sapos; sin renacuajos acuaticos. Posturas pe- quenas pero con grandes huevos ( 17 eleutherodactylinos y Syncope). 10. Huevos depositados en la espalda de la hembra en el agua y tienen desor- rollo directo. Posturas moderadamente pequenas, con huevos grandes (Pipa pipa ) . 11. Huevos llevados sobre la espalda de hembras no acuaticas y teniendo de- sarrollo directo. Posturas pequeiias y huevos grandes {Hemiphracfus prohos- cideiis ) . Las modificaciones del modo general de reproduccion en lagos parecen estar todas en relacion con el incremento de la sobrevivencia de los huevos y de las largas. Esto es logrado a traves de nu- merosas y pequeiias posturas, reniocion de huevos y/o larvas del agua, incre- mento del tanano de los recien nacidos, o a traves de cuidado parental. El incre- mento de la especializacion reproductiva resulta en una reduccion de la fecundi- dad pero en una mayor sobrevivencia. Generalistas, en sentido reproductivo, con alta fecundidad estan mcjor adapta- dos a ambientes no-predecibles. Entre los lagartos, hay 13 especies que paracen tener reproduccion con- tinua, y siete discontinua; se desconoce la situacion para las nueve restantes es- pecies. Generalmente aquellas especies con reproduccion continua son pequeiias (longitud corporal de hembras gravidas 26-100 mm); en cambio las de reproduc- cion discontinua son un poco mas grandes (longitud corporal 48-115 mm). Solo Mahiiya mabouya es vivipara. Aquellas de reproduccion continua tienen multiples posturas por airo, mien- tras que aquellas de reproduccion dis- continua tienen solo una postura por ano. Dos grandes clases de estrategias re- productivas existcn entre los lagartos. Aquella con una madurez mas temprana y multiples posturas se encuentra mayor- itariamente en lagartos de posturas pe- queiias. El tamaiio de la postura puede ser un reflejo del tamaiio del lagarto (microteiidos), eapacidad de soporte de las lamelas adhesivas (geckcSnidos y Ano- lis), o la necesidad de un rapido escape (Enyalioides cofanorum y Vlica timhro). Tamanos grandes y movimientos lentos perniiten posturas mayores en Enyali- oides Jaticeps, PoJychrus marmoratus y Kentropyx peJviceps. La otra estrategia, con una madurez tardia y posturas uni- cas, se da en Ameiva ameiva y Mahuya mahouya. Los patrones de reproduccion no son evidentes en culebras, con los datos re- cogidos al menos. La escasez de hem- bras preriadas sugiere que la reproduc- cioii intraindividual puede ser anual o bianual. Las culebras viviparas (8 es- pecies) tienen nidadas mas grandes (x^ 25.2) que las 45 especies oviparas (x=: 7.6). Las delgadas serpientes arboreales (Dipsas, Imantodes, Leptodeira, Lepto- phis, y OxyheJis) no tienen mas de seis huevos. Quizas estas serpientes se repro- ducen mas frecuentemente que aquellas oviparas de cuerpos pesados o que las viviparas. Bajo el regimen climatico sin esta- ciones de Santa Cecilia la reproduccion en la mayoria de las especies es aciclica. Hay una continua infusion de jovenes en la poblacion y generaciones .superpues- tas. La continuidad de reproduccion tiende a disminuir la fluctuacion pobla- cional y la tasa de extincion. Por cllo, la crianza continua puede ser una de las razones de la alto riqueza en especies en las selvas hiimedas del tropico. En comparacion con especies de am- bientes estacionales, la mayoria de los anfibios y reptiles de las selvas lluviosas del tropico emplean muclias mas estra- tegias reproductivas, se reproducen mas frecuentemente, maduran mas temprano, tienen un mas alto potencial reproduc- tivo por unidad de tiempo, y tienen poblaciones mas estables, pero tienen mas baja sobrevivencia y una corta \'ida. 344 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY El estudio de la estructura de la comunidad consistio primariamente en un amilisis de la distiibucion del ha- bitat, de la actividad diaria, y de dieta y alimentacion. Informacion en relacion a los renacuajos, los cantos nupciales de los aniiros, y relaciones entre los tamanos de las especies tambien fueron iitiliza- dos. La comunidad piiede ser dividida en varios componentes: 1) diurnos y terrestres, 2) diurnos y arboreos, 3) noc- turnos y terrestres, 4) nocturnos y arbo- reos, y 5) acuaticos. Analisis del sobre- montaje de nichos de cada uno de los componentes rcvela una amplia superpo- sicion para varios recursos entre determi- nados tipos adaptativos y determinados niveles troficos. En la detenninacion del niimero de especies que pueden coexistir en Santa Cecilia, lo siguiente fue consid- erado: 1) el espectro y abundancia de recursos disponibles, 2) la estabilidad de los recursos, 3) la extension de la utili- zacion del recurso por los miembres de la comunidad, y 4) la cantidad de sobre- montaje en la utilizacion del recurso por los miembros de la comunidad. El examen de estos criterios revela que en Santa Cecilia hay una alta diver- sidad abundancia de recursos disponi- bles y que el amplio sobremontaje en la de estos recursos indica que la compe- tencia interespecifica no es importante para la regulacion de las especies y sus tamaiios poblacionales. Por el contrario, se postula qut^ las poblaciones (espec- cialmente aquellas de tipo r) estan limi- tadas por la predacion fluctuaciones ape- riodicas del ambiente. La comunidad heipetologica no esta estructurada como las comunidades de aves. Las aves tienen especializaciones para dietas restringidas, diferentes es- trategias para la persecucion o busqueda del alimento, y complejas interacciones sociales intraespecificas e interespecifi- cas. En contraste, la mayoria dc los an- fibios y reptiles (especialmente sapos y lagartos) son generalistas con escaso o nulo comportamiento social. Por cllo, no es sorprendente que muchos modelos ecologicos que han tenido su origen con- ceptual en evidencias ornitologicas pare- cieran no aplicar a la comunidad herpe- tologica tropical. En comparacion con las comunidades herpetologicas de ambientes estacionales, las comunidades de las selvas tropicales sin demarcaciones estacionales gozan de un mayor espectro de recursos y una major dimensionalidad. Estas diferen- cias por si solas pueden explicar la mayor ricjueza en especies que hay en las comu- nidades tropicales. Ocho patrones de distribucion son evidentes entre las especies que compo- nen la herpetofauna de Santa Cecilia. Cuatro de estos patrones incluyen a la totalidad de la cuenca amazonica; 28 anfibios y 56 reptiles (45% del total) caen dentro de estos cuatro patrones. Los otros cuatros patrones se reducen al alto Amazonas; son exhibidos per 66 anfibios y 35 reptiles ( 55% del total ) . Al comparar la herpetofauna de Santa Ce- cilia con aquellas de Iquitos, Peru; de Belem, Brasil; y de la Isla Barro Colo- rado, Panama, se encuentra un alto grado de parecido faunistico entre Santa Ce- cilia e Iquitos y uno menor entre Santa Cecilia y Barro Colorado. Sudamerica se separo de Africa cerca de 90 y de Australia cerca de 38 millones de aiios atras. Durante la mayor parte del Terciario el continentc se movio en un arco hacia Norteamerica, y una coneccion de tierra continua entre Cen- troamerica y Sudamerica se formo aprox- imademente 5.7 millones de anos atras. Ann cuando algiin intercambio faunis- tico ocurrio entre Norte y Sudamerica a traves de un archipielago, Sudamerica estuvo efectivamente aislada de otros continentes en la mayoria del Terciario. Durante estc tiempo, miembros de la mayoria de las familias de los anfibios y reptiles evolucionaron en Sudamerica. De los 25 grupos familiares de Santa Ce- cilia, 16 son de origen gondwanalandico; nueve otros invadieron Sudamerica desde Norteamerica — cuatro previo al Plio- ceno via archipielago y cinco luego del EQUATORIAL HERPETOFAUNA 345 establecimiento de la coneccion de los continentes en el Plioceno. Se asume que al final del Plioceno (1.8 millones de ancs atras) todas las familias y la may- oiia, sino todos, de los generos de an- fibios y reptiles ahora conocidos en Santa Cecilia existian ya en Sudamerica en o alrededor de la cuenca amazonica. La biota de la cuenca amazonica durante el Terciario y el Cuaternario estuvo influenciada por un lago que se retiraba gradualniente en la cuenca y per el levantamijento de los Andes. La selva pluviosa se disperse sobre la tierra emergente pero a causa de la fluctuacion entre periodos secos y humedas del cliina se tuvo contracciones periodicas de la selva lluviosa; las mas recientes fases aridas ocurren hace 11.000 y 2500-4000 anos atras. Durante estas fa.ses aridas la selva pluviosa persistio en refugios aisla- dos; uno de los mas grandes (Refugio Napo) abarcaba a Santa Cecilia. Las especies animales que habitan le selva lluviosa sobrevivieron en los refugios forestales y alii se diferenciaron. Si su diferenciacion resulto en aislamiento re- productive de otras poblaciones aisladas, la subsecuente reunificacion de la selva permitio a las recientemente fomiadas especies vivir cu i)arapatria si ellas com- petian por recursos limitados, o en sim- patria si los recursos necesarios no eran restringidos. Recursos abundantcs pcr- mitieron la acumulacion de la immensa comunidad herpetologica en Santa Ce- cilia. El presente trabajo es solo una intro- duccion al estudio de comunidades her- petologicas tropicales. Mas estudios cuantitativos c intensos son necesarios. Estudios detallados sobre los ciclos re- productivos y sobre la frecuencia de los cruzamientos individuales son requcri- dos urgentemente. Desgraciadamente el hombre esta destruyendo gradualniente la biota amazonica. Se requiere en forma urgente colectar series adecuadas de es- pecimenes para estudios taxonomicos y distribucionales de la biota, pero esta clase de colecta compensatorio no per- mitira estudiar las comunidades. Si se quiere tener un entendimiento de las comunidades de las selvas lluviosas trop- icales en la cuenca amazonica es nece- sario establecer grandes reservas natur- ales que provean sitios de estudios y preservacion de esta rica biota para la humanidad. LITERATURE CITED Anderssox, L. G. 1899. Catalogue of Linnean type-specimens of snakes in the Royal Museum in Stockholm. Bih. Svenska Vet.-Akad. Hand!., 24(6): 1-35. Andrews, R., Raxd, A. S. 1974. Reproductive effort in anoline hzards. Ecology, 55: 1317-1.327. AxTHOXY, J. 1955. Essai .sur revolution anato- mique de I'appareil venimeux des ophi- diens. Ann. Sci. Xat. (Zool. ), Ser. 11, 17:7-53. AxELROD, D. I., Bailey, H.P. 1968. Cretaceous dinosaur extinction. Evolution, 22:595- 611. Bailey, J. R. 1966. Modes of evolution in Xew World opisthoglyph snakes. Mem. Inst. Butantan, 33:67-72. Beebe, W. 1944. Field notes on the lizards of Kartabo, British Guiana, and Caripito, Venezuela. Part 2, Iguanidae. Zoolog- ica, 29:195-216. Beebe, W. 1945. Field notes on the lizards of kartabo, British Guiana, and Caripito, Venezuela. Part 3, Teiidae, Amphis- baenidae and Scincidae. Ibid., 30:7-32. Beek, K. J., Bha.mao, D. L. 1969. Nature and geography of South American soils. Jn FrriKAU, et al. (Eds.). Biogeography and ecology in South America. Junk, The Hague, 1:82-112. Brame, a. H., Jr., Wake, D. B. 1963. The salamanders of South America. Contr. Sci., Los Angeles Co. Mus., (69): 1-72. Bkame, a. H., Jr., Wake, D. B. 1971. New species of salamanders (Genus Bolito- nlossa) from C'^lombia, Ecuador, and Panama. Ibid., (219): 1-34. Breoer, C .M., Jr. 1946. Amphibians and rep- tiles of the Rio Chucunaque drainage, Darien, Panama, with notes on their life histories and habits. Bull. Amer. Mus. Nat. Hist., 86:375-436. 346 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Brown, K. S., Jr. 1972. The heliconians of Brazil ( Lepidoptera : N\mp]ialidae ) . Part III. Ecology and biology of Hcli- coniiis nattcrcri, a key primitive species near extinction, and connnents on the evolutionary development of Heliconitis and Eueides. Zoologica, 57:41-69. Browx, K. S., Jr., Mielke, O. II. H. 1972. The heliconians of Brazil (Lepidoptera: Nyniphalidae). Part II. Introduction and general comments, with a supple- mentary revision of tlie tribe. Ibid., 57: 1-40. Cald\\'ell, J. P. 1973. Tropical tree frog com- mimities: Patterns of reproduction, size, and utilization of structural habitat. Un- pub. Ph.D. Dissertation, Uni\'. Kansas: 1-197. Cei, J. M. 1968. Distribution et specialisation des batraciens Sudamericans. In Dela- MARE DeBOUTTEVILLE, C, RaPOPORT, E. H. (Eds.). Biologic L'Amerique Aus- trale. IV. Cent. Nat. Recher. Sci. Paris: 199-214. Cody, M. L. 1974. Competition and the struc- ture of bird communities. Princeton Univ. Press, New Jersey: 1-318. Cody, M. L. 1975. Towards a dieory of conti- nental species diversities. In Cody, M. L., Diamond, J. M. (Eds.). Ecol- ogy and evolution of communities. Har- vard Univ. Press, Cambridge: 2 14-257. Cole, L. C. 1954. The population conse- quences of life history phenomena. Quart. Rev. Biol., 29:103-1.37. CoLWELL, R. K., Futuyma, D. J. 1971. On the measurement of niche breadth and overlap. Ecology, 52:567-586. CoNAXT, R. 1955. Saurian shell crusher. Na- ture Mag., Feb. 1955, unpaginated 2 pp. CoNNELL, J. H. 1975. Some mechanisms pro- ducing sti"ucture in natural communities: a model and e\idence from field experi- ments. In Cody, M. L. and Diamond, J. M. (Eds.). Ecology and evolution of communities. Harvard Press, Cam- bridge: 460-490. Cracraft, J. 1974. Continental drift and vertebrate distribution. Ann. Rev. Ecol. Syst., 5:215-261. Crump, M. L. 1971. Quantitative analysis of the ecological distribution of a tropical heri^etofauna. Occas. Paper Mus. Nat. Hist. Univ. Kansas, ( 3 ) : 1-62. Crump, M. L. 1972. Territoriality and mating behavior in Dcndrohates ^rantilifcrus ( Anura : Dendrobatidae ) . Herpetolog- ica, 28:195-198. Crump, Martha L. 1974. Reproductive strat- egies in a tropical amnan community. Misc. Publ. Mus. Nat. Hist. Univ. Kan- sas, (61): 1-68. Damuth, J. E., Fairbridge, R. W. 1970. Equatorial Atlantic deep-sea sand and ice-age aridity in tropical South Amer- ica. Bull. Geol. Soc. Amer., 81:189-206. Dexgo, G. 1969. Problems of tectonic rela- tions between Central America and tlie Caribbean. Trans. Gulf Assoc. Geol., 19:311-320. DiETZ, R. S., HoLDEx, J. C. 1970. Reconstiuc- tion of Pangaea: breakup and dispersion of continents, Permian to present. Jour. Geophys. Res., 75:4939-4956. Dixon, James R. 1973. A systematic review of the teiid lizards, genus Bachia, with remarks on Hctcrodactyliis and Anoto- saura. Misc. Publ. Mus. Nat. Hist. Univ. Kansas (57): 1-47. Dixon, J. R., Soini, P. 1975. The reptiles of the upper Amazon Basin, Iquitos region, Peru. I. Lizards and amphisbaenians. Contr. Biol. Geol. Milwaukee Publ. Mus., (4): 1-58. Dixon, J. R., Soini, P. 1976. The reptiles of tlie upper Amazon Basin region, Peru. II. Crocodilians, Turtles and Snakes. Ibid., (12): 1-91. Dobzhansky, T. 1950. E\olution in the trop- ics. Amer. Sci., 38:209-221. DuELLMAN, W. E. 1958. A monographic study of the colubrid snake genus Leptodeira. Bull. Amer. Mus. Nat. Hist., 114:1-152. DuELLMAN, W. E. 1965. A biogeographic ac- count of the herpetofauna of Michoacan, Mexico. Univ. Kansas Publ. Mus. Nat. Hist., 15:627-709. DuELLMAN, W. E. 1966. The Central Ameri- can Herpetofauna: An ecological per- .spective. Copeia, 1966(4):700-719. DuELLMAN, W. E. 1967. Comtship isolating mechanisms in Costa Rican hylid frogs. Herpetologica, 23: 169-183. DuELLMAN, W. E. 1968. The taxonomic status of some American hylid frogs. Ibid., 24:194-209. DuELLMAN, W. E. 1970a. Identity of the South American hylid frog Garbcana garhei. Copeia, 1970(3):534-538. DuELLMAN, W. E. 1970b. The hylid frogs of Middle America. Monog. Mus. Nat. Hist. Univ. Kansas, (l):l-753. DuELLMAN, W. E. 1971a. A taxonomic review of South American hylid frogs, genus Fhnjnohijas. Occas. Pap. Mus. Nat. Hist. Univ. Kansas, (4):1-21. DuELLMAN, W. E. 1971b. The identities of some Ecuadorian Inlid frogs. Herpeto- logica, 27:212-227. ' DuELLMAN, W. E. 1972a. A new species of //(//« from Amazonian Ecuador. Copeia, 1972(2) :265-271. EQUATORIAL HERPETOFAUNA 347 DuELLMAX, W. E. 1972b. Soutli American frops of the Hyla rostnita jfnnip (Am- phibia, Anura, Hyhdae). Zool. Mede- del., 47:177-192. DuELLMAX, W. E. 1972c. The systematic status and life liistoiy of H\ila rhodo- pcpla Giinther. Heipetologica, 28:369- 375. DuELLMAX, W. E. 1973a. Nest-building lulid frogs. HISS News-Jour., 1:72, 75. DuELLMAx, W. E. 1973b. Frogs of the IhjJa geographica group. Copeia, 1973(3): 515-533. DuELLMAX, W. E. 1973c. Descriptions of new lizards from tlie upper Amazon Basin. Herpetologica, 29:228-231. DuELLMAX, W. E. 1974a. A reassessment of the taxonomic status of some Neotropical hylid frogs. Occas. Pap. Mus. Nat. Hist. Univ. Kansas, (27): 1-27. DuELLMAx, W. E. 1974b. Ta.\onomic notes on PhijUomcchisa (Anura: Hylidae) from the upper Amazon Basin. Herpetologica, 30:105-112. DuELLMAx, W. E., Crump, M. L. 1974. Speci- ation in frogs of the Hijla pariiceps group in tlie upper Amazon Basin. Oc- cas. Pap. Mus. Nat. Hist. Univ. Kansas, (23): 1-40. DuELLMAx, W. E., Lescure, J. 1973. Life history and ecology of the hylid frog Ostcoccphahts tautinus, with obser\a- tions on lar\al beha\ior. Ibid., (13): 1-12. Duellmax, W. E., Trueb, L. 1976. The sys- tematic status and relationships of the h>lid frog Nijctimautis rugiccps. Ibid., (58): 1-14. Duxx, E. R. 1944. The lizard genera Anadia and Ptijchoglossiis in Colombia. Cal- dasia 3:63-68. Duxx, E. R. 1948. American frogs of the fam- ily Pipidae. Amer. Mus. No\itates, (1384): 1-13. Duxx, E. R. 1949. Notes on the South Amer- ican frog genus Edulorhina. Ibid., (1416): 1-10. Edmund, A. G. 1969. Dentition. In CANS, C. (Ed.). Biology of the Reptilia, I. Academic Press, London: 117: 200. Edwards, S. R. 1974. Taxonomic notes on Soutli American dendrobatid frogs of the genus Colostcthus. Occas. Pap. Mus. Nat. Hist. Univ. Kansas, (30): 1-14. EiDT, R. C. 1969. The climatology of South America. In Fittkau, E. J., ct al. (Eds.). Biogeography and ecology in South America. Junk, The Hague, 1: 54-81. Emiliaxi, C, Gaertner, S., Lidz, B. 1972. Neogene sedimentation on the Blake Plateau and the emergence of the Cen- tral American isthmus. Paleogeog. Pa- leoclim. Paleoecol., 11:1-10. EsTES, R. 1975. Fossil Xenoptis from the Paleocene of South America and the zoogeography of pipid frogs. Herpeto- logica, 31:263-278. EsTES, R. Price, L. I. 1973. Iguanid lizard from the Upper Cretaceous of Brazil. Science, 180:748-751. EsTES, R., Reic, O. a. 1973. The early fossil record of frogs. A re\iew of the evi- dence. In Vial, J. E. (Ed.). Evolu- tionary biology of the anurans: contem- porary research on major problems. Univ. Missouri Press, Columbia: 11-63. Estes, R., Wake, M. H. 1972. The first fossil record of caecilian amphibians. Nature, 239:228-231. Fisher, R. A. 1930. The genetical theory of natural selection. Ed. 2. Dover Press, New York: 1-287. Fitch, H. S. 1968. Temperature and behavior of some equatorial lizards. Herpeto- logica, 24:35-38. Fitch, H. S. 1970. Reproductive cycles in lizards and snakes. Misc. Publ. Mus. Nat. Hist. Univ. Kansas, (52): 1-247. FouQUETTE, M. J., Jr. 1960. Isolating mechan- isms in three sympatric tree frogs in the Canal Zone. Evolution, 14:484-497. FuEXTES, E. R. 1976. Ecological convergence of lizard communities in Chile and Cali- fornia. Ecology, 57:3-17. FuGLER, C. M., Schwaxer, T. D. 1968. Sex- ual dichromatism in the genus Uraccn- troii Kaup in eastern Ecuador. Heqjeto- logica 24:253-255. Gadgil, M., Bossert, W. H. 1970. Life his- storical consequences of natm-al selec- tion. Amer. Natur., 104:1-24. Goodlaxd, R. J. A., Ihwtx, H. S. 1975. Ama- zon Jungle: Green Hell to red desert. Elsevier Sci. Publ. Co., New York: 1-155. GooDMAX, D. E. 1971. Territorial behavior in a neotropical frog, Dcndrobatcs grann- liferus. Copeia, 1971(2) :365-370. GosxER, K. L. 1960. A simplified table for staging anuran embryos and larvae witli notes on identification. Herpetologica, 16:183-190. Graxt, N. K. 1971. South Atlantic, Benue Trough, and Gulf of Guinea Cretaceous triple junction. Bull. Geol. Soc. Amer., 82:2295-2298. Haffer, J. 1969. Speciation in Amazonian forest birds. Science, 165:131-137. 348 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Haffer, ]. 1970. Geologic-climatic history and zoogeographic significance of the Uraba region in nortliwestern Colombia. Cal- dasia, 10:60.3-636. Haffer, J. 1974. Avian speciation in tropical Sou til America. Publ. Nuttall Omith. Club, (14): 1-390. Haffer, J. 1975. Avifauna of northwestern Colombia, South America. Bonner Zool. Monog., (7): 1-182. Hairstox, N. G. 1969. On the relative abun- dance of species. Ecology, 50:1091- 1094. Harrixgtox, H. J. 1962. Paleogeographic de- velopment of South America. Bull. Amer. Assoc. Petrol. Geol., 46:1773- 1814. Hexxessy, p. a. 1968. A study of pigmenta- tion in a hylid frog, Ht/Ia geop.raphica ( Spix, 1824 ) , with special reference to changes during ontogeny. Unpub. Ms. Thesis, Univ. Kansas: 1-72. Heyer, W. R. 1972. The status of Leptodac- tijlus pumilio Boulenger (Amphibia, Leptodactylidae ) and the description of a new species of Leptodactyhts from Ecuador. Contr. Sci. Los Angeles Co. Mus., (231): 1-8. Heyer, W. R. 1973. Systematics of the mar- moratus group of the frog genus Lepto- dactyhts (Amphibia, Leptodactylidae). Ibid., (251): 1-50. Heyer, W. R. 1974a. Relationships of the marmoratus species group (Amphibia, Leptodactylidae) within tlie subfamily Leptodactylinae. Ibid., (253): 1-46. Heyer, W. R. 1974b. Vanzoliniu.s, a new genus proposed for Lcptodactylus dis- codactylus (Amphibia: Leptodactyli- dae). Proc. Biol. Soc. Washington, 87: 81-90. Heyer, W. R. 1976. Studies in larval amphib- ian habitat partitioning. Smithsonian Contr. Zool., 242:1-27. HOLDEN, J. C, DiETZ, R. S. 1972. Galapagos core, NazCoPac triple junction and Car- negie/Cocos ridges. Nature, 235:266- 269. HoOGMOED, M. S. 1973. Notes on the herpeto- fauna of Surinam IV. The lizards and amphisbaenians of Surinam. Junk, The Hague: 1-419. HuRLBERT, S. H. 1971. The nonconcept of species diversity: a critique and alterna- tive parameters. Ecology, 52:577-586. HuTCHixsox, G. E. 1957. Concluding remarks. Cold Spring Harbor Sym. Ouant. Biol., (22):415-427. HuTCHixsON, G.E. 1959. Homage to Santa Rosalia, or why are there so many kinds of animals. Amer. Nat., 931:145-159. IxGER, R. F., Greex-^berg, B. 1966. Annual reproductive patterns of lizards from a Bornean rain forest. Ecology, 47:1007- 1027. Jaxzex, D. H. 1973. Sweep samples of tropi- cal foliage insects: Effects of seasons, vegetation types, elevation, time of day, and insularity. Ibid., 54:687-708. Larsox, R. L., Ladd, J. W. 1973. Evidence for tlie opening of the South Atlantic in the early Cretaceous. Nature, 246:209- 212. Laurext, R. F. 1973. A parallel survey of equatorial amphibians and reptiles in Africa and Soutli America. In Meggers, B. J., Ayexsu, E. S., Duckworth, W. D. (Eds.). Tropical forest ecosystems in Africa and South America: A compara- tive review. Smithsonian Inst. Press, Washington, D.C.: 259-266. Leox, J. R., Ruiz, L. J. 1971. Reproduccion de la lagartija, Amciva hifrontata ( Sau- ria: Teiidae). Carib. J. Sci., 11:195- 201. Lescure, J. 1976. Etude de deu.x tetards de Phyllobates ( Dendrobatidae) : P. fe- moralis (Boulenger) et P. pictiis ( Bib- ron). Bull. Soc. Zool. France, 101(2): 299-306. Levixs, R. 1968. Evolution in changing en- vironments. Princeton Univ. Press, Princeton, New Jersey: 1-120. Levitox, a. E., Axdersox, C. 1966. Further comments on the beha\'ior of the Pana- manian microteiid Echinosaitra honida. Herpetologica, 22:160. Lloyd, M., Ghelardi, R. J. 1964. A table for calculating the "equitability" component of species diversity. Jour. Anim. EcoL, 33:217-225. LovERiDGE, A. 1929. A new Anadia from Co- lombia with remarks on other members of the genus. Proc. Biol. Soc, Washing- ton 42:99-102. LowE-McCoxxELL, R. H. 1975. Fish commu- nities in tropical freshwaters. Their dis- tribution, ecology, and evolution. Long- man, New York. 1-338. LuTZ, B. 1960. Fighting and an incipient no- tion of territory in male tree frogs. Co- peia, 1960(1 ):6I-63. Lyxch, J. D. 1968. Two new frogs of the genus Kleuthcrodactylus from eastern Ecuador (Amphibia: Leptodactylidae). J. Herp., 2:129-135. Lyxch, J. D. 1970. A new eleutherodactyline frog from Amazonian Ecuador. Proc. Biol. Soc. Washington, 83-221-225. EQUATORIAL HERPETOFAUNA 349 Lynch, J. D. 1974. New species of frogs (Lep- todactylidae: Elcutlwroclactyhis) from the Amazonian lowlands of Ecuador. Occas. Pap. Mus. Nat. Hist. Univ. Kan- sas, (31): 1-22. Lynch, J. D. 1975. The identity of the frog Elcuthcrodactylus conspicillatus ( Gun- ther), with descriptions of two related species from northwestern South Amer- iac (Amphibia, Leptodactylidae) Contr. Sci. Nat. Hist. Mus. Los Angeles Co.; (272): 1-19. Lynch, J. D., Duellman, W. E. 1973. A re- view of the centrolenid frogs of Ecuador, with descriptions of new species. Occas. Pap. Mus. Nat. Hist. Uni\'. Kansas, (16): 1-66. MacArthur, R. H. 1957. On the relative abundance of bird species. Proc. Nat'l Acad. Sci., 43:293-295. MacArthur, R. H. 1965. Patterns of species diversity. Biol. Rev., 40:510-533. MacArthur, R. H. 1969a. Patterns of commu- nities in the tropics. Biol. T- Linnean Soc, 1:19-30. MacArthur, R. H. 1969b. Species packing and what competition minimizes. Proc. Nad. Acad. Sci., 64:1369-1371. MacArthur, R. H. 1970. Species packing and competitive equilibrium for many spe- cies. Theoret. Pop. Biol., 1:1-11. MacArthur, R. H. 1971. Patterns in terres- trial bird communities. In Earner. D. S., et al. (Eds.). A\ian biology, I. Academic Press, New York: 189-221.' MacArthur, R. H. 1972. Geographical ecol- ogy. Harper and Row, New York: 1-269. MacArthur, R. H., Wilson, E. O. 1963. An equilibrium theory of insular zoogeog- raphy. Evolution, 17:373-387. MacArthur, R. H., Wilson, E. O. 1967. The theory of island biogeography. Prince- ton Univ. Press, Princeton, New Jersey: 1-120. Malfait, B. T., Dinkelman, M. G. 1972. Circum-Caribbean tectonic and igneous acti\'ity and the evolution of the Carib- bean plate. Bull. Geol. Soc. Amer., 82:251-272. Margalef, D. R., 1968. Perspectives in eco- logical tlieory. Uniw Chicago Press, Chicago: 1-111. Marshall, A. J., Hook, R. 1960. The breeding biology of equatorial vertebrates: repro- duction of the lizard Agama agarna lio- notus Boulenger at lat. O'^Ol'N. Proc. Zool. Soc. London, 134:197-205. May, R. M. 1975. Patterns of species abun- dance and diversity. 7/! Cody, M. L.. Diamond, J. M. (Eds.). Ecology and evolution of connnunities. Harvard Univ. Press, Cambridge :81-120. Medem, F. 1952. Palcosiichus tiigonattis (Schneider) en Colombia (Noticia pre- liminar). Lozania, 5:1-12. Medem, F. 1963. Osteologia craneal, distribu- cion geografico. y ecologia de Mclano- stichiis niger (Spi.x) (Crocodylia: Alli- gatoridae). Rev. Acad. Colombiana Cien. Exact. Fisicas Nat., 12(45): 1-19. Medem, F. 1969. Estudios adicionales sobre los Crocodylia y Tcstudinata del Alto Caqueta y Rio Caguan. Caldasia, 10: 329-353. Miller, R. S. 1969. Competition and species diversity. In Diversity and Stability in ecological systems. Brookhaven Svmp. Biol., (22):63-70. Moreau, R. E. 1963. Mcissitudes of the Afri- can biomes in the late Pleistocene. Proc. Zool. Soc. London, 141:,395-421. Moreau, R. E. 1969. Climatic changes and tlie distribution of forest \'ertebrates in West Africa. J. Zool., 158:39-61. MiJLLER, P. 1973. The dispersal centers of terrestrial vertebrates in the Neotropical Realm. Biogeographica, 2. Junk, The Hague: 1-244. Myers, C. W. 1965. Biology of the ringneck snake, Diadopliis punctatus, in Florida. Bull. Florida St. Mus. 10:43-90. Myers, C. W. 1974. The systematics of Rlia- dinaca ( Colubridae), a genus of New World Snakes. Bull. Amer. Mus. Nat. Hist., 153:1-262. Myers, C. W., Rand, A. S. 1969. Checklist of amphibians and reptiles of Barro Colo- rado Island, Panama, with comments on faunal change and sampling. Smith- sonian Contr. Zool., (10):1-11. Neill, W. T. 1964. Vi\iparity in snakes: some ecological and zoogeographic considera- tions. Amer. Nat., 98:35-55. Oliver, J. A. 1947. The seasonal incidence of snakes. Amer. Mus. Novitates, (1363): 1-14. Oliver, J. A. 1948. The relationships and zoo- geography of the genus Thalcrophis Oliver. Bull. Amer. Mus. Natur. Hist., 92:157-280. Peters, J. A. 1956. An analysis of variation in a South American snake, Catesby's snail-sucker (Dipsas cateshyi Sentzen). Amer. Mus. No\itates, (1783): 1-41. 350 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Peters, J. A., Orces-V., G. 1960. Lcptophis ciiprctis Cope, a valid South American colubrid species. Beitr. Neotrop. Fauna, 2:139-141. Peters, J. A., Doxoso-B.-vrros, R. 1970. Cata- logue of tlie Neotropical Squamata: Part II. Lizards and amphisbaenians. Bull. U.S. Natl. Mus. 297:1-293. Peters, J. A., Orejas-Miraxda, B. 1970. Cat- alogue of tlie Neotropical Squamata: Part I. Snakes. Ibid., 297:1-347. Patrick, R. 1966. The Catherwood Founda- tion Peruvian-Amazon Expedition. Lim- nological Studies. Monog. Acad. Nat. Sci. Philadelphia, (14): 1-495. Petersen, U. 1958. Structure and unplift of the Andes of Peru, Bolivia, Chile, and adjacent Argentina. Bol. Soc. Geol. Peru, 33:57-143. PiANKA, E. R. 1966. Latitudinal gradients in species diversity: a review of concepts. Amer. Nat., 100:33-46. PiANKA, E. R. 1967. On lizard species diver- sity: North American flatland deserts. Ecology, 48:333-351. PiANKA, E. R. 1970. On r and K selection. Amer. Nat., 104:592-597. PiANKA, E. R. 1973. The structure of lizard communities. Ann. Rev. Ecol. Syst., 4:53-74. PiANKA, E.R. 1975. Niche relations of desert lizards, hi Cody, M. L., Diamond, J. M. (Eds.) Ecology and evolution of communities. Harvard Univ. Press, Cambridge:292-314. Pope, C. H. 1961. The giant snakes. Knopf, New York: 1-290. Prance, G. T. 1973. Phytogeographic support for the tlieory of Pleistocene forest ref- uges in the Amazon Basin, based on evidence from distribution patterns in Caryocaraceae, Chrysobalanaceae, Di- chapetalaceae and Lecytliidaceae. Acta Amazonica, 3(3):5-28. Presch, W. 1973. A review of the tegus, liz- ard genus Tupinambis (Sauria: Teiidae) from South America. Copeia, 1973(4)- 740-745. Pyburn, W. F., Heyer, W. R. 1975. Identity and call of the frog Lcptodactiilus stcno- dcma. Ibid., 1975 (3):585-587. Raub, G. B. 1960. On the unique sound pro- duction of the Surinam toad, pipa pipa. Ibid., 1960 (4):368-369. Rabu, G. B., Rabb, M. S. 1960. On the mating and egg-laying beha\ior of the Surinam toad, Pipa pipa. Ibid., 1960 (4):271- 276. Rabb, G. B., Rabb, M.S. 1963. Additional ob- servations on breeding behavior of the Surinam toad, Tipa pipa. Ibid., 1963 (4):636-642. Rabb, G. B., Snedigar, R. 1960. Observations on breeding and development of the Surinam toad, Fipa pipa. Ibid., 1960 (l):40-44. Rand, A. S., Humphi^ey, S. S. 1968. Interspe- cific competition in the tropical rainfor- est: ecological distribution among liz- ards at Belem, Para. Proc. U.S. Natl. Mus., 125:1-17. Raven, P. H., Axelrod, D. I. 1972. Plate tec- tonics and Australasian paleobiogeog- raphy. Science, 176:1379-1385. Raven, P. H., Axelrod, D. I. 1974. Angio- spemi biogeography and past continental movements. Ann. Missouri Bot. Card., 61:539-673. Reig, O. 1968. Peuplement en vertebres tetra- podes de I'Amerique du sud. In Dela- MARE DeBOUTTEVILLE, C, RaPOPORT, E. H. (Eds.). Biologic I'Amerique Australe. IV. Cent. Nat. Recher. Sci. Paris: 214-260. Rivero, J. A. 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool., 126:1-207. Rosen, D. E. 1975. A vicariance model of Caribbean biogeography. Syst. Zool., 24:431-464. RosENzwEiG, M. L. 1975. On continental steady states of species diversity. In Cody, M. L., Diamond, J. M. (Eds.). Ecology and evolution of communities. Harvard Univ. Press, Cambridge: 121- 140. RossMAN, D. 1973. Miscellaneous notes on the South American water snake genus HcUcops. HISS News-Journal, 1:189- 191. RossMAN, D. 1976. Revision of the South American colubrid snakes of tlie Heli- cops pastazae complex. Occas. Pap. Mus. Zool. Louisiana State Univ., (50): 1-15. Salthe, S. N., Duellman, W. E. 1973. Quan- titative constraints associated with re- productive mode in anurans. In Vial, J. L. (Ed.). Evolutionary biology of the anurans: contemporary research on major problems. Univ. Missouri Press, Columbia: 229-249. Salthe, S. N., Mecham, J. S. 1974. Repro- ductive and courtship patterns. In Lofts, B. (Ed.). Physiology of Am- phibia, II. Academic Press, San Fran- cisco: 309-521. Saul, W. G. 1975. An ecological study of fishes at a site in upper Amazonian Ecua- dor. Proc. Acad. Nat. Sci. Philadelphia, 127:93-134. EQUATORIAL HERPETOFAUNA 351 Savage, J. M. 1955. Descriptions of new colu- brid snakes, genus Atmctus, from Ecua- dor. Proc. Biol. Soc. Washington, 68: 11-20. Savage, J. M. 1960. A revision of the Ecua- dorian snakes of tlie cokibrid genus Atractiis. Misc. Publ. Mus. Zool. Univ. Mich., (112): 1-86 Savage, J. M. 1968. The dendrobatid frogs of Central America. Copeia, 1968 (4): 745-776. Savage, R. M. 1973. The geographic distribu- tion of frogs: patterns and predictions. In Vial, J. L. (Ed.). Evolutionary biology of the anurans: contemporary research on major problems. Univ. Mis- souri Press, Columbia: 351-445. Savage, J. M. 1974. The isthmian link and tlie evolution of Neotropical mammals. Contr. Sci. Nat. Hist. Mus. Los Angeles Co., (260): 1-51. Schmidt, K. P. 1952. The Surinam Coral Snake Micnirus surinamensis. Fieldiana, Zool. 34:25-34. Schmidt, K. P., Walker, W. R., Jr. 1943a. Peruvian snakes from the University of Arequipa. Zool. Ser. Field Mus. Nat. Hist. 24:279-296. Schmidt, K. P., Walker, W. R., Jr. 1943b. Snakes of tlie Peruvian coastal region. Ibid., 24:297-324. ScHOENER, T. W. 1965. The evolution of bill size differences among sympatric con- generic species of birds. Evolution, 19: 189-213. ScHOENER, T. W. 1967. The ecological signifi- cance of sexual dimorphism in size in the lizard Anolis conspersus. Science, 155:474-477. ScHOENER, T. W. 1969. Size patterns in West Indian Anolis lizards: I. Size and spe- cies diversity. Syst. Zool., 18:386-401. ScHOENER, T. W. 1970. Size patterns in West Indian Anolis lizards: II. Correlations with the sizes of particular sympatric species — displacement and convergence. Amer. Nat., 104:155-174. ScHWABE, G. H. 1969. Towards an ecological consideration of the South American continent. In Fittkau, E. J., et al. (Eds.). Biogeography and ecology in South America. Junk, The Hague, 1: 113-136. Sexton, O. J., Ortleb, E. P., Hathaway, L. M., Ballinger, R. E., Light, P. 1971. Reproductive cycles of tliree spe- cies of anoline lizards from Isthmus of Panama. Ecology, 52:201-215. Sexton, O. J., Turner, O. 1971. The repro- ductive cycle of a neotropical lizard. Ibid., 52:159-164. Shannon, C. E. 1948. The mathematical theory of communication. In Shannon, C. E., Weaver, W. The mathematical theory of communication. Univ. Illinois Press, Urbana: 1-117. Sherbrooke, W. C. 1975. Reproductive cycle of a tropical lizard, Neusticunis ecpleo- piis Cope, in Peru. Biotropica, 7:194- 207. Sill, W. D. 1968. The zoogeography of the crocodilia. Copeia, 1968 (l):76-88. Silverstone, p. a. 1975. A revision of the poison-arrow frogs of the genus Dendro- bates Wagler. Nat. Hist. Mus. Los An- gles Co. Sci. Bull., (21): 1-55. Simmons, J. E. 1975. The female reproductive cycle of the teiid lizard A7neiva ameiva petersii Cope. Herpetologica, 31:279- 282. Simpson, G. G. 1950. History of the fauna of Latin America. Amer. Sci., 38:361-389. Stebbins, R. C, Hendrickson, J. R. 1959. Field studies of Amphibians in Colom- bia, South America. Univ. California Publ. Zool., 56:497-540. Stuart, L. C. 1948. The amphibians and rep- tiles of Alta Verapaz, Guatemala. Misc. Publ. Mus. Zool. Univ. Michigan, (69): 1-109. Taylor, E. H., Peters, J. A. 1974. The caecil- ians of Ecuador. Univ. Kansas Sci. Bull., 50:333-346. Tedford, R. H. 1974. Marsupials and the new paleogeography. In Ross, C. A. (Ed.). Paleogeographic provinces and provin- ciality. Soc. Econ. Paleo. Miner, Spec. Publ., (21): 109-126. Terborgh, J. 1975. Faunal equilibria and the design of wildlife preserves. In GoL- LEY, F. B., Medina, E. (Eds.). Trop- ical ecological systems. Trends in ter- restrial and aquatic research. Springer- Veriag, New York: 369-380. Tinkle, D. W. 1969. The concept of repro- ductive effort and its relation to the evo- lution of life histories of lizards. Amer. Natur., 103:501-516. Tinkle, D. W., Wilbur, H. M., Tilley, S. G. 1970. Evolutionary strategies in lizard reproduction. Evolution, 24:55-74. Truer, L. 1974. Systematic relationships of neotropical homed frogs, genus Hemi- phractus (Anura: Hylidae). Occas. Pap. Nat. Hist. Univ. Kansas, (29): 1-60. Trueb, L., Duellman, W. E. 1971. A synopsis of neotropical hylid frogs, genus Osteo- cephalus. Ibid., ( 1 ) : 1-47. TscHOPP, H. J. 1953. Oil explorations in the Oriente of Ecuador, 1938-1950. Bull. Amer. Assoc. Petrol. Geol., 37:2303- 2347. 352 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Turner, F. B. 1962. The demography of frogs and toads. Quart. Rev. Biol., 37:303- 314. UzzELL, M. 1965. Teiid Hzards of the genus Echinosauia. Copeia, 1965 (l):82-89. Van der Hammen, T. 1966. The Pliocene and Quaternary of the Sabana de Bogota (the Tilata and Sabana formations). Geol. Mijnbouw, 45:102-109. Van der Hammen, T. 1975. The Pleistocene changes of vegetation and climate in tropical South America. J. Biogeogr., 1:3-26. Van der Hammen, T., Gonzalez, E. 1964. A pollen diagram from the Quaternary of the Sabana de Bogota (Colombia) and its significance for the geology of the northern Andes. Geol. Mijnbouw, 43: 113-117. Vanzolini, E. 1951. Amphisbaenia fuliginosa. Contribution to the knowledge of the Brazilian lizards of the family Amphis- baenidae Gray, 1825. 6. On the geo- graphic distribution and difi^erentiation of Amphisbaena fuliginosa Linne. Bull. Mus. Comp. Zool., 106:1-67. Vanzolini, P. E. 1961. Notas bionomicas so- bre Dracaena guianensis no Para (Sau- na, Teiidae). Pap. Avul. Dept. Zool., Sao Paulo, 14:237-241. Vanzolini, P. E. 1968. Geography of the South American Gekkonidae (Sauria). Arq. Zool. Sao Paulo, 17:85-111. Vanzolini, P. E. 1973. Paleoclimates, relief, and species multiplication in equatorial forests. In Meggers, B. J., Ayensu, E. S., Duckworth, W. D. (Eds.). Trop- ical forest ecosystems in Africa and South America: A comparative review. Smithsonian Inst. Press, Washington, D.C.:255-258. Vanzolini, P. E., Ab'saber, A. N. 1968. Di- vergence rate in South American lizards of the genus Liolaemus ( Sauria, Iguani- dae). Pap. Avul. Zool. Sao Paulo, 21: 205-208. Vanzolini, P. E., Williams, E. E. 1970. South American anoles: The geographic differentiation and evolution of the Ano- lis chrtjsolepis species group ( Sauria, Iguanidae). Arq. Zool. Sao Paulo, 19: 1-124. ViTT, L. J., Congdon, J. D. 1975. Reproduc- tive effort and body shape in lizards: resolution of a paradox. Heip. Review, 6:74 (abstract). VuiLLEMiER, B. S. 1971. Pleistocene changes in tlie fauna and flora of South America. Science, 173:771-780. Wake, D. B. 1966. Comparative osteology and evolution of the lungless salamanders, family Plethodontidae. Mem. South California Acad. Sci., 4:1-111. Walker, C. F. 1973. A new genus and species of microhylid frog from Ecuador. Occas. Pap. Mus. Nat. Hist. Univ. Kansas, (20): 1-7. Walker, C. F., Duellman, W .E. 1974. De- scription of a new species of microhylid frog, Chiasmocleis, from Ecuador. Ibid., (26): 1-6. Whitmore, F. C, Jr., Stewart, R. H. 1965. Miocene mammals and Central American seaways. Science, 148:180-185. Whittaker, R. H. 1970. Communities and ecosystems. Macmillan Co., London :1- 162. Wilbur, H. 1972. Competition, predation, and tlie structure of the Ambijstoma - Rana sylvatica community. Ecology, 53:3-21. Wilbur, H. M., Tinkle, D. W., Collins, J. P. 1974. Environmental certainty, trophic level, and resource availability in life history evolution. Amer. Nat., 108:805- 817. Williams, G. C. 1966. Adaptations and natural selection. Princeton Univ. Press, Prince- ton, New Jersey: 1-307. Zaret, T. M. 1976. Fish in the inland tropics. Science, 192:672-673. AVAILABLE MISCELLANEOUS PUBLICATIONS IN HERPETOLOGY UNIVERSITY OF KANSAS MUSEUM OF NATURAL HISTORY 52. Reproductive c\cles in lizards and snakes. B\' Henry S. Fitcli. Pp. 1-247, 16 fig- ures in text. June 19, 1970. Paper bound, $6.15 postpaid. 53. Evolutionary relationships, osteology, and zoogeography of leptodactvloid frogs. By John D. Lynch. Pp. 1-238, 13i figures in text. June 30, 1971. Paper bound, $5.95 postpaid. 55. Middle American lizards of the genus Ameiva (Teiidae) with emphasis on geo- graphic variation. By Arthur C. Echternacht. Pp. 1-86, 28 figures in text. De- cember 14, 1971. Paper bound, $2.15 postpaid. 57. A systematic review of the Teiid lizards, genus Bachia, with remarks on Heter- odact)'lus and Anotosaura. By James R. Dixon. Pp. 1-47. 15 figures in text. February 2, 1973. Paper bound, $1.15 postpaid. 59. Systematics and evolution of the Andean lizard genus Pholidobolus (Sauria: Teiidae). By Richard R. Montanucci. Pp. 1-52, 8 figures in text. May 14, 1973. Paper bound, $1.30 postpaid. 61. Reproductive strategies in a tropical anuran community. By Martha L. Crump. Pp. 1-68, 13 figures in text. November 15, 1974. Paper bound, $1.70 postpaid. 62. A demographic study of the ringneck snake (Diadophis punctatus) in Kansas. Pp. 1-53, 19 figures in text. April 3, 1975. Paper bound, $1.30 postpaid. 65. The biology of an equatorial herpetofauna in Amazonian Ecuador. By William E. Duellman. Pp. 1-352, 198 figures in text, 4 color plates. August 30, 1978. Paper bound, $15.00 postpaid. MONOGRAPHS IN HERPETOLOGY 1. The hylid frogs of middle America. William E. Duellman. Pp. 1-753, 324 text figures, 72 color plates. 1970. Two volumes, cloth bound, $15.00 post^oaid. 4. Recollections of an herpetologist. By Edward H. Taylor, A. Byron Leonard, Ho- bart M. Smith, George R. Pisani. Pp. 1-160, 6 figures in text, 1 plate. Paper bound, $6.00 postpaid. c:^, cme Bookbinding Co.. Inc. 300 Summer Stfe«jt B«8ton, Mass. 0??!') Date Due