<. io^- XU^t^^^-^^^ MUS. COM P. ZOOU. LIBRARY OF KANSAS MISCELLANEOUS UNIVERSITY PUBLICATION MUSEUM OF NATURAL HISTORY SEP 5 1978 NO. 65 HARVARD UNIVERSITY
The Biology of an Equatorial Herpetofauna 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 142 143 Hyla funerea ( Cope ) 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 coriacea Peters Phyrnohyas ( ) 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 208 208 Mabitya mabouya" (Lacepede) 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) 234 235 Clelia clelia clelia ( Daudin ) 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) 255 255 Rhadinaea brevirostris ( Peters ) 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 Today many biologists are concerned moon and is carrying out exploratory primarily with theoretical aspects of pop- probes of outer space; yet there remain ulation biology, ecology, and evolution. on earth two nearly unknown ecosystems Models of species diversity, niche —the depths of the oceans and the low- breadth and overlap, resource partition- land tropical rainforest. Technological ing, competition, predator-prey, and r problems limit the exploration of oceans, and K selection abound in the literature. but the tropical lowlands? Is the "fore- But how do these models apply to the boding jungle," to which the exaggerated tropics? Where and how can they be writings of travelers and adventurers all- tested? Although inferences frequently too-often refer, beyond the limits of mod- are made in the literature, base line data ern technology? are wanting in most cases. In the Nineteenth Centuiy a few nat- The present report is an attempt to uralists ventured into the Amazon Basin present base line data and preliminary in South America. The vast collections interpretations. Hopefully, they will pro- and provocative writings of Henry W. vide future investigators with the in- Bates, Alexander von Humboldt, Richard formation and impetus for more inten- Spruce, and Alfred R. Wallace should sive and varied ecological and popula- have provided the impetus for continued tional studies, which in trnn will furnish and intensive exploration and study. Af- the bases for testing models. This report rica, with its relatively small Congo concerns the amphibians and reptiles of Basin, was the "dark continent" to be Santa Cecilia, a small area in the upper explored, colonized, and exploited, Amazon Basin. It deals with the amphib- whereas South America, which so richly ians and reptiles for two reasons. First, deserves the appelation given to Africa, I am a herpetologist and am most knowl- languished. Although the short-lived edgeable about those groups. Second, "rubber boom" late in the last century amphibians and reptiles are ideal organ- left an opera house in Manaus, the Ama- isms for faunal study in the tropics. They zon Basin remained a biological tierra are closely tied to the environment, usu- incognita. ally are independent of given species of numerous and Now the Amazon Basin is being ex- plants, and are sufficiently well known to be studied feasi- ploited—before it has been adequately enough choice of the area was explored and studied. Human popula- bly. The study tion pressure coupled with food and en- completely fortuitous. ergy shortages seem to make it "practi- During the past ten years numerous cal" to settle the vast region, constiTict publications dealing with the herpeto- roads, clear the forests, cultivate the soil, fauna of the Santa Cecilia region have and drill for oil. The lateritic soil is appeared. Many of these have been leached and eroded after two or three descriptions of new species: Eleuthero- crops; the rivers are polluted; and we dactyhis croceoinguinis and variahilis are losing forever a part of the world's (Lynch, 1968); E. orphnolaimtis (Lynch, biota. For the most part biologists are 1970); E. martiae, pcmhdus, and qiiaqtia- still ignorant of the fauna and flora of versus (Lynch, 1974); E. lanthanites the Amazon Basin and have little accu- (Lynch, 1975); CoJostethus satili (Ed- rate conception of the ecology of the wards, 1974); Hyla ])revifrons (Duell- rainforest. Ironically, the Biome Study man and Crump, 1974); }Iijlo cruentom- Committee of the International Biologi- ma (Duellman, 1972a); Centrolenella cal Program never succeeded in initiating midas, munozorum, and resplendens Chiasmo- a study of the tropical rainforest. (Lynch and Duellman, 1973); 10 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
the Oriente cleis anatipes (Walker and Duellman, portation between Quito and and also for other courtesies. Ec- 1974); Syncope antenori (Walker, 1973); many stationed at Emjaliokles cofonorum and Alopo^Iossiis uadorian anny personnel atriventris (Duellman, 1973c); and HeJi- Santa Cecilia aided us in many ways; I indebted to Colonel Ra- cops petersi (Rossman, 1976). Other am especially of mirez. Our work in was papers deal with the taxonomic status Quito greatly various taxa: Phrynohyas coriacea facilitated by personnel at the Universi- (Duellman, 1968); Hyla funerea (Duell- dad Catolica del Ecuador; I especially thank Father Gustavo Dra. man, 1971b); Hyla fl.(irhci (Duellman, Maldonado, 1970a); Hyla rhodopepla (Duellman, Olga Hcrrera de MacBryde, and the late 1972c); Hyla punctata, rossalleni, leuco- Francisco Leon. phyllata group and Sphaenorhynchus Muiiozlandia was "discovered" by carneus (Duellman, 1974a); and Ama- Charles M. Fugler, fomierly of Auburn zonian PhyUomcdusa (Duellman, 1974b). University. I will be forever grateful to Material from Santa Cecilia has been Fugler for his telephone call to me in used in the following systematic reviews: October 1966 in which he suggested Hyla geographica group (Duellman, Santa Cecilia as a study site. 1973b); Hyla parviceps group (Duell- The data and specimens upon which man and Crump, 1974); Hyla rostrata this report is based were accumulated group (Duellman, 1972b); Osteocepha- mostly by field parties from The Univer- lus (Trueb and Duellman, 1971); Hemi- sity of Kansas. I thank each person for phractus (Trueb, 1974); Nyctimantis their contribution and list them in de- (Duellman and Trueb, 1976); and Ecua- scending order of the amount of time dorian caecilians (Taylor and Peters, spent at Santa Cecilia: Martha L. 1974). Fitch (196S) reported on body CiTunp, John E. Simmons, Linda Trueb, temperatures of several species of lizards. Stephen R. Edwards, James W. Wad- a anal- H. Charles F. Crump ( 1974) provided thorough dick, Thomas Fritts, in the ysis of reproductive strategies Walker, John D. Lynch, Henry S. Fitch, anurans, and Simmons (1975) analyzed Abraham Goldgewicht K., Werner C. A. the female reproductive cycle in Ameiva Bokermann, Joseph T. Collins, Arthur C. ameiva. Echternacht, and Robert W. Henderson. In addition to these persons, all of whom ACKNOWLEDGMENTS were primarily concerned with studies on the herpetofauna, I am indebted to Field studies were initiated at Santa Philip S. Humphrey, Stephen R. Humph- Cecilia because of the existence primarily rey, Frank B. Cross, Gerald R. Smith, and of a base "Muiiozlandia." Our camp, especially William G. Saul for their con- work there was made possible by Ing. tributions made incidental to their stud- hous- Ildefonso Murioz B., who provided ies on the birds and fishes. Charles M. and a He and his ing laboratory. family Fugler, George Key, and Terry D. made it for us to devote full possible Schwaner collected amphibians and rep- time to studies while biological enjoying tiles at Santa Cecilia; their specimens but comfortable, accommodations rustic, were deposited at Auburn LTniversity in the jungle. For their gracious hospi- Museum and the Museum of Compara- which included numerous tality, fiestas tive Zoology at Harvard University. I am I sincere and despedidas, extend my indebted to Schwaner for field notes and miichissimas to the Muiioz fam- gracias photographs and to Robert H. Mount ily. and Ernest E. Williams for the loan of I am grateful to the personnel of Gulf specimens. Charles F. Walker kindly Oil Ecuador and Texaco Petroleum Com- made available material that he collected pany, Ecuador, for providing air trans- and deposited in the University of Mich- EQUATORIAL HERPETOFAUNA 11
of the was igan Museum of Zoology. Ronn Altig, part laboratory work sup- a from the General Re- Mississippi State University, collected at ported by grant Santa Cecilia in 1973-1975; I am grateful search Fund, The University of Kansas. to him for making material available to A grant from the National Science Foun- me. dation (GB-35483) made possible the Some of the limited information pro- completion of the research. vided herein on the geology and vegeta- During the preparation of this report tion was derived in part from infonna- I have been assisted by Jan Caldwell, tion provided by Richard Govett of the Albert Fisher, Julian C. Lee, John E. Texaco Petroleum Company, John Simmons, and especially Martha L. Dwyer of the Missouri Botanical Garden, Crump. I sincerely thank them for their and Bruce MacBryde, fonnerly of the careful and extensive work. I am in- Universidad Catolica del Ecuador. I was debted to James R. Dixon and John D. fortunate in being able to spend time in Lynch for critically reviewing the taxo- the field with each of them. nomic accounts and to Martha L. Crump The identification of the material col- for her provocative discussions and criti- lected at Santa Cecilia has been a long cal review of the manuscript. I am grate- and arduous task that required visits to ful to Juan R. Leon and Jaime E. Pefaur many museums; I am grateful to curators for translating the keys and summary of these collections in the United States, into Spanish. Many of the photographs Europe, and South America for their hos- in this report were taken by Crump and for to pitality and provision of working space. Simmons; I thank them permission Specimens of some groups have been reproduce the photographs, and I am studied by others, who have provided grateful to Simmons for his painstaking me with identifications and insights into work in the darkroom. the systematics of these groups. Thus, I Indeed I have been fortunate in hav- to Edward H. this am grateful Taylor (cae- ing the opportunities to undertake cilians), Federico Medem (pleurodire research and in having the cooperation of turtles), Charles W. Myers (Leima- so many associates. But my most cher- A. (lophis and Rhadinaea) , Douglas ished fortune is my wife, Linda Tmeb. W. Ronald Rossman (Helicops), Heyer She accompanied me on four trips to (LeptodacfyJits), and Charles F. Walker Santa Cecilia, assiduously worked in the and D. (microhylids), especially John field, and collaborated in several prelim- for his efforts on the difficult Lynch inary reports. She has provided many genus Eleutherodoctyhis. I am extremely ideas for the organization, analysis, and of the aid the she has appreciative provided by interpretation of the data, and late A. Peters, who on oc- But James many critically read the entire manuscript. shared with me his extensive are casions her major contributions to this report knowledge of the Ecuadorian herpeto- her illustrations, the usefulness of which fauna. less colorful shall, I fear, long outlive my field studies were a labor of The variously sup- prose. Piers has been love, ported by the Herpetological Research which I can never repay in kind. Fund, F. William Saul Fund, and the Watkins Museum of Natural History SCOPE OF STUDY Grants, all of The University of Kansas. The material herein is the Crump's field work in 1971-72 was sup- presented result of 48 man months of field work ported by the National Science Founda- Cecilia on the Rio tion (GB-29557). Visits to European at Santa Aguarico, Ecuador, smaller museums were made possible by a grant Provincia Napo, plus data from from the Penrose Fund of the American collections of specimens and four localities along the Rio Philosophical Society (No. 5063), and nearby 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 described in detail, and most never have Santa Ceciha consisted of about 3 square been illustrated. I present descriptions kilometers. During the course of the field of the animals based on living colors and work, efforts were made to collect series illustrations of most of the species. The of each species from every month of the tadpoles of 46 frogs are described. As an year and to obtain data on the micro- aid to the identification of species of habitat, activity, and life history of each amphibians and reptiles in the upper species. As a result of the field work, Amazon Basin, illustrated keys are of- 9035 specimens (+292 lots of tadpoles) fered in English and Spanish. representing 185 species were obtained; In attempting to analyze reproduc- 7765 lots of tive and I have sum- specimens ( +2
44 . There is a possible in vertebrates and to order in ( ) corresponding disparity in the number of and arthropods, save for gryllotalpids and species specimens from different months 1 . Conse- blattids, which were distinguished from ( Fig. ) conclusions other orthopterans. The numbers of each quently, concerning activity or of a are kind of prey in a given stomach were reproduction given species biased the few data from some noted, and the percentage volume of by each kind was estimated. months (January and December espe- if some females of Climatic data at Santa Cecilia were cially). Accordingly, a are in every obtained from a rain gauge in a clearing given species gravid and and a maximum-minimum thermometer month, except January December, months from which only one or two non- kept in a thatch-roofed building. Daily females are available, I have been readings were taken at about 0800 hr gravid inclined to state that the species breeds each day. Because of the richness and the On the other hand, complexity of the flora, no attempt was throughout year. the absence of females in July is made at a floristic analysis; the vegeta- gravid considered to be valid. tion was studied only in a structural context. Plant collections from Santa Ce- Although field studies were conducted cilia exist in the Shaw Herbarium of the throughout the year, the collecting of Missouri Botanical Garden in St. Louis tadpoles was sporadic. Consequently, 14 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
MAN DAYS SPECIES SPECIMENS xlO 300-
^200
100-
+ 4- +- +- 4- + -+- JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC
Fig. 1. —Comparative monthly rates of collecting effort and results at Santa Cecilia. data on occurrence of tadpoles was used river in the Oriente is the Rio Napo, only in a positive way—i.e., absence of which flows southeastward to join the data on tadpoles in a given habitat or at Rio Marafion near Iquitos, Peru, and a time of the year was not considered form the Rio Solimoes. The latter flows to be reliable. eastward to Manaus, Brasil, where it is joined by the Rio Negro to form the DESCRIPTION OF THE AREA great Rio Amazonas. One of the major tributaries of the The Amazon Basin, with an area of Rio Napo is the Rio Aquarico; the latter about 6,000,000 square kilometers, strad- is formed by the confluence of the Rio dles the equator and extends from the Cofanes and the Rio both of base of the Andes about 3000 kilometers Chingual, which drain the eastern face of the Andes eastward to the Atlantic Ocean. With in northern Ecuador. The confluence is the exception of some low hills and pla- at an elevation of 600 meters; from this teaus, the elevation diminishes gradually the Rio flows in from about 500 meters at the base of the point Aguarico rapidly a boulder-strewn bed to a below Andes to sea level at the mouth of the point Santa Cecilia (340 m). Throughout the Amazon. Although Ecuador is a small remaining two-thirds of the length of the country with a coastline on the Pacific river, it is broad, primarily silt-bottomed, Ocean and with the Andes bissecting the and to an elevation of about 200 country from north to south, approxi- drops meters at its confluence with the Rio mately one-third of the country lies in Amazon Basin (Fig. 2), a region referred Napo. to locally as the Oriente. The major Geologically the region of the upper EQUATORIAL HERPETOFAUNA 15
Fig. 2.—Map of Ecuador showing location of the upper Rio Aguarico area.
Rio Aguarico consists of deep Tertiary soils occur in extreme eastern Ecuador, alluvial deposits (2500-3000 m) above but apparently these either are absent or Cretaceous sandstones in the studied. The marine ( Tschopp, inconspicuous region 1953; Harrington, 1962). Following the beaches along the Rio Aguarico are of or river aver- teiTninology Beek and Bramao ( 1969), quartz sand pebbles. The two principal soil types occur in the age size of the latter varies from about region. Closest to the Andes and con- 30 cm near the base of the Andes to tinuing eastward in the well-drained about 10 cm at Santa Cecilia. areas are reddish brown laterites, where- Whereas the primary study area con- as red-yellow podzols occur in poorly sisted of about 3 square kilometers at drained areas removed from the Andes. Santa Cecilia, four other sites along the Pale yellow latosols and low humic gley Rio Aguarico were included as secondary 16 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
77°00' '76°45'
0°00'
Fig. 3.—Map of the upper Rio Aguarico showing location of study sites.
420 study areas; together these encompassed Puerto Ore.—00°03'N, 77°irW; an airhne distance of about 85 kilometers m. This was a temporary placer mining and an elevational range of 320-570 camp on the south bank of the Rio Agua-
3 . The five sites are de- rico. The area meters ( Fig. ) mostly supported primary scribed below: forest (1968). Dwreno.—00°02'S, 76°30'W; 320 m. Santa Cecilia.—00° 03'N; 76°59'W; 340 This small Cofan Indian village is on the m. Prior to 1965 this was a small Quec- south bank of the Rio Acjuarico. A few chua Indian village scattered along the hectares around the village were culti- north bank of the Rio Aguarico. In 1965, vated; elsewhere there was primary for- Texaco-Gulf established a petroleum ex- est (1967). ploration camp and built an airstrip. Lago Agrio.—00°03'N, 76°53'W; 340 When field studies were initiated in No- m. This is now the major petroleum cen- vember 1966 most of the area was cov- ter in the Oriente of Ecuador with a large ered by primary forest (Fig. 4). The airfield, refinery, and town of about 2500 exceptions were the clearings for the air- people. When field work was done there field, exploration camp, and the two-hec- the area was for- tare which ( 1969), virgin primary compound "Muiiozlandia", est. This site should not be confused with served as our base camp. Adjacent to a small lake by the same name about 3 the village of Santa Cecilia were some kilometers to the northwest. fields of pidtanos and ijuca. All human Puerto Libre.—00° 12'N, 77°29'W; 570 habitation and disturbance were between and the much smaller m. This is an intermittent placer mining the Rio Aguarico Rio 1-2 kilometers to the camp with a grass airstrip situated in a Conejo lying north. The Rio flows northward narrow valley at the base of the Andean Conejo of foothills. In 196(8 there was secondary into the Rio San Miguel, a tributary which is a river forest in the immediate vicinity of the the Rio Putumayo, large the Rio Solimoes. camp; elsewhere the area was covered by flowing eastward into in each primary forest. Beginning 1966 succeeding year EQUATORIAL HERPETOFAUNA 17
Fig. 4.—Aerial pliotograph of the Rio Aguarico encompassing the village of Santa Cecilia on the north bank of the river at the left edge of the picture and the airstrip (diagonal clearing in middle of picture). The darker irregular tones are forest; the pale areas are swamps domi- nated by Hdiconia. Photographed 2 February 1966 (courtesy of U.S. Air Force). brought more people into Santa Cecilia mation in the following sections pertains and more destruction of the forest, so only to the area of Santa Cecilia. that by 1971 many fanns of 50 or more hectares had been hacked out of the Physiography and Hydrography forest. In October 1971 the road between The terrain is flat with nu- Quito and Lago Agrio was completed; generally merous small streams and depressions; the oil companies moved to Lago Agrio, the total relief is about 20 meters. The and the Ecuadorian army took over the airstrip occupies a narrow ridge between oil camp (Fig. 5). With the completion the Rio Aguarico and Rio Conejo. From of the road, most of the families that had Mufiozlandia eastward the ridge gradu- been living on the river bank moved to ally diminishes to the Rio Aguarico, but the road; in so doing, they abandoned upstream there is a bluffs along the river. their homes and cultivated areas. Thus, The higher ground encompasses most of Santa as a no Cecilia, village longer the area between the Rio Conejo and the exists. of its former inhabitants Many Rio Aguarico west of the airstrip. now own 50-hectare homesteads along The Rio Aguarico is a "white water" the road. river with a width of about 100 meters Unless specified otherwise, the infor- at Santa Cecilia. According to Saul 18 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
>n.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 meters in width. In undisturbed areas 2.0 meters per second, and the surface the river is concealed by the forest. The temperature varies between 18° and 20° velocity is 0.3-0.6 meters per second, and C. Water level fluctuations of about 5 the surface temperatures are 22-23° C. meters were common and mostly asso- (Saul, 1975). Other streams in the area ciated with heavy rains on the Andean are small (< 3 meters wide) and shal- slopes. At times of high water, the size low. They vary in gradient from some of the island at Santa Cecilia was spring-fed rivulets that descend over rock markedly reduced, and local residents and gravel from the ridge to the Rio claimed to have seen the entire island Aguarico to silt-bottomed streams drain- covered with water. In contrast, at times ing lakes and swamps. of low silt water a broad, boulder and bar Two pennanent lakes are present in connects the of the island western end the area. The southern lake is the larg- with the north bank of the river (Fig. 6), est (± 150 X 300 m), whereas the resulting in an extensive backwater la- northern is about 100 m in diameter. on the north of goon side the island. Both lakes are deep; actual depths arc The Rio Conejo is a shallow (up to unknown but exceed 10 m. Originally 2 meters), meandering stream about 6 these eutrophic lakes were bordered by EQUATORIAL HERPETOFAUNA 19
M^ 20 xMISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Fig. 7.—The west shore of tlie lower lake at Santa Cecilia.
2 3 TEMPERATURE 600 X 35° < RAINFALL Q Z <
Z) -500 3 z 30' O o < >- _lX (-z o 25"- 400 z o z < LlI2 2
20'- --300
cr3 < < q: q: Ui Q. 2 200 —1 — —I 1— -+- JAN FEB MAR APR MAY JUN JUL AUG 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 Mean/Day Days Heavy Clear Maximum Minimum Jan 372.5 12.0 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 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, cerned more with the structure of the 165, and 4 mm, 3 days with no rain, and vegetation than with the floristics. then 86, 38, and 46 mm. Generally, rain Santa Cecilia is near the western lim- falls more frequently by day than at its of the largest expanse of tropical rain- night. Rain was recorded on 215 days forest in the world. This biotope called in the period July 1971-June 1972; of the Hijlea by most South American biol- these, rain fell during the day and night ogists is referred to locally as the selva. on 47 days and only at night on 40 oc- The region of Santa Cecilia is at a suffi- casions. Sometimes rainfall is extremely ciently high elevation to preclude the localized during brief but heavy showers, seasonal flooding of great expanses of whereas at times of extensive heavy forest. Thus, the hajiaJes of the lower cloud cover the entire region receives elevations of the Amazon Basin, char- rain. acteristic as far inland as Iquitos, Peru, are absent. Vegetation Prior to 1965 we can assume that, with Although plant collections have been the exception of a few clearings along the made for the Shaw Herbarium of the river and scattered swamps, the entire Missouri Botanical Garden in St. Louis region was covered with primary rain- and the herbarium at the Universidad forest (Fig. 9-10). This forest is best Catolica in Quito, no published summary developed on level, well-drained ground. of the flora is available. Because the In such places the largest trees form a physiognomy determines the numbers canopy 30-35 meters above the ground. and kinds of habitats available for ani- Some stilt palms protrude through the mals, the following commentary is con- 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. shallow, extensive root systems. Growth Most such trees have extensive but- seems to be limited by the ability of the tresses, that may extend 3 meters up the shallow soil to hold the towering trees. which be to 2 meters in trunks, may up These forest giants frequently fall over, diameter In some areas (Figs. 11-12). with the uprooting of the extensive root definite stratification can be found. A system creating broad shallow craters secondary stratum, when present, is 15- and the interwoven system of lianas tear- 20 meters above the ground. This usually ing a large swath in the forest. Thus, 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- but- liads are uncommon. The large, Fig. 11. —Buttressed base of large tree in pri- tressed trees forming the canopy have 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
marsh. Fig. 14.—A forest-edge situation showing dense shrubby growth; in the foreground is a
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 are organized by major types of habitat. rains to large semi-permanent swamps These are defined as follows: (Fig. 16). The swamps in primary forest Primary Forest.—Mature forest char- characteristically have numerous spiny acterized by nearly continuous canopy, palms, whereas the swamps in secondary stratification of vegetation, and deep forest usually support dense growths of mulch layer. Heliconia, the immense leaves of which Secondary Forest. — Successional attain lengths of 2 meters, exclusive of stages and partially lumbered primary stems (Fig. 17). forest. The cutting of the large trees re- sults in a that Man-made clearings are either culti- physiognomy resembling of inteiTnediate successional vated or maintained as clearings by man stages. Forest —The or grazing animals. Some maiz and pine- Edge. ecotone, usually resulting from human disturbance, be- apple are grown, but pldtanos and ijtica tween forest and clearings. are the most common crops. As a result Bamboo.—Any of the stands of bam- of topographic irregularities, there are boo in forest or cleared areas. low-lying areas in clearings that contain Clearing.—Uncultivated clearings, water. The marshes in these open de- usually man-made and characteristically pressions contatn sawgrass, small Heli- supporting a variety of grasses. conia, and various aroids (Fig. 18). Ctdtivated Fields.—Any cleared areas Habitats bearing crops. Swamp.—Forest depressions contain- Throughout the accounts of the species ing water either ephemerally or perma- and in the ecological analysis, the data 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 29
Marsh Water-filled depressions in ways (Rio Aguarico and Rio Conejo) are clearings. included in this category. Lake.—This includes category only Stream.—All flowing water not in- the two lakes in the area. pennanent cluded in the fonner category. River. —Only the two major water-
IDENTIFICATION OF THE SPECIES
One of the most difficult problems Uno de los problemas mas dfficiles that faces biologists working with the que han enfrentado los biologos a traba- large Amazonian biota is the identifica- jar con la abundante biota amazonica tion of the species. The following keys ha sido la identificacion de las especies. are offered as an aid to the identification Las siguiente claves se entregan como of the species of amphibians and reptiles una ayuda para la identificacion de las from the upper Rio Aguarico. They especies de anfibios y reptiles de la re- should be useful for the herpetofauna of gion del alto Rio Aguarico. Ellas de- the entire upper Amazon Basin in Ecua- berian ser igualmente utiles para la her- dor, southern Colombia, and northern petofauna de toda la Hoya Amazonica Peru. Identifications made by use of the tanto en Ecuador, el sur de Colombia, keys should be checked against the de- como en norte de Peru. Las identifica- scriptions given in the accounts of the ciones hechas a traves de estas claves species. deberian ser confrontadas con las de- The keys have been designed for use scripciones dadas en los registros de es- in the field with living or freshly pre- pecies. served Unless stated other- specimens. Estas claves han sido disefiadas para colors to animals. All wise, pertain living su uso en el campo, trabajando con characters used in the are visible keys especimenes vivos o recientemente pre- with no more magnification than that servados. Si no menciona otra cosa, las a 10 hand lens. The provided by X keys colores corresponden a animales vivos. are satisfactory for specimens of all ages, Todos los caracteres usados en las claves that some will except juvenile frogs pre- son visibles con no mas aumento que el sent difficulties. dado por una lupa manual de lOX. Las All character the most states, except claves con satisfactorias para especi- obvious are illustrated. Internal ones, menes de toda edad, excepto para al- such as dentition and hemi- structures, gunos sapos juveniles que presentaran have been avoided. The most con- penes, un poco mas de dificultad. scutellation character in snakes is fusing Todos los caracteres senalados, ex- the loreal, a scale on the side of the head cepto los mas obvios, son ilustrados. Las between the nasal and the preocular estructuras internas, tales como denti- 41a). Some snakes lack either the (Fig. cion y hemipenes, han sido evitados. De loreal scale or the scale. If the preocular los caracteres de escutelacion el que mas scale that is is than present higher long, confunde en las culcbras es la escama it is the preocular (Fig. 41c), whereas, loreal, ubicada lateralmente en la cabeza if it is than wide," it is the loreal longer entre las escamas nasal y la preocular (Fig. 43b). (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). 48 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Key to the Turtles Clave de la Tortugas
1. Neck folds laterally to lie along mar- 1. Cuello plcgado lateralmente fuerca gin of shell 2 de la concha 2
Neck withdraws straight back into Cuello se retrac directamente dentro shell ^ 5 la concha 5
2. Head broad, flat, with fleshy probos- 2. Cabeza ancha, chata, con proboscis cis and many long barbels; carapace carnosa y muchas barbillas; cara- broad, depressed, with ridges of pacho ancho, deprimido, con las keeled scutes salientes de los escutelos quillados Chelus fimhriatus (p. 192) CJielus fim])riatns (p. 192)
Not as described 3 Diferente a lo descrito 3
3. Two rounded, longitudinal ridges on Dos crestas longitudinales redon- carapace; lateral edges of carapace deadas en el carapacho; margenes upturned laterales del carapacho dispuestos Platemys platycephah (p. 193) hacia arriba Platemys platycephala (p. 193) Not as described 4 Diferente a lo descrito 4
4. Margin of posterior plastral indenta- 4. Margcn de la identacion plastral tion concave (Fig. 27a); plastron posterior concava (Fig. 27a); plas- dark; throat dusky tron oscuro; region gular terrosa Mesoclemmys gibha (p. 192) Mesoclemmys gihJ)a (p. 192)
Margin of posterior plastral indenta- Margen de la identacion plastral tion convex convexa ( Fig. 27b ) ; plastron posterior (Fig. 27b); plas- pale with or without dark spots; tron palido con o sin manchas as- throat cream with black spots curas; region gular crema con man- Phrynops geoffroanus tuherosus chas negras (p. 193) Phrynops geoffroanus tuherosus (p. 193)
a \ / \ / b
Fic. 27.—Posterior margins of plastra: A. Mesoclemmys gibba, B. Phrynops geojfroaniis. 192 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
— Cat. Chelon. Syncope antenori Walker Cliclt/s finiJ)iiata Boulenger, Rhynchoceph. Crocod. Brit. Mus.:209. Syncope antenori Walker, 1973, Occas. Pap. Clielus jxmhnatus—Mertens, Miiller, and Rust, Mus. Nat. Hist. Univ. Kansas, 20:3 [Type 1934, Bl. Aqua.-Terr. Kunde, 45:65. locality.—Puerto Libre, Proxincia Napo, Materia].—Santa 3. Ecuador]. Cecilia, Identification.—This flattened turtle, Material.—Puerto 11. Libre, commonly called matamata, with a broad —This dull mi- Identification. minute, head and fleshy proboscis is unlike any crohylid frog differs from all other frogs other Amazonian turtle. The snout and in the area four toes and, by having only parietal region are orange-tan; the rest additionally, from all other microhylid of the head and neck are dull dark frogs by having a visible tympanum. brown. The limbs are dark brown with This small frog has a robust body and a tan stripe along the anterior edge of broadly truncate snout in dorsal view. the forelimb. The carapace is dark The distinct tympanum is about half of brown with a yellowish tan middorsal the diameter of the eye. The fingers bear stripe; the tips of the keels on costal and lateral fringes; the four toes (the nomial and marginal scutes are tan. The plas- first toe absent) have terminal discs and tron is dull tan. 6 330 mm; 9 345 mm. lack webbing. The dorsum is dull brown; Occurrence.—All of the specimens the venter is grayish brown with small were obtained from the Rio Aguarico by bluish white flecks. Males lack nuptial Indians. excrescences. S 12 mm; 9 14 mm. Life History.—No data are available. Occurrence.—Eight individuals were in primary forest and three in secondary Mesoclemmys gibba (Schweigger) forest in All were found at - July. night Emys gihha Schweigger, 1812, Konigsberg. of low herbs and seven four on leaves Arch. —Naturgesch. Math., 1:299 [Type lo- on the ground. cality. unknown]. Mesoclemmj/s ^ihha—Gray, 1873, Ann. Mag. Life History.—Five females contained Nat. Hist., (4) 11:306. 5-6 (X=5.4) ovarian eggs 1.2 mm in Material.—Santa 5. diameter. The presence of few relatively Cecilia, —This side-necked tur- large ovarian eggs suggests that in this Identification. tle has a smooth, broad that is species eggs may be terrestrial and un- carapace It differs from dergo direct development. flared laterally. Vhrynops in a dark ( Fhry- Mating Call.—No call was associated by having plastron light nop.s) and from Platemys by having a with this species. The absence of vocal smooth carapace (two rounded longi- slits in adult males perhaps is indicative tudinal ridges in Platemys). Further- that this small frog is mute. more, Mesoclemmys has a shallow round- Food.—Examination of seven stom- ed indentation in the posterior margin of achs revealed the presence of ants and the carapace, whereas Phrynops has a mites in six; one of these also contained deep angular indentation. The carapace a small beetle. The seventh individual and plastron are dark brown to black, contained a beetle, a and a diplopod, and the head and neck are brown. 9 pseudoscorpion. 214 mm. Occurrence.—Two were in TURTLES brought by natives, and two shells were found by CHELmAE a forest pond. Another was in a marsh Chelus fimbriatus (Schneider) in a clearing. Life History.—No data are available. Tcsiudo fiml}riata Schneider, 1783, Allgem. Remarks.—The two recently killed Naturgescli. Schildkr.:349 [Type locality. — found a in Surinam]. females were by pond primary EQUATORIAL HERPETOFAUNA 193
— 8. forest in April; only shells remained. Material. Santa Cecilia, Conceivably these individuals were seek- Identification.—This brightly colored turtle is from other ing oviposition sites when they were at- readily distinguished tacked by predators. The proximity of side-necked turtles in the area by its the turtles to the large permanent pond elongate carapace having two rounded suggests that this species may inhabit longitudinal ridges, bordering a median such ponds and lakes. depression. The shells of MesocJemmijs and Plirynops lack ridges, and CheJiis has three keeled The Phrynops geoffroanus tuberosus (Peters) ridges. carapace, plastron, and skin are dark brownish Platcmtis tiihcrosa Peters, 1870, Sber. Akad. black with an to Wiss. Berlin, 1870:311 [Type locality.— orange-tan periphery Cotinga River, Mt. Roraima, Guyana]. the shell and broad orange stripes on the tiihcrosa—Miillcr, 1939, Phninops pcoffroana limbs and dorsum of the head, c? 161 16:95. Physis, mm; 9 156 mm. Material.—Puerto Ore, 1; Santa Ce- Occurrence.—A juvenile having a cilia, 1. carapace length of 55 mm was in a patch Identification. —Plirynops ^cojfroanus of sunlight in a shallow rivulet in pri- has a broad, smooth carapace that is mary forest, and one adult was crawling flared peripherally. The carapace has a along a trail by day. One adult was in deep angular indentation in the posterior shallow water in the Rio Aguarico by margin. The carapace is black. The day, and two adults were lying on the plastron and ventral flange of the cara- bottom of shallow swamps at night. pace are creamy yellow to orange with Life History.—No data are available. black spots. The limbs are dull gray KiNOSTERNIDAE with pale red to creamy white spots. The head and neck are dull olive green with Kinosternon scorpioides (Linnaeus) fine reticulations and a black line black Testudo scorpioides Linnaeus, 1766, Systema from the snout through the orbit and Naturae, Ed. 12, 1:352 [Type locality.— onto the neck. The throat is pale tan Surinam]. Kinosternon scorpioides—Gray, 1831, Synops. with irregular black markings. A pair Rept., 1:34. of white chin barbels are long creamy Cinosternon scorpioides scorpioides—Sieben- present. The only other side-necked tin-- rock, 1907. Sher. Akad. Wiss. Wien, 116: tle in the area with a smooth carapace is 576. Mesoclemmys gihha which has a black Material.—Santa Cecilia, 9. — turtle with plastron with a shallow rounded indenta- Identification. This small tion in the posterior margin. 9 316 mm. a high carapace with three low, longitu- Occurrence.—Both specimens were dinal keels withdraws the neck into the obtained from the Rio Aguarico by na- shell. Other species with longitudinal or keels on the CheJus tives. ridges carapace ( Life Histori/. —No data are available and Platemys) have low, flattened shells, from Santa Cecilia. Medem (1969:334) and fold the neck laterally under the other turtle reported nests containing 10, 11, and 15 margin of the shell. The e22:s found in March and November in with a high carapace is Geochelone, Caqueta, Colombia. which has short limbs with unwebbed toes; the toes of Kinosternon are webbed. Platemys platycephala (Schneider) The carapace is dull grayish brown; the is dull brown. The Testiido platycephala Schneider, 1792, Schr. plastron yellowish with Ges. Naturf.— Freunde Berlin, 10:261 [Type top of the head is yellowish orange locality. "East Indies"]. black flecks; the rest of the head, neck, Platcntys platycephala—Boulenger, 1889, Cat. and limbs are dull gray. 6 161 mm; Chelon. Rhynchoceph. Crocod. Brit. Mus.: 227. 9 155 mm.