GUNTHER KOHLER Reptiles of Central America|Impressum Cover photos: Front cover, left: Polychrus gutturosus (Rio San Juan, Nicaragua), Right top: Bothriechis aurifer (Guatamala). Photo: M. Burger Right bottom: Diploglossus monotropis. Photo: R. W. Van Devender Photo p. 4: Cloud forest (Cerro Kilambé, Jinotega, Nicaragua). Back cover: Rain forest (Saslaya National Park, Atlantico Norte, Nicaragua). Lampropeltis triangulum (Patuca National Park, Olancho, Honduras). Manin QL 656 Al KEY x Aoo3 Kéhler, Gunther Reptiles of Central America Offenbach: Herpeton Verlag, 2003 ISBN 3-936180-02-4 Editorial Board: AJ Gutman, West Hartford; J. R. McCranie, Miami; Peter Stafford, London; Harlan Walley, Dekalb; Larry David Wilson, Miami All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including. photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. No liability is assumed with respect to the use of information contained herein. © 2003 Herpeton, Verlag Elke Kéhler, Rohrstr. 22, D-63075 Offenbach, Germany Photos and drawings: Gunther Kéhler, if not otherwise indicated Layout; Elke Kohler, OffenbachReptiles of Central America Gunther KohlerGunther Kéhler Reptiles of Central America 644 Color photographs 103 Drawings and diagrams 210 MapsContents Foreword Introduction . oe Comments on the Usage of this Book... 10 Classification of the Reptiles of Central America 12 The Environment... ow 16 Geological History and Zoogeography of the Reptile Fauna ... 24 Crocodilians (Crocodylia) wo... 28 Turtles (Testudines) . 33 Chelydridae 34 Dermatemydidae 35 Emydidae 36 Kinosternidae 41 Testudinidae .. 46 Cheloniidae, Dermochelyida 47 Lizards (Sauria) «0.0.0... 54 Anguidae . 5D Eublepharidae 67 Gekkonidae 68 Gymnophthalmidae sone 80 Helodermatidae ...... Iguanidae Scincidae .. Teiidae Xantusiidae Xenosauridae Worm Lizards (Amphisbaenia) .. Snakes (Serpentes) Blind Snakes (Scolecophidia) Anomalepididae 0... Leptotyphlopidae ‘Typhlopidae .. Macrostomata ..... Boidae .... Loxocemidae . Tropidophiidae ...... Colubridae Elapidae Viperidae Table of Scalation Characters in Snakes . . sn Snakebite ~ Prevention and First Aid ...... Mimiery in Snakes .. Acknowledgments Literature Cited .. Glossary Inde ecsecse . 306 312 ww B58 . 361Foreword Foreword Middle America constitutes one of the biodiversity hotspots of the world—a region characterized by a high level of species richness and endemism. It is understand- able, thus, that this area, extending from the Isthmus of Tehuantepec in Mexico to the southern extent of the Panamanian isthmus, should have attracted considera- ble interest in this era of biodiversity decline. The realization among natural scientists that the earth’s biota is under attack by forces most humans would describe as involving simply the business of living has generated an unprecedented interest in the cataloguing of the extent of that biota before it disappears The herpetofauna of Middle America, too, is being studied by a multinational group of specialists, including herpetologists from the length and breadth of Mexico and Central America, the United States, and, in the case of the present work, Germany. This study has resulted in the publication within the last decade of a number of detailed faunal works by Julian C. Lee, James R. McCranie and me, and Jay M. Savage and summary works by Jonathan A. Campbell, Celina Duefias and co- workers, Roberto Ibaiiez and co-workers, and the author of the current work, Gunther Kéhler on various portions of Middle America, These works have formed the basis for the two predecessors of the current work, as well as this work itself. In the year 2000, Dr. Kéhler published the first volume of the Reptilien und Amphibien Mittel- amerikas, which dealt with the croco- dilians, turtles, and lizards of this region. He followed this book in 2001 with a second volume dealing with the snakes of Middle America. These two books, written in Dr. Kéhler’s native language, are now combined, updated, and translated into his adopted language, English. Although the book is entitled The Reptiles of Central America, it covers the same area as its pre- decessors. The presentation of this infor- mation in English should increase its appeal to a broader readership The two previous books dealt with 3 croco- dilians, 25 turtles (including six marine species), 215 lizards, three amphisbaeni- ans, and 268 snakes. The present. work increases the species number by fifteen lizards and eight snakes to a total of 537 species. This increase of 23 species has occurred, however, within the last two years! With the continuation of the herpe- tofaunal surveys currently underway throughout Middle America, the number of constituent species will continue to rise. As it stands now, however, the current total of 537 species is only 55 fewer than the 592 species listed on the Center for North American Herpetology website for all of the United States and Canada! Clearly, Middle America has a very rich herpeto- fauna. The present book continues the publishing tradition begun by Dr. Kohler in 1993 with a book on basilisks of publishing his major works under the banner of Herpeton Verlag Elke Kéhler, the publishing com- pany operated by his spouse. In the last decade, Dr. Kéhler has produced, some- times alone, sometimes with coauthors, an impressive series of colorful books of both scholarly and popular interest on subjects as broad-ranging as reptile egg incubation, systematics of spiny lizards (Sceloporus) and spiny-tailed iguanas (Ctenosaura), and the systematics and distribution of the members of the Nicaraguan herpetofauna. The Herpeton Verlag website, in fact, cur- rently lists ten books under Dr. Kéhler’s authorship, two of which are German and Spanish versions of a treatment of the bio-logy of Iguana iguana, the Green Iguana. Several of these books are directed toward herpetoculturists, whereas others target professional and amateur herpetologists. Gunther Kohler is the Head of the Department of Herpetology at the Research Institute Senckenberg in Frankfurt am Main, Germany, and Director of the Conservation Project Utila Iguana, Honduras. He is continuing a tra- dition that extends back to the days of Oskar Béttger and, more recently, Robert Mertens. Dr. Kohler is continuing and expanding on the interest Mertens had in the herpetofauna of Middle America, espe- cially that of E] Salvador. Currently, Dr. Kohler is carrying on several major pro- jects dealing with the herpetofauna of El Salvador (in collaboration with Eli Greenbaum and Milan Vesely), Honduras (in collaboration with James R. McCranie and me), and Nicaragua, and the biology and conservation of the Utila Spiny-tailed Iguana, Ctenosaura bakeri. Since 1995, when he was awarded his Ph.D. and began work at the Research Institute Sencken- berg, he has rapidly made a name for him- self as a major player in the studies on the herpetofauna of biodiverse Middle America. This assessment is evidenced by Fig. 1, Review Workshop for the Global Amphibian Workshop at La Selva Biological Station, Costa Rica, in November 2002. Foreword the twelve books, 100 popular articles, and 110 scientific papers on reptiles and amphibians he has produced to date. On a more personal note, I am pleased to call Gunther Kéhler both colleague and friend. Recently, we both had a chance to participate in the Mesoamerican Review Workshop for the IUCN-supported Global Amphibian Workshop, which occurred at the La Selva Biological Station in eastern Costa Rica, While there, in addition to par- ticipating in the very important work of determining the conservation status of the almost 700 species of amphibians occupy- ing Mexico and Central America, we had a chance to renew our friendship while ram- bling about in search of amphibians and reptiles to photograph on the station grounds. During the five days we spent at the station, the first time we had been in the field together, I became impressed with Dr. Kéhler’s enthusiasm, dedication, breadth of knowledge about the Middle American herpetofauna, and even his cre- dentials. He holds both Ph.D. and DVM degrees! It was also during this time at La Selva that we began to trace the outlines of a project that will lead to another book, this one on the amphibians and reptiles of the Honduran Bay Islands and Cayos Cochinos, with our mutual colleague James R. MeCranie. I look forward to this project with anticipation. The book for which this is the foreword is a masterful synthesis of information from not only the two books that preceded it, but also from the broad array of scientific liter- ature that appeared both before and after the publication of its predecessors, produe- ing a Literature Cited section of more than 1000 entries. The species lists and keys for identification have all been revised and updated and will prove useful to anyone who has a doubt about what herp is in hand. The text is accompanied by profes- sional quality color photographs of about 400 of the 537 species and expertly doneForeword line drawings of diagnostic features. In addition, there are many attractive habitat photos and generalized distribution maps that will broaden the reader’s understand- ing of the places these creatures live. The 6 and 1/2 by 9 inch size also is convenient for portage into the field. Thus, finally, after more than three decades, we have a fitting modern treatment of the herpeto- fauna of Middle America, which has been wanting since the classic work by James L Peters and his co-workers, covering only the squamate reptiles, appeared in 1970 and quickly grew out of date. Middle America is an important reservoir of herpetodiversity, much of which evolved in situ, and it de- serves the kind of attention Gunther Kohler has provided. This book will provide the basis for more extensive treatments of this region’s herpetofauna that are needed to demonstrate the vast biogeographic significance of this region and the need to protect it for perpetuity. This foreword was written while I was on a research leave at the University of Miami, which leave was supported by NIGMS (National Institute of General Medical Science) Grant No. 1R25GM0083-02 to Dr. Michael S. Gaines, chairperson of the Department of Biology. Larry David Wilson Miami, Florida 12 January 2003Introduction The present book, which is based on KOHLER (2000a, 2001a), attempts to pro- vide a summary of the widely scattered information on Central American reptiles. The region that is covered by this book includes the seven countries that consti- tute political Central America (Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, and Panama), as well as the southeastern part of Mexico (the area east of the Isthmus of Tehuantepec, including the Yucatan Peninsula; Fig. 3). This part of Mexico belongs to the zoogeographically impor- tant Central American landbridge (Savace, 1982). Although the Swan Islands belong politically to Honduras, they are not inclu- ded because they are biogeographically part of the West Indies, instead of Central America. The history of herpetological research in Central America dates back to the early nineteenth century. Early milestones in the history of Central American herpetolo- gy were the “Herpetologia Mexicana” of Arend F. A. WIEGMANN (1834), the “Biologia Centrali-Americani” of Albert C. L. G. GUNTHER (1885-1902) and the “Mission Scientifique au Mexique et dans VAmérique Centrale” of DUMeRIL, Bocourr & Mocquarp (1870-1909). Early milestones of Central American snake literature include BOULENGER’s Cata- logues on the turtles, lizards and snakes in the British Museum (BOULENGER 1885, 1889, 1893, 1894, 1896a). In 1970, important works such as the two volume “Catalogue of Neotropical Squamata” of James A. Prrers and co-workers appeared. In the interval of about 30 years since these two publications, a large body of papers dealing with the various aspects of Central American reptiles has been published. This body includes works on certain genera or larger groups of turtles Introduction Caribhean Sea Belize Costa Rica Fig. 3. The zoogeographic region of Central Ameriea (shaded), (e.g. Ernst & Barpour 1989, Iverson 1992), lizards (e.g. UzzELL 1965a, 1965b, EcHTERNACHT 1971, OrreDAL 1974, Harris & Kiuce 1984, Harris 1994), snakes (e.g. DUELLMAN 1958, Downs 1967, Myers 1974, CAMPBELL & LAMAR 1989, CAMPBELL & Bropte 1992, Roze 1996, Savack & SLOWINSKI 1996), Additionally, a number of regional checklists and compilatory works (Meyer & WILSON 1973, H. SmirH & SMITH 1973, 1976, 1977, 1979, Savack & VILLA 1986, CAMPBELL & VANNINI 1989, KOHLER 1999c, 2001b, PEREZ SANTOS 1999), as well as thorough monographs of selected coun- tries, have been published (TayLoR 1951, 1954, Lee 1996, CAMPBELL 1998a, SrarrorD & MEYER 2000, SAVAGE 2002) Finally, in 1999, a masterful taxonomic reference for the snakes of the world (Colubridae not treated in the first volu- me) was published (McDIARMID et al. 1999). I generally follow the taxonomic arrangement proposed in that work. Central America is the link between North and South America and is of great zoogeo- graphic importance because its fauna con-10 Introduction tains species from both continents, in addi- tion to a high number of endemic species. [As presently understood, the reptile fauna of Central America consists of 533 species representing 163 genera and 29 families (Table 1). This enormous diversity is due to the exceptional vegetational variation found in the region, which makes it a mosaic of rain forests, pine forests, cloud forests, dry forests, cactus forests, savan- nas, swamps, and mangroves. This diversi- ty, and the fact that some areas have so far not been scientifically explored, renders Central America a great challenge for her- petological research. Much research remains to be done on the diversity, systematics, and ecology of Central American reptiles. The recent discovery of new taxa has even necessitated the description of new genera (CAMPBELL & SmirH 1998, KOHLER et al, 2001b). However, both scientific investigation and efforts to preserve biodiversity are in a race against time as the pace of habitat destruction increases in Central America Group Ped Crocodiles - Turtles iC Lizards 10 Amphisbaenians 1 Snakes 9 Total 29 Comments on the Usage of this Book ‘The intent of this work is to provide an identification guide for both scientists and travelers to the numerous reptile species found in Central America. Many Central American species of reptiles are so distinc- tive in coloration, as well as body size and shape, that an animal in question often can readily be identified by photographs or drawings alone. If the identification remains questionable by this method, identification keys should be consulted. Additionally, the distribution maps can be included in the identification process, because many species have very restricted distributions. The identification keys included in the present book consider all families, genera, and species that are known to occur in Central America, even though this is not explicitly stated in the respective key hea- dings. Subspecies are mentioned in the species accounts, but are not considered in the identification keys or in the distribution maps. The keys are dichotomous, meaning that they provide pairs of contrasting alternatives, Such a key works using step by step elimination of alternatives until eed ee 2 3 14 25 50 : 230 1 3 86 276 153 537 ‘Table 1. Taxonomic composition of the reptile fauna of Central America.only a single alternative remains. It is important to remember when using the identification key that the juveniles of many lizard and snake species have dif- ferent coloration and pattern than the adults. Any key based on distinguishing coloration will not be able to definitively identify the juveniles. The technical terms and morphological character designations used in the keys can be frustrating to non- specialists, but I am optimistic that anyone willing to invest some effort in learning the terms will be able to use the keys with con- fidence and success. The glossary included in the book will be helpful in this regard, as will the illustrations accompanying the keys. Following the introduction to each major group of Central American reptiles, the fol- lowing information is provided in the spe- cies accounts: the current scientific name with author and year of d ‘iption, follow- Introduction ed by the citation of the original deserip- tion and type locality; the maximum size (carapace length, total length [TL] or snout-vent length [SVL]), and the known geographic distribution. For some species, additional notes on ecology and reproduc- tion are included. Detailed anatomical descriptions are beyond the scope of the book. The recommendations for further reading at the end of each generic account serve the reader as a guide to the addition- al references. The species accounts are sha- ded with gray for easy recognition. The index at the end of the book will help in finding the pages where the keys, species accounts, photographs (bold page num- bers), and distribution maps (italic page numbers) of the respective species are loca- ted. The keys and maps included in this book are based mostly on the information provided in the publications listed at the end of each generic account supplemented by my own unpublished data. Fig. 4. Bothriechis schlegelii (Bartola, Rio San Juan, Nicaragua), 11Classification of the Reptiles of Central America Classification of the Reptiles of Central America _ Chelydridae fe idae = Emydidne ae Chelydra (1) Dermatemys (1) Rhinoclemmys (6) Claudius (1) Terrapene (1) Kinosternon (6) Trachemys (1) Staurotypus (2) Caretta (1) Dermochelys (1) Chelonia (1) Eretmochelys (1) Lepidochelys (2) ‘The taxonomic levels: Hl Class © Family Mi Order Genus (in parentheses is the number of species P Pi Bi Suborder that occur in Central America) 12Anguidac Abronia (18) Celestus (8) Coloptychon (1) Diploglossus (3) Gerrhonotus (1) Mesaspis (2) Anolis (1) Basiliscus (4) Corytophanes (3) Ctenonotus (1) Ctenosaura (12) Daetytoa (7) Enyalioides (1) Iguana (1) Norops (72) Laemanctus (2) Morunasaurus (1) Phrynosoma (1) Polychrus (1) Sceloporus (18) Urosaurus (1) Classification of the Reptiles of Central America ‘Eublepharidae Gekkonidac Coleonyx (2) Aristelliger (1) Anadia (2) Gehyra (1) Bachia (2) Gonatodes (1) Echinosaura (2) Hemidactylus (5) Gymnophthalmus (1) Lepidoblepharis (2) Leposoma (2) Lepidodactylus (1) Neusticurus (1) Phyllodactylus (3) Prionodactylus (1) Sphaerodactylus (9) Ptychoglossus (3) Thecadactylus (1) Eumeces (1) ‘Ameiva (6) Lepidophyma (1) Mabuya (1) Aspidoscelis (6) Mesoscincus (2) Cnemidophorus (1) Scineella (1) Sphenomorphus (4) Xenosaurus (1) Heloderma (1) ORE. Balinyer 1314 Classification of the Reptiles of Central America Anomalepis (1) Leptotyphlops (2) Ramphotyphiops (1) 4 Helminthophis (1) Typhlops (4) | Liotyphlops (1) | jo Aghistrodon ( Atropoides (2) Bothriechis (9) Bothrops (2) Cerrophidion (2) Crotalus (1) Lachesis (2) Porthidium (6) Micrurus (16) i venomous! __ [ Macrostomata % a e Boa (1) Loxocemus (1) Trachyboa (1) Corallus (2) Ungaliophis (2) Epierates (1)Classification of the Reptiles of Central America Se Dianna eC Nateetaae CREED", ic @ ‘Nerodia (1) Iphicos (6) Chironius (3) mniop: Coluber (1) Dipsas (8) Storeria (1) Amastridium (2) Dendrophidion (4) Imantodes (5) ‘Thamnophis (4) Atraetus (5) Dryadophis (3) Leptodeira (5) Chapinophis (1) Drymarchon (1) Ninia (8) Clelia (3) Drymobius (4) Pliocereus (2) Erythrolamprus (2) Elaphe (1) Rhadinaea (20) Geophis (18) Ficimia (2) Sibon (8) Hydromorphus (2) Lampropeltis (1) ‘Trimetopon (6) Liophis (2) Leptodrymus (1) ‘Tropidodipsas (3) Manolepis (1) Leptophis (7) Urotheca (5) Conophis (2) Omoadiphas (1) Masticophis (1) Crisantophis (1) Oxyrhopus (1) Onybelis (4) Diaphorolepis (1) Phimophis (1) Pituophis (1) Enutiophis (1) Peeudoboa (1) Peeustes (1) Enulius (3) Siphlophis (1) Salvadora (1) Nothopsis (1) Tretanorhinus (2) Senticolis (1) Rhinobothryum (1) ‘Tripanurgos (1) ‘Symphimus (2) Scolecophis (1) Xenodon (1) ‘Seaphiodontophis (1) Spilotes (1) Stenorrhina (2) Tantilla (22) Tantillita (3) Trimorphodon (1) ‘The taxonomic levels: (Suborder |} Infraorder @ Family i Subfamily Genus (in parentheses is the number of species that occur in Central America) 1516 The Environment The Environment The Environment A chain of roughly 100 volcanoes stretches along the Pacific coast of Central America from eastern Chiapas, Mexico, to central Panama. Of these, about 25 retain signs of activity. The volcanic peaks are highest in the north, where two in Guatemala reach elevations of more than 4000 m. The various mountain complexes that form the physiogeography of Central America can be divided into two major massifs. The northern block includes the Mesa Central of Chiapas, the highlands of central and southern Guatemala, and those of Honduras, El Salvador, and northern Nicaragua, ‘These are separated from the southern highlands by the Nicaraguan Depression. The southern highlands include the Cordilleras de Guanacaste, Central, and Talamanca. The two large Nicaraguan lakes (Lagos de Managua and Nicaragua) are the largest inland bodies of water in Central America, with, surface areas of 1035 km? and 8157 km2, respectively. The diverse landscape of Central America sup- ports very different regional climatic situations (DUELLMAN 1966, SruaRT 1966). On the Pacific versant of nuclear Central America the dry season extends from January through April. A subhumid corri- dor, made up predominantly of dry forest or the remains thereof, stretches along the Pacific versant of Central America from the Isthmus of Tehuantepec through northwestern Costa Rica. Generally, the Caribbean versant experiences an equa- ble, tropical climate with relatively mesic conditions throughout the — year. Exceptions are the northern part of the Yucatan Peninsula, as well as some interi- or valleys, which experience a more arid climate. The wettest regions of Central America are found in southeastern Nicaragua and northeastern Costa Rica, which receive an excess of 6000 mm preci- pitation per year. (Honduras), Sandy beaches Sandy beaches (Fig. 5) extend intermit- tently along much of both the Pacific and the Caribbean coast, especially in bays that are protected from the breakers by offshore reefs, Commonly, a narrow strip of coconut palms occurs along the beach. Dunes support the growth of scrubby xero- phytic vegetation, such as low shrubs, aga- ves, and matted cacti (Rhipsalis sp.). Among others, the heliothermie race run- ners (Cnemidophorus), spiny-tailed igu- anas (Ctenosaura), and spiny lizards (Sceloporus) inhabit the sandy beaches. These exposed sandy habitats are also important nesting sites for sea turtles and spiny-tailed iguanas (Ctenosaura). Mangroves Mangroves (Fig. 6) are found along the coastline at many localities in Central America. The red mangrove (Rhizophora mangle) is more salt water tolerant, and usually occurs immediately along the edges of lagoons and bays. Behind the stands of red mangrove, the much taller black mangrove (Avicennia germinans)Fig. 6. Mangrove with epiphytes on Isla de Utila, Honduras. and white mangrove (Laguneularia race- mosa) often form dense swampy forests. A tangle of aerial and adventitious roots characterizes this ecosystem. Mangrove swamps provide habitat for the American ‘ocodile (Crocodylus acutus), the Utila anole (Norops utilensis), and the Utila iguana (Ctenosaura bakeri). Dry forest Dry forests are typical of the Pacific ver- sant of Central America and a few interior valleys of Honduras and Guatemala, The climate is characterized by low precipita tion (500 to 1200 mm) that falls primarily during the rainy season from June through November. During the dry season, when precipitation is negligible, a majori- ty of the trees shed their leaves, permit- ting sunlight and wind to desiccate the ground layer. Only the large rivers contain water throughout the year, whereas smal- ler streams are intermittent or dry during the dry season. Plants and animais of this extreme habitat are well adapted to the The Environment prolonged drought periods. Dry forests consist of a thick ground cover of su lents and drought-resistant shrubs, Tree heights generally do not exceed 20 m, and many of the trees are armed with thorns, ‘A curious aspect of this formation is the presence of tree cacti, which often reach heights equal to that of trees (Fig. 7). Savanna The savanna that occurs in Central America includes wet savanna (e.g. on the Honduran island of Utila) and pine savan- na (“Sabanas de Pinos Caribe”). Pine Fig. 7. Dry forest at Tehuantepec, Mexico (400 m elevation) Fig. 8. Pine savanna at Alamikamba, Nicaragua (100 m elevation).18 The Environment savanna is found on the Caribbean ver- sant of Honduras (from the Laguna Ebano southeastward) and Nicaragua (from the Rfo Grande de Matagalpa northward), ‘This habitat is dominated by open stands of Caribbean pine (Pinus caribaea) that abruptly change over to broadleaf forest. Rain forest Evergreen tropical rain forest is among the most diverse ecosystems on the planet, and in many regions its scientific explora- tion is still in the beginning stages. In the northern parts of Central America, rain. forest is found mainly in the Caribbean lowlands. In Costa Rica and Panama, it also covers large areas of the Pacific ver- sant. During the day, temperatures in the tropical rain forest reach 30-35°C. During the night, temperatures drop to about 25°C. Relative humidity is very high (80- Fig. 9. Rain forest at Bartola, Rio San Juan Nicaragua (30 m elevation). Fig. 10, Buttress tree in the rain forest at Bar- tola, Rio San Juan, Nicaragua (30 m elevation) 100%). Rainfall tends to be considerable throughout the year, and heavy rains are most common in the early afternoon. Several levels of vegetation can be disting- uished in tropical rain forest, from the dimly lit forest floor, the scrub level, the trees of different sizes, up to the tallest trees of the forest, which reach high above the canopy. The diversity of tree species is extraordinary and can exceed 50-100 per Fig. 11. Cloud forest at Volean Mombacho, Nicaragua (1100 m elevation).The Environment Fig. 12. View of the cloud forest on Cerro Kilambé, Nicaragua. hectare. Many reptiles occur exclusively in primary lowland and premontane rain forests and disappear with deforestation. Cloud forest A world of its own, cloud forest is the type of natural vegetation that can be found above 1200 m elevation in many of the mountain ranges of Central America. In this type of habitat, temperatures are rela- tively low (5-15 °C, depending on altitude) and humidity is constantly high. The trees in cloud forest are densely covered with epiphytes, such as bromeliads, orchids, ferns, and mosses, Tree ferns, which can reach several meters in height, are also found frequently. Small streams channel the water on its way to the lowlands. At very high altitudes along exposed ridges, a special sub-type of cloud forest grows, the so-called elfin forest. Subject to low tem- peratures, high precipitation, and high wind speeds, trees in the elfin forest usually attain less than 1-3 m in height. The fauna of cloud forests tends to be less diverse than that of the lowlands, but there is a high number of endemic species, cies that only occur in one particular area, such as a summit or a mountain chain 13. A cloud forest trail at Monteverde, Costa Rica (1500 m elevation). 19The Environment ssssces::=c:seencueecemes seer Elevation Map ‘BHSqEST BIO[[FPIOD w oooe< fl Ww 0002-000T ; | w000T-0 | neaqeld wepemsyeny, i sederyg op renuag esoyy \ a 20The Environment Map of Vegetational Zones sexy pnor fi (quessaa o1y1oeg) 1SeI0] UeI UaareAg x Quesiaa weaqqied) 3010} ures uosaszong A qso10j urea jouoseog [I aaousueyy | | durems ‘seysrepy a wuuvacs outa fl qsounj yuooug ysoroy Sag 2122 The Environment === Climate in Central America The climate in Central America is high- ly location dependent, with the yearly cycle largely characterized by variations in precipitation levels. On the Caribbean side, the heaviest rains fall during December and January, while the Pacific side receives virtually no rain during the northern winter (the dry sea- son on the Pacific side lasts from January until April). In general, the Caribbean side of Central America is more humid and receives more precipi- tation than the Pacific side. The following climate diagrams illus- trate the yearly climate in selected areas. The mean monthly precipitation is indicated by bars. The mean daily temperature can be read from the upper curve and the mean nightly temperature from the lower curve (data from MULLER 1996, except for Utila, which was collec- ted personally). Precitatin ew) Temperate CC) \ Guatemala City “7 San Salvador Managua” Precipitation (mn) Lo 4 JFMAMJJASOND ae Belize City tila Tegucigalpa Christobal Terparaure °C)(oy ames a oR 8 8 & & - a 3 8 8 = 2 ee The Environment gucigalpa, Hondui € a 8 8 & 2 2 «2 eo ea 8 8 2 2 © eo24 Geological History Geological History and Zoogeography of the Reptile Fauna of Central America Savaci (1966, 1982, 2002) provided masterful analyses and discussions on the composition, origins, and history of the Central American herpetofauna and developed a theory explaining the present distribution pattern of the amphibians and reptiles. Earlier biogeographers have regarded Central American as a faunal transition zone between the American con- tinents (e.g. SIMPSON 1950, K. ScHMDr 1954, DARLINGTON 1957, STUART 1964). However, Savack (1966, 1982, 2002) demonstrated that the Central American herpetofauna forms a distinct zoogeo- graphic entity, both in regard to its history and present composition. The following discussion is based largely on the latter three publications, supplemented with information from works on the geology of the region by Maurair & DINKELMAN (1972) and MARSHALL et al. (1979). The Central American herpetofauna is composed of genera with an origin in the region (autochthonous genera), as well as those that dispersed from North and South America into Central America. Savace (1966, 1982) concluded that three major and one minor historical source units had contributed to the Central American herpetofauna. ‘The most important historical unit is the “Middle American Element” (SAVAGE 1966, 1982) that is made up of genera that are primarily tropical Mesoamerican in distri- bution (many Central American endemics) and have their closest relatives either in Central or South America. The ancestors of this historical unit usually evolved in the region itself or in South America, and reached far into North America during the early Tertiary period when a tropical mesic climate with warm temperatures prevailed, When the climate generally became cooler and dryer during the late Cenozoic era, they became restricted to tropical Mesoamerica. Examples of the “Middle American Element” include the Kinosternidae, Helodermatidae, Xantusi- idae, and Xenosauridae. These families are all known as fossils from early Tertiary deposits in North America near 40° latitude north (SavaGr 1966) A second major historical unit is the “Old Northern Element” (SavaGE 1966, 1982), which is composed of genera with a pri- marily extratropical distribution in Eurasia and/or North America. During the early Tertiary period, this group's ances- tors experienced a more or less continuous and circumpolar distribution on the northern landmasses. The gradual clima- tic changes during the Cenozoie era forced this group's ancestors southward and caused their fragmentation into geograph- ic isolates. As a result, this stock contains a unique Mesoamerican component of endemic taxa that have evolved with the autochthonous “Middle American Element” from the Eocene epoch onward. A third major historical unit is the “South American Element” (SavaGr 1966, 1982), which is composed of genera with primari- ly South American distributions and rela- tionships. Their ancestors evolved in South America and migrated various distances into Central America after the closure of the Panamanian portal during the Pliocene. Consequently, this unit is a rather recent contributor to the current Central American herpetofaunal diversity. Finally, a fourth less important historical unit, the “Young Northern Element” (Savack 1966, 1982) can be differentiated. This unit is represented by a few genera (including Sceloporus and Urosaurus) witha primarily extratropical distribution associated with semiarid to arid regions in southwestern North America. A brief summary of the geological history of Central America facilitates the under- standing of the current distribution pat- terns of reptiles in the region. The land connection that existed between the American continents during the late Cretaceous and early Tertiary periods (70 to 50 million years ago) broke up in the Eocene epoch (54 to 35 million years ago) due to inundation of the Isthmian Link (= lower Central America). This so-called Panamanian portal separated Nuclear Central America from South America during a period of time that lasted for about 45 million years. It was not before the early Pliocene (some 5 million years ago) that the closing of the Panamanian portal reconnected the American conti- nents. The uplift of the Central American mountain massifs began in the Pliocene and continues today. The major physiogeo- graphic changes caused significant clima- tic shifts. During the Pliocene, the climate became generally cooler and more arid. Additionally, the Pleistocene glacials and interglacials were extremely important events. In the early Tertiary period (e.g. during the Paleocene epoch, 65 to 54 million years ago), the North and South American conti- nents were connected by a land bridge with low relief that supported a general- ized tropical reptile fauna that extended northward to at least 40° latitude. It can be assumed that all families of reptiles that currently inhabit the Neotropics were already present at that time. During the Eocene, flooding of the southern parts of the intercontinental land bridge (from what is now central Nicaragua to north- ern Colombia) eliminated the connection of North and South America. Soon, two groups were present in the northern re- gion: the “Middle American Element” and a subunit of the Old Northern Element, the so-called “Central American Complex” Geological History (Savacr 1966, 1982), The latter group reached Middle America in the Eocene through migration southward along the developing Rocky Mountains and became isolated from their congeners in the eastern and western areas of what is now the United States by the gradual develop- ment of a more arid climate across north- ern Mexico, In the Oligocene epoch, condi- tions remained similar to those of the Eocene in Nuclear Central America, al- though some uplift of the northern moun- tain massifs may have occurred The Miocene epoch (25 to 5 million years ago) was the time of greatest change during the Tertiary period in Central America, both in regard to climate and vegetation, as well as changes to the land relief. In the Miocene, with the disappear- ance of the Panamanian portal, a contin- uous land connection between the conti- nents was re-established for the first time since the early Eocene, making faunal exchange possible. The South American influence is present- ly restricted mostly to lower Central America (i.e., the region of what is today Costa Rica and Panama). Only a few spe- cies of South American origin have reached northern Central America (e.g., Gymnoph thalmus speciosus), whereas most South American elements dispersed only into Panama (e.g., the genera Morunasaurus and Trachyboa) or Costa Rica (e.g., the genera Anadia, Dactyloa, and Neu- sticurus). Many species of Central or North American origin have migrated far into South America (e.g. the genera Crotalus, Iguana, Norops) The Pliocene (5 to 1.8 million years ago) was the most important period in the evo- lution of the Central American reptile fauna that led to the modern species com- position. At the beginning of this epoch, representatives of both historical units (Middle American Element and Central American Complex) were widely distrib- uted over much of the Central American26 Zoogeography of the Reptile Fauna == region, with speciation centers in the uplifting mountain massifs. The moun- tains formed a climatic divide that led to the differentiation of species communities adapted to the more arid conditions on the Pacific versant, as well as those associated with the more mesic habitats on the Caribbean versant. ‘These modern species dispersed along routes that were controlled by climatic and physiogeographic factors (Fig. 14). Mountains restricted the migration of low- land species and provided dispersal routes for highland species. Two types of barriers 0-1000 m 1000-2000 22000 m. dispersal routes mesic climate dispersal routes, arid climate = climatic barriers 11000-2000 m. 22000 m dispersal routes | Sy Ma highland species Fig, 14. Principal dispersal routes of amphibi- ans and reptiles in Contral America during the past two million years. After SAVAGE (1966) are effective for lowland species: highland massifs and climatically unsuitable areas. An example of the latter type of barrier is the xeric northern part of the Yucatén Peninsula that prevented mesic-adapted species from moving from the Petén forests into the region. An area of very complex lowland climatic filters occurs along the Pacific versant of Costa Rica and western Panama. From the late Pliocene onward, the overall picture of relief and climate in Central America resembled, for the most part, the present physiogeography, with the excep- tion of the marine embayment of Nicaragua and the still flooded northern part of the Yucatan Peninsula, both of which became more and more dry due to decreasing sea levels. During the glacial periods of the Pleistocene (1.8 million years ago to the present), the summits of the highest Central American mountains were cov- ered with glaciers. Based on recent studies on fossil pollen profiles (CoLiNvaux 1993, 1996), it is assumed that the average temperature in Central America dropped 6 to 8°C below the present level during glacial periods, resulting in environments in the lowlands most closely resembling those of midelevation forests (500 to 1600 m) at present (SAVAGE 2002). During these periods the distributions of the highland species were compressed downward (by about 800 m), where they co-occurred with lowland species (Savace 2002). During interglacial periods, the highland species moved back up the cordilleras as tem- peratures returned to levels equivalent to those at present (SruaRT 1951, CAMPBELL 1984, SAVAGE 2002), It is assumed that the cooling and warming cycles in the latter part of the Cenozoic, in association with the continued uplift of the cordilleras, provided a major impetus for speciation (Savace 2002) During the turbulent geological history of Central America, several effects have con-Zoogeography of the Reptile Fauna Oligocene & Miocene Pliocene I Old Northern Element Generalized Tropical Fauna 1 Midate American Element IB South American Element i Central American Complex Fig. 15. Geological history of Central America since the Paleocene. After Savace (1966). tributed to the evolution of the region's reptile fauna (SAVAGE 1966): Peninsular effects A characteristic of peninsulas is that spe- cies can invade, but are hindered from further migration by the surrounding sea. Nuclear Central America formed a pen- insula during most of the Tertiary period and accumulated successive invaders from the north. The Panamanian portal served as an effective barrier, preventing these invaders from migrating southward. From the Eocene through the Pliocene epochs, this peninsula served as a center of spe- cies differentiation. Marine portal effects The formation of marine portals can frag- ment the former continuous distribution of a species, thus stimulating independent differentiation of the isolated populations. With the inundation of the Isthmian Link, species that were formerly continuously distributed in South and North America became isolated on the two sides of the Panamanian portal, with differentiation proceeding in Nuclear Central America independently from that in South America, Examples for such sister groups are the genera Ameiva / Cnemidophorus, Caiman ! Crocodylus, and Dactyloa / Norops. Insular effects Due to isolation, islands stimulate specia- tion, During the Tertiary period, particu- larly in the Miocene epoch (24-5 million years ago), the numerous islands associa- ted with the Isthmian Link / Panamanian portal zone of instability supported insu- lar evolution. Land bridge effects ‘The formation of land bridges makes pos- sible the invasion of species previously hindered from immigration hy the exist- ence of marine portals. After the re-estab- lishment of the Isthmian Link in the Pliocene, invasion from the south by spe- cies previously excluded from Central ‘America began and continues to the pres- ent. 27seprpos0.13, Crocodylia Crocodilians (Crocodylia) The order Crocodylia is represented in Central America by three species in two families. Typical of all species of the order, these are large, robust, semiaquatic rep- tiles. They inhabit lakes, lagoons, swamps, and large rivers. These strictly carni- vorous reptiles feed on invertebrates and vertebrates and exhibit necrophagy and even cannibalism (LEE 1996). Fig. 16, Adult Crocodylus acutus (Rio Tareol Costa Rica). Photo: A. Flaschendriger Fig. 17. Adult Morelet’s crocodile (Crocodylu Photo: R. Cedeno V. 28 In Central America, the American croco- dile (Crocodylus acutus) inhabits predom- inantly brackish coastal environments (lagoons, marshes, and mangrove swamps) with individuals penetrating further inland along major rivers, Morelet’s crocodile (Crocodylus moreletii) and the spectacled or brown caiman (Caiman crocodilus) live primarily in freshwater s, such as lakes, ponds, and rivers. Fig. Cano Spectacled caiman (Caiman crocodilus) at ‘egro, Costa Rica, Photo: H. Maass Fig. 19. Speetacled caiman (Caiman crocodilus) at Cano Negro, Costa Rica. Photo: H. MaassFig. 20, Juvenile Crocodylus acutus (Rio Patuea, Honduras). All crocodilians are oviparous and lay hard-shelled eggs (usually seasonally, and ingle clutch per year). The spectacled caiman and Morelet’s crocodile build mound-type nests made of dead vegetation, sticks, and mud for their eggs, whereas the American crocodile excavates a shallow hole in the ground. In Chiapas, Mexico, the egg-laying season of the American croc- odile is during the months of March, April, and May with individual nests also observed as early as February (ALVAREZ peL Toro 1974), The female digs a hole, approximately 70 cm in diameter and 20 to 50 cm deep. Preferably, ni are made on sandy embankments above the water- line or high tide mark. Crocodylus acutus produces 15 to 105 (usually 20 to 60) eggs per clutch that hatch after an incubation period of 85 to 111 days (KOHLER 1997). The mound-type nests of Caiman crocodilus can reach a height of one meter and a diameter of more than two meters (STATON & Dixon 1977). After oviposition, the fe- male closes the nest cavity and crawls over it several times to condense the material. However, air pockets prevail in the nest throughout the incubation period. Due to fermentation and decomposition of the plant material, constant humid tempera- tures of 32-34 °C are produced within the mound. Caiman crocodilus (egg-laying Crocodylia 21. Juvenile Spectacled caim aragua), n (Bartola, sason from carly August through early November; GRENARD 1991) produces clut- ches of 10 to 23 eggs that hatch after an incubation period of 88 to 104 days. In Crocodylus moreletii (egg-laying season from April through June), the clutch size varies from 20 to 45 eggs and incubation takes 78 to 98 days (ALVAREZ DEL Toro 1974, KOHLER 1997). As far as we know, all crocodilians exhibit parental care (GREER 1971, KUSHLAN & SIMON 1981). During the incubation peri- od, the female remains in the immediate 'y of the nest and attacks potential egg predators (MopHa 1967, Corr 1971), The female responds to vocalizations made by juveniles from within the nest by opening it and transporting the hatchlings to water (POOLEY 1977). In crocodilians, the incubation tempera- ture determines the sex of the hatchlings. In most species, high and low tem- peratures produce male hatchlings, where- as intermediate temperatures produce females (DEEMING & FERGUSON 1988, KOHLER 1997). In Central America, all three species are endangered due to overhunting and destruction of habitat. National laws to protect crocodilians exist in all countries, 29seyypoo0a 30 Crocodylia Furthermore, Crocodylus acutus and C. moreleiii are listed in Appendix 1, and Caiman crocodilus in Appendix 2 of the Convention on International Trade in Endangered Species (CITES) Fig. 23 Juvenile American crocodile (Crocodylus acutus) at Rio Patuca, Honduras, Alligatoridae Caiman crocodilus (Linnagus 1758), Systema Naturae, ed. 10: 206; type locality: unknown. Maximum total length 2 m, Southern Mexico along most of the coasts of Central America into South America, sea level to 200 m eleva- tion. Recently, Busack & Panpya (2001) demonstrated that no subspecies of C. croco- dilus should be recognized. The same authors raised the former subspecies yacare to full species status, Crocodylidae Crocodylus acutus Cuvier 1807, Ann. Mus. Hist. Nat., Paris, 10: 55; type locality: Santo Domingo. Maximum total length 4 m (rarely up to 6 m). From southern Florida, USA, across the Greater Antilles, as well as from Sinaloa, Mexico to South America (Colombia and Venezuela), sea level to 500 m elevation Crocodylus_moreletii DuMERT. & BIBRON 1851, Cat. Méthod. Coll. Rept: 28; type locali- ty: Lake Petén, Guatemala. Maximum total length 3 m (rarely up to 4 m), Tamaulipas, Mexico, across most of the Yucatan Peninsula to southern Belize, sea level to 500 m eleva- tionFig. 24. Crocodylus acutus submerged in a small stream (Rio Patuca, Olancho, Honduras). LG Caiman crocodilus Crocodylus acutus Crocodylia 31 jilians Cc32 Crocodylia a Caiman crocodilus Crocodylus acutus Fig. 26 Characteristics important in the identification of crocodylians. Drawings by M. Vesely. Key to crocodilians 1 a A transverse ridge just anterior to eyes on top of head (Fig. 26); no teeth of lower jaw visible when mouth closed . : Caiman crocodilus b No transverse ridge just anterior to eyes on top of head (Fig, 26); 4" tooth of lower jaw lies laterally in an open groove and is visible when mouth is closed .. 2 2 a Primary subcaudal whorls not interupted, by incomplete secondary whorls (Fig. 27a); shout relatively slender, about twice as long as wide ... _.Crocodylus acutus b Primary subcaudal whorls interupted by incomplete secondary whorls (Fig. 27b); snout relatively broad, length about 1.5 times width .......... Crocodylus moreletit Further Reading Wermurh & MBRTERNS 1961, New. 1971, BRAzaitis 1973, ALVAREZ DEL. TORO 1974; Ross & Ross 1974; BurretauT 1979, Ross 1987, DENSMORE & OWEN 1989, Ross & GARNETT 1989, GRENARD 1991, PEREZ-HIGAREDA et al, 1991, Ekwst et al, 199. b, Crocodylus moretetii Fig. 27. Ventral surface of tail. Incomplete secondary subcaudal whorls in Crocodylus moreletii shaded,~ Testudines Turtles (Testudines) nuchal plate bridge marginal plates axillar plate inguinal plate ©. moveable hinge — Fig. 28. Characteristies important in the identifi- cation of turtles a, Lateral view b. Dorsal view; ¢. Ventral view; moveable hinge M=marginal plate; Jeural plate; =vertebral plate. pectoral plate anal plate femoral plate abdominal plate ‘Turtles are represented by 25 species in 14 genera and 7 families in Central America, the kinosternids and emydids being the most speciose with 8 species each. Key to genera of turtles 1 a Forelimbs modified as oar-like flippers; Fig. 29. Plastrons. two or fewer claws on each foreleg; oceanic — After WermuTH & species Sea Turtles (see key on page 53) Mnxrsns (1961), b Forelimbs not modified as oarlike flip- pers; more than two claws on each foreleg; species living on land or in fresh or brackish water habitats ... 2 2 a Plastron with 11 or fewer plates .. 8 b Plastron with 12 plates 3a Plastron not reduced and not cruciform .. 4 b Plastron very reduced and cruciform (Fig. 29b-d) ae : (Fig. Chelydra —. Stawrotypus 4. ClaudiusTestudines: 4 a Plastron with two moveable hinges (Fig. 28¢ and 30), chin barbels present susseanees Kinosternon b Plastron rigid, without moveable hinges (Fig. 28a); chin barbels absent . Dermatemys (in part) Chin barbels absent .. Chelydra b Chin barbels present (Fig. 28¢) . 6 6 a Plastron rigid, without moveable hinges (Fig. 294); bridge connected to carapace by ligaments . Claudius b Plastron with moveable hinge; bridge connected vo carapace by an osseous suture Staurotypus a » 7 a Plastron with moveable hinge; plastron and carapace in ntact, bridge absent - Terrapene hout moveable hinge a8 8 a Hind limbs elephantine (columnar); cara- pace dome-shaped; without webbing bet- ween toes .. Geochelone b Hind limbs not: columnar: carapace rela- tively flat, only slightly convex; webbing between toes more or less developed ....... 9 b Plastron rigid, w’ bridge present ... 9 a Limbs unicolor dark gray; snout slightly upturned, somewhat tubular. Dermatemys (in part) b Limbs with some kind of pattern (mostly pale stripes); snout not as above 10 10a Pattern of carapace consists of yellow rings with dark center; hind feet with extensive webbing ... Trachemys b Pattern of carapace without yellow rings, or, if yellow rings are present, then hind h little or no webbing -Rhinoclemmys Fig. 30, All species of Kinosternon have a plas- tron with hinges that enable a complete occlu- sion of the shell (K. leucostomum). 34 Fig. 31. Chelydra serpentina (Rio San Juan, Nicaragua) Chelydridae Chelydra The snapping turtle (Chelydra serpentina) can reach 47 cm carapace length with a weight of 20 kg and should be handled with great care. It is able to deliver quick strikes with its powerful jaws. Snapping turtles inhabit ponds, lakes, and slow- moving rivers with muddy bottoms. Mostly nocturnal, these animals usually rest in shallow water with only the nostrils and eyes protruding above the surface during the day. At night, this turt- le walks slowly on the bottom of its watery habitat searching for food. Snapping turt- les feed on almost anything they can swal- low, particularly insects, shrimp, crabs, fish, and frogs, but also crocodile and turt- le hatchlings, snakes, birds, and small mammals, as well as plant matter such as algae and other water plants and fruits. The female digs a nest for her eggs (in April through June) on a sandy embank- ment above the waterline, preferably in aspot with some protection by low vegeta- tion. Twenty to 30 eggs (exceptionally up to 83 eggs in very large females) are laid per clutch, The eggs measure 23 to 33 mm (weight 7 to 15 g) and the juveniles (cara- pace length 24 to 31 mm) hatch after an incubation period of 55 to 125 days (ERNST & Barsour 1989). Chelydra serpentina (Lannazus _ 1758), Systema Naturae, ed. 10: 199; type locality’ “Galidis regionibus”. Carapace length to 47 cm, Widely distributed across southeastern US.A., Mexico, and Central America to Ecuador, sea level to 1160 m elevation. The subspecies Chelydra s. acutirostris PETERS 1862 (northern Honduras across lower Central America to northern South America) and Chelydra 8, rossignonii (BOCOURT 1868) (central Veracruz, Mexico, across the base of the Yueatan Peninsula to western Honduras) oceur in Central America. Further Reading Exwst & Ginuons 1988 ‘i Chelydra serpentina 1B Dermatemys mawii Dermatemydidae Dermatemydidae Dermatemys The family Dermatemydidae consists of a single genus with one species. Dermat- emys mawii is found in lagoons, rivers, and lakes from southeastern Mexico through Belize. The species also tolerates brackish water. These nocturnal turtles are highly adapted to their aquatic environment, and are capable of performing prolonged dives. On land, however, locomotion is difficult, as is lifting the head. Adult Central American river turtles feed almost exclusively on plant material (algae and other water plants and fruits that fall into the water). The egg-laying period lasts from April through September with the females burying their clutches of 6 to 20 eggs (57-70 x 30-34 mm) on sandy embankments near the shoreline. Dermatemys mawii Grav 1847, Proc. Zool. Soe. London 1847: 55; type locality: unknown. Carapace length to 65 em. From central Veracruz, Mexico, through Tabasco, northern Chiapas, Mexico, and northern Guatemala to Belize, sea level to 300 m elevation, Further Reading Iverson & MITTERMEIER 1980, Fig. 82. Central American river turtle (Dermatemys mawii) in Tabasco, Mexico, Photo: J. HaftEmydidae Emydidae Rhinoclemmys The species of Rhinoclemmys are distribu- ted across the tropics and subtropics of Mexico, Central America, and northern South America. Some species (¢.g., R. fune- rea) are highly aquatic, whereas the majority of Neotropical wood turtles have a semiaquatic or even terrestrial lifestyle. In the Biosphere Reserve, Bosawas, Nicaragua, I encountered several R. annu- lata within the rain forest, some 100 meters from the nearest body of water. On the other hand, I have recorded R. funerea in the same general area, but exclusively in rivers. There, it can frequently be seen basking on partially submerged logs. Neotropical wood turtles are highly herbi- yorous and only occasionally ingest in- sects, They often forage on land at night, feeding predominantly on broadleaf plants, grasses, and fruits. Most Central ‘American species produce small clutches with one or two eggs; only R. melanosterna and R. pulcherrima lay larger clutches (up to 5 eggs per clutch) (ERNST & BARBOUR 1989, KOHLER 1997). Rhinoclemmys annulata (GRAY 1860), Proc. Zool. Soc. London 1860: 231; type locality: Hsmeraldas, Ecuador. Carapace length to 200 mm. Eastern Honduras to northwestern South America, sea level to 920 m elevation. Rhinoclemmys areolata (DuMBRIL & BIBRON 1851), Cat. Méthod. Coll. Rept: 10; type loca~ lity: El Petén, Guatemala. Carapace length to 200 mm. Southern Veracruz, Mexico, to extre- me western Honduras, sea level to 300 m ele- vation. Rhinoclemmys funerea (Core 1876), J. Acad. Nat. Sci. Philadelphia (2) 8: 154; type locality: Puerto Limén, Costa Rica. Carapace length to 325 mm. Rio Coco (border Honduras / Nicaragua) to central Panama, sea level to 600 m elevation. Rhinoclemmys _melanosterna (GRAY 1861), Proe, Zool. Soe. London 1861: 205; type locali- ty: Cherunha, Gulf of Darién, Colombia. Carapace length to 290 mm. Southeastern Panama to northwestern Ecuador, sea level to 400 m elevation. Rhinoclemmys pulcherrima (Gray 1855), Cat. Shield Rept. Brit. Mus. 1: 25; type locality: Mexico, Carapace length to 200 mm. Sonora, Mexico, to Costa Rica, sea level to 1160 m ele- vation, The subspecies Rhinoclemmys p. pul- cherrima (Guerrero and Oaxaca, Mexico), R. p. incisa (BOCOURT 1868) (Oaxaca, Mexico, to northern Nicaragua), and R. p. manni DUNN 1930¢ (southern Nicaragua to Costa Rica) occur in Central America, Rhinoclemmys rubida (Cope 1870), Proc. Amer. Philos. Soc. 11: 148; type locality: Juchitan, Oaxaca, Mexico. Carapace length to 230 mm. Pacific coast of Mexico from Jalisco to western Chiapas, sea level to 750 m. Only the nominate subspecies occurs in Central America, Fig. 33. Rhinoclemmys rubida. Photo: R. Wicker = A - Fig. 34. Juvenile Rhinoclemmys funerea (Rio San Juan, Nicaragua),Emydidae | es Rhinoclemmys funerea oS I Rhinoclemmys melanosterna Fig. 35. Rhinoclemmys pulcherrima (Rio Patuca, Olancho, Honduras). \ iRhinoclemmys annulate Rhinoclemmys areolata \ £4 Wi Rhinoclemmys rubida Fig. 36. Rhinoclemmys areolata. Photo: R. D. Bartlett Fig. 87. Rhinoclemmys areolata. (Calakmul Biosphere Reserve, Campeche, Mexico). Photo: H. Bahena B. Fig. 38. Rhinoclemmys annulata is more terrestrial than other species of the genus. 3738 Emydidae Key to Rhinoclemmys 1 a Hind feet with extensive webbing 0... 2 b Hind feet with little or no webbing .....4 2 a Iris pale gray without. transverse dark bar; posterior surface of thigh with verti- cal orange bars .. Rhinoclemmys melanosterna b Iris with transverse dark bar; ipatiern of thigh not as above oe 3 3 a One or two red supratemporal stripes present on each side reaching snout; eara- pace with or without ocellated pattern vou: Rhinoclemmys pulcherrima b No red supratemporal stripes; carapace without ocellated pattern . = Rhinoclemmys funerea 4 a Dorsal surface of head with distinct large yellow marking (often horseshoe-shaped; Fig. 33); bridge darkly pigmented ... .. Rhinoclemmys ru' b Dorsal surface of head not as above; bridge pigmented or not ..... 5 5 a Dorsal surface of head uniformly dark brown; supratemporal stripe, if present, indistinct carapace with broad keel Rhinoclemmys annulata b One or two pale supratemporal stripes present on each side of head posterior to orbit; keel on carapace, if present, narrow .. Rhinoclemmys areolata Further Reading Ens? 1978, 1980a-c, 1981a-c, PEREZ-HIGAREDA MITH 1987. Fig. 39. Rhinoclemmys pulcherrima (near Portegolpe, Guanacaste, Costa Rica). Photo: H. Bringsoe Fig. 40. Terrapene carolina yueatana (Calakmul Biosphere Reserve, Campeche, Mexico). Photo; H. Bahena B, Terrapene Four species of the genus Terrapene are currently recognized, with only 7: carolina occurring in southern Mexico (ERNST & McBreen 1991). On the Yucatan Peninsula, this species inhabits wet savannas and pasture land, as well as thorn forest and evergreen rain forest, with a preference for open habitats (LEE 1996). Although box turtles exhibit a ter- restrial life style, individuals can be ob- served sitting in shallow water. Juveniles appear to have a predominantly carni- vorous diet, while adults are predominant- ly herbivorous. The carnivorous diet of box turtles includes snails, worms, insects, spi- ders, salamanders, and frogs. Copulation is preceded by an extensive courtship, which can be divided into three phases (EVANS 1953): (1) a cireling, biting, and shoving phase; (2) a preliminary mounting phase; and (3) a copulatory phase. The females produce clutches of 2 to 4 eggs that hatch after 50 to 70 days, depending on ambient temperature (KouLER 1997).Terrapene carolina (LINNA! Naturae, ed. 10: 198; type lo fe Carapace length to 200 mm. Widely distribu- ted over the central and eastern U.S.A. with isolated populations in Mexico (Tamaulipas, San Luis Potosi, Veracruz, and the Yucatén Peninsula), sea level to 100 m elevation. The subspecies 7 c. mexicana (GRAY 1849b) (Ta- maulipas to Veracruz, Mexico) and 7! c. yuca- tana (BOULENGER 1895) (Yucatan Peninsula) occur in the area covered by this book . Further Reading Mu.steap 1969 (revision of Terrapene); ERNST & McBreen 1991, H. Suri et al. 1996. C Terapone colna a Fig. 41, Terrapene carolina yucatana (Campeche, México). Photo: Emydidae Fig. 42. Juvenile Trachemys scripta (Alamikamba, Nicarague Trachemys ‘The slider turtle (Trachemys seripta) has the widest geographic distribution of any non-marine turtle species. Due to its popu- larity as a pet, it represents the most well known turtle species. At the same time, there is still considerable disagreement among scientific authorities regarding its taxonomy on both the generic and subspe- cific level. I follow Cari H. Ernst (in Gipnoxs 1990: pp. 57-67), who places this species in the genus Trachemys and recog- nizes the subspecies 7. s. venu (Caribbean versant of southern Mexi and Yucatan Peninsula), 7: s. grayi (Pacific versant of the Isthmus of Tehuantepec to western Guatemala), and 7. s. ornata (Honduras to Panama) in Central ‘America, The position of JOHN M. LEGLER (in GipBons 1990; pp. 82-105), who assigns the species to the genus Psewdemys, with the following Central American subspecies P. s, venusta (Caribbean versant from 3940 Emydidae southern Mexico through most of Central America), P. s. grayt (Pacific versant of the Isthmus of Tehuantepec to border of El Salvador), and P. s, emolli (Nicaraguan Jakes and adjacent Costa Rica), seems unjustified. According to the studies of SemDEL & SMITH (1986) and SEIDEL & JACKSON (1990), the generic name Trachemys is correct for scripta and a few more species from the Caribbean region. Recently, SEIDEL (2002) proposed to recog- nize some of the former subspecies of T. scripta as distinct species. According to his taxonomic arragement, two species of slider turtles are recognized in Central America: 7. venusta and T. emolli. However, the taxonomy of Central American slider turtles may be subject to further revisions in the near future. Therefore, I have adopted a conservative approach and have used the name T, serip- ta for the time being. Slider turtles are diurnal and exhibit a very aquatic lifestyle. They occupy most freshwater habitats, with a preference for quiet waters with soft bottoms, such as ponds, lakes, and rivers. This turtle can frequently be seen basking on partially submerged rocks, roots, and logs. Juveniles feed mostly on fish and aquatic invertebrates, such as insects, worms, and snails, The food preferences of these ani- mals change with age toward a more herb- ivorous diet, including algae and other water plants, Adults are virtually omni- vorous. Slider turtles are preyed upon by crocodilians and humans. During the egg-laying period, which is in the dry season (December through May), the females bury their clutches of 9 to 25 eggs in open places that are exposed to the sun. The female digs a hole of 10 to 15 cm depth that is usually somewhat wider at the bottom. A female can produce up to six clutches in one season. Juveniles (cara- pace length 30-33 mm) hatch after an incubation period of 65 to 85 days. Females reach sexual maturity at an age of five to seven years and a carapace Fig. 43. Adult female Slider turtle (Trachemys seripta) at Rio Patuca, Honduras, length of 200 to 240 mm (Mout & LEGLER 1971, Ernst & BarBour 1989, Voct 1990), Trachemys scripta (ScHOEPFF 1792), Hist. ‘Testud.: 16; type locality: unknown, Carapace length ‘to 60 em, Widely distributed across central and eastern U.S.A. and southward through Mexico and Central America to Colombia and Venezuela, sea level to 1000 m elevation. Sce text above for subspecies. Further Reading Mow & Li ER 1971, Gipsons 1990, SEIDEL 2002 IB Trachmys scriptaKinosternidae Claudius ‘The genus Claudius is monotypic; the sole species is C. angustatus, the chopontil or narrow-bridged musk turtle. Narrow- bridged musk turtles are characterized by a small plastron that is connected to the carapace by ligaments. These turtles are known for their pugnacious attitude, and should be handled with great care in order to avoid severe bites. Claudius angustatus is an aquatic species that principally in- habits areas with seasonally flooded grasslands and forages in these grass- lands during the flooding (FLORES-VILLELA & ZuG 1995). It is most active around dawn. Claudius angustatus is strictly carnivorous and preys on invertebrates and small vertebrates. During the rainy season, some individuals migrate consider- able distances and can then occasionally be encountered on land during the day- time (L&E 1996). After the floods, C. angus- tatus reproduces. The clutch size varies from 1 to 8 eggs, which hatch after an incubation period of 94 to 150 days depending on environmental conditions (Ernst & Barsour 1989, KOHLER 1997). The Narrow-bridged musk turtle is Fig. 44. Claudius angustatus (Tabasco, Mexico). Photo: J. Haft Kinosternidae reported to estivate sequestered in during the dry season and resume acti with the onset of the rainy season (ALVAREZ DEL TORO 1983) Claudius angustatus Core 1865, Proc. Acad. Nat. Sci. Philadelphia 17: 187; type locality: ‘Tabasco, Mexico. Carapace length to 165 mm, Central Veracruz, Mexico, across. northern Guatemala to Belize, sea level to 200 m ele- vation. Further Reading IVERSON & BERRY 1980. x 1B Claudius angustatus Fig. 45. Claudius angustatus (Rio Hondo, Quintana Roo, Mexico), _ Photo; H. Bahena B. 4142 Kinosternidae Kinosternon Mud turtles (genus Kinosternon; about 20 species) are small to medium-sized, pre- dominantly aquatic chelonians that are widely distributed over North, Central, and South America. They are character- ized by one or two moveable hinges in the plastron that enable complete closure of the shell. The Central American species of Mud turtles live aquatically, mostly in ponds, rivers, and marshes from sea level to slightly more than 1000 m elevation. Frequently, individuals can be encoun- tered on land when they wander from one body of water to another. On the morning of 20 April 1990, during a stay at Tortuguero, Costa Rica, I observed a spec- imen of Kinosternon leucostomum in a clearing within the rain forest. When a mud turtle is lifted, it usually retreats into its shell and closes the shell completely with the aid of the plastron hinges (Fig. 30). After a while, some individuals might stick out their heads and attempt to bite whoever holding it. This is often accom- panied by the excretion of a malodorous fluid from their cloacal bladders. In regions with a pronounced annual cli- matic cycle, mud turtles spend the dry sea- son estivating while buried in mud (eg., Fig. 46. Kinosternon acutum. Photo: J. Buskirk Fig. 47. Kinosternon leucostomum (Rio San Juan, Nicaragua). Fig, 48, Kinosternon ereaseri (Calakmul, Campeche, Mexico) Photo: R. Cedeno V. creaseri in the northern Yucatan Peninsula; Lee —_1996). Kinosternids feed mostly on aquatic inver- tebrates, such a , worms, and in- sects. Some species (e.g., K. leucostomum) also consume a large quantity of plant material (seeds, fruits, flowers, and leaves); however, whenever available, they also prey on small vertebrates (fishes, frogs, lizards, and snakes) including carrion During the reproductive season, the fe- male produces several clutches, which, depending on the species, contains one to ten eggs (KOHLER 1997). KinosternonKinosternidae Kinosternon acutum GRay 1831, Synops. Rept. Cataphracta 1: 34; type locality: un- known, Carapace length to 120 mm. Central Veracruz, Mexico, through — northern Guatemala to Belize, sea level to 200 m ele- vation. Kinosternon angustipons Lecter 1965, Univ. Kansas Publ, Mus. Nat. Hist. 15 (13): 617; type locality: Los Diamantes, Limon Province, Costa Rica. Carapace length to 120 mm, Southeastern Nicaragua to northern Panama, sea level to 260 m elevation. Kinosternon creaseri HARTWEG 1934, Oce. Pap. Mus, Zool., Univ. Michigan 277: 1; type locality: “one mile south of the Hacienda, Chichen Itza, Yucatan,” Mexico. Carapace length to 121 mm. Northern portion of Yucatan Peninsula, sea level to 100 m eleva- tion, Fig. 49. Kinosternon scorpioides (Gualaca, Chiriqui, Panama). Photo: M. Lundberg Kinosternon leucostomum DUMBRIL & BIBRON 1851, Cat. Méthod. Coll. Rept.: 17; type locali- ty: N. Orleans [in error]; Mexique; Rio Usumacinta, Carapace length to 174 mm. Central Veracruz, Mexico, through Central America to northern South America, sea level to 1500 m elevation. Two subspecies: Kinosternon 1. leucostomum (Veracruz, Mexico, to Nicaragua) and K. 1. postinguinale Core 1887 (Nicaragua to Ecuador) Kinosternon scorpioides (LiNNABUS 1766), Systema Naturae, ed. 12: 362; type locality: Surinam. Carapace length to 270 mm. Southern Tamaulipas, Mexico, through Central America to South America, sea level ‘to 1500 m elevation. The subspecies K. s. scor- pioides (Panama southward), K. s. abaxillare om Baur 1925 (Central Depression of Chiapas), K. 8, albogulare Duéxit. & Bocourr 1870) (Honduras to Panama), K. s cruentatum DumeriL & BrBRON 1851 (Mexico to Honduras) occur in Central America. ¥ Kinostemon angustipons IB Kinostemon leucostomum \ IB Kinosternon creaseri ¥ Kinosternon acutum ® Kinosternon scorpioidesKinosternidae Fig. 50. Kinosternon scorpioides (near Porte- golpe, Guanacaste, Costa Rica). 1300 Key to Kinosternon 1 a Bridge very narrow, its width less than 21% of carapace length ossstneenm Kénosternon angustipons b Bridge less narroy 21% of carapace le its width more than Yet sn couce 2a Length of gular scute about half the length of anterior moveable lobe of plas tron (Fig. 52a,b) 3 bb Length of gular scute distinctly leas than half the length of anterior moveable lobe of plastron (Fig. 52¢,4) 1... 4 3 a Anterior moveable lobe of plastron as long as or shorter than fixed lobe (ig, 52a) Soouiianinn Kinosternon acutum b Anterior moveable lobe of plastron con- spicuously longer than fixed lobe (Fig 52b) Kinosternon creaseri 4 a Anterior moveable lobe of plastron con- spicuously longer than fixed lobe (Fig. 52c); axillary scute usually not in contact with inguinal scute; carapace never with more than one keel; a pale postorbital stripe usually present (Fig. 47) ... Kinosternon leucostomum b Anterior moveable lobe of plastron as long as fixed lobe (Fig. 52d); axillary seute usually in contact with ‘inguinal scute; carapace usually with three keels; no pale postorbital stripe i Kinosternon scorpioides Further Reading Iverson 1976 (Kinosternon in Belize); IVERSON 1980a, b, 1983b, Fig. 51, Kinosternon leucostomum (Tortuguero, Costa Rica). anterior moveable lobe fixed lobe posterior moveable lobe a. K. acutum b. K. ereaseri ie ©. K, leucostomum gular scute axillary scute Inguinal scute J. K, scorpioides Fig. 52. Plastron of Kinosternon species Axillary and inguinal scutes orange Adopted after Lex (1996)Staurotypus The genus Staurotypus includes two spe- cies, the Pacific coast giant musk turtle (S. salvinii) and the Mexican giant musk turtle (S. triporeatus), both of which occur in Central America. These are aggressive turtles that need to be handled with great caution. With their strong, sharp jaws, these animals can cause serious bite wounds. The strongly developed dorsal keels help to stabilize the carapace, there- by making it more difficult for predators (e.g., Crocodylus moreletii) to break the strong shell. The species of this turtle genus are strictly aquatic inhabitants of lakes and slow moving rivers with soft bottoms and rich aquatic vegetation. Giant musk turtles are not great swim- mers; these animals prefer to walk around on the bottom of their aquatic habitat. They are carnivorous, feeding on crabs, worms, snai aquatic insects, amphib- ians, and fishes. Additionally, smaller turtles (especially of the genus Kino- sternon) are preyed upon by the giant musk turtle and their shells broken with the aid of their strong jaws (CAMPBELL 1998a). During the reproductive season, the female buries several clutches, each with three to ten eggs, on the riverbank. The hatchlings open the eggshells after 120 to 207 days of incubation. At that time, they have a shell length of 25 to 31 mm (S, salvinii) or 30 to 35 mm (S. tripor- catus) (Ernst & Barpour 1989, KOHLER 1997). Kinosternidae Fig. 53. Staurotypus triporcatus (Tabasco, Mexico). Photo: J. Haft Staurotypus salvinii Gray 1864, Proc. Zool. Soc, London 1864: 127; type locality: Huamuchil, Guatemala. Carapace length to 250 mm. Pacific versant of northern Central America from Oaxaca, Mexico, to El Salvador, sea level to 700 m elevation. Staurotypus triporcatus (WiEGMANN 1828), Isis von Oken 21: 364; type locality: Rio Alvarado, Veracruz, Mexico. Carapace length to 379 mm. On Caribbean versant from Veracruz, Mexico, through the base of Yucatan Peninsula to western Honduras, sea level to 300 m elevation. IB Staurotypus triporcatus WH Staurotypus salvinii_ Te Staurotypus salvinii (ventral view; uintla, Guatemala) Fig. 54. near Iztapa, E 45,46 Kinosternidae Key to Staurotypus 1 a Dorsolateral keels extend the length of carapace from anterior to posterior mar- ginals (Fig. 55a); anterior moveable lobe of plastron shorter than posterior moveable lobe ... ... Staurotypus triporeatus b Dorsolateral keels do not extend the length of carapace, but end on Ist and 4th plourals (Fig. 55b); anterior moveable lobe of plastron longer than posterior moveable lobe onnven Staurotypus salvinit Further Reading DEAN & BICKHAM 1983, IVERSON 1983a, 1985. a. Staurotypus triporcatus _. Staurotypus salvini Abb. 55. Carapace with keels (shaded). Fig. 56. Staurotypus salvinii (near I2tapa, Escuintla, Guatemala), Fig. 57. Adult Geochelone carbonaria (Darién, Panama) Photo: H. Birkahn Testudinidae Geochelone ‘The red-footed tortoise (Geochelone carbo- naria), which occurs in Panama, is the only South American tortoise that reaches as far north as southern Central America (LEGLER 1963). It is otherwise distributed mostly in tropical South America. The spe- cies was introduced by humans on several Caribbean islands, such as Isla de Maiz Grande, Nicaragua (PrircHarD & TREBBAU 1984). In Panama, the red-footed tortoise is found mostly in forested areas and does not occur in the humid savannas (Lec! 1963). These diurnal animals dig burrows in the ground into which they retreat at night, Red-footed tortoises feed mainly on grasses, flowers, fallen fruit, and carrion. In Panama, this tortoise is hunted and eaten by the local people, as is the case in other parts of its range to the south. The egg-laying season is from June through September. During this period, the fe- males bury their clutches with 2 to 15 eggs in the ground. Females probably produce several clutches per season. The eggs are somewhat clongate (40-59 mm x 34-48 mm) and have a rigid shell. After an incu- bation period of 116 to 185 days, the ju-veniles hatch; they have a shell length of 39 to 45 mm (ERNST & BARBOUR 1989, KOHLER 1997). Reproductive maturity is reached at a shell length of 200 to 250 mm R 1963) Geochelone carbonaria (Sprx 1824), Anim, Spec. Nov, Testud. Brasil.: 22; type locality: “Habitat sub cognomine ‘Capitary’ (2) ad flu men Amazonum,” South America. Carapac length to 51 cm. Southeastern Panama through most of the Amazon Basin south- ward to Rio de Janeiro and westward to Paraguay and to northern Argentina, sea level to 300 m elevation. Further Reading LEGLER 1963; Ernst & LEuTERITZ 1999, Fig. 58. Juvenile Geochelone carbonaria (Cordoba, Colombia) Photo: M. Lundberg Cheloniidae, Dermochelyidae turtle (Eretmochelys reef, Photo: J. Pichler Sea Turtles (Cheloniidae, Dermochelyidae) Sea turtles were already to be found around the world during the Cretaceous period, with fossil finds dating back at least 200 million years. Thus, the ancestors of our modern-day sea turtles lived side-by-side with the dinosaurs, but were able to outlive the gigantic Plesiosaurus and Ichthyosaurus. The modern species came into existence around 10 ~ 60 million years ago and comprise, along with marine iguanas and sca snakes, the only marine reptiles. All species of sea turtles have forelimbs that have evolved into paddle- shaped fins, with only one or two claws. Further adaptations to the aquatic life style are salt glands, by means of which superfluous salt, which has been ingested through feeding in sea water, is expelled. These animals can reach adult weight of between 35 and 500 kg. 4748 Cheloniidae, Dermochelyidae ‘The females craw] up onto certain be: at dusk, in order to dig a nest and deposit their eggs. After filling in the nest, the female returns to the sea during the same night, The young sea turtles hatch after an incubation period of approximately two months and abandon the nest together for the sea during the night. After attaining sexual maturity a number of years later, the adult females return to the original beach for nesting. However, this generali- zation of a sea turtle’s life cycle only applies in a general way. The eight known species of sea turtles vary considerably with regard to their geographical location, mating cycles, and behavior. Moreover, the various populations of a particular species exhibit a number of differences due to geo- graphical distribution. The following in- formation about the nesting seasons of the individual species relates in each case to Central America. Six species of sea turtles from two families and five genera nest on the beaches of Central America. There used to be mass landings of sea turtles (so- called “arribadas” or “arribazones”) in many Central American countries (Costa Rica, Honduras, Nicaragua, and Panama to name a few), in which thousands of females of the species Lepidochelys oliva- cea and others came onto land all at once in order to lay their eggs on one beach. This phenomenon only takes place in Costa Rica presently (CORNELIUS 1995). Among the most important factors that have led to the rapid decrease in sea turtle populations are their massive commercial use (the collection of eggs and killing of adult turtles), the plundering of nests by feral animals (dogs and pigs), shrimp fishing, and sea pollution (cases of death due to ingestion of tar and plastic refuse). Turtle products are offered in numerous tropical and subtropical countries as sou- venirs for tourists. It is completely ir- responsible and illegal to sell and abuse sea turtles or products made from these animals as souvenirs, in light of their endangered status Fig. 60. Green turtles (Chelonia mydas) copulating Photo: J. Pichler Fig. 61, Lepidochelys olivacea nesting at Playa Bavaria, Guanacaste, Costa Rica Fig. 62. Sea turtle hatchlings on their dangerous walk to the sea. Photo; J. PichlerFig. 63. Green turtle (Chelonia mydas) with shark sucker Photo: J. Pichler The loggerhead sea turtle (Caretta caretta) is found worldwide, primarily in subtropical regions, and less in tropical cli- mates. It is found primarily on the Caribbean side of Central America. The most important nesting grounds of this species are located in Florida and South Carolina in the U.S. while in Central America few specimens come onto land to lay their eggs, mostly in the northeastern portion of the Yucatén Peninsula, In Caretta caretta, egg laying has been docu- mented from April to July. After a single short mating phase, a female produces 2 — 5 clutches, each with 40 190 eggs at intervals of approximately two week: From these, the young will hatch after 68 days of incubation (MAGNUSON ct al 1990, MARQuEZ 1990). While it was earlier proposed that two spe- cies of the genus Chelonia (green turtles) populated the coastal regions of Central America (C. mydas on the Caribbean side and C. agassizii on the Pacific side), it is currently believed that only a single spe- cies is represented in the genus (ERNST & Barwour 1989, Kart & Bowen 1999), The shell of C. mydas can measure up to 140 cm in length. Important nesting grounds Cheloniidae, Dermochelyidae | Fig. 64, Loggerhead sea turtle (Caretta caretta). Photo: R. D. Bartlett are located along the coast of the Mexican state of Michoacan, as well as in El Salvador (Jiquilisco), Guatemala, and Costa Rica (Tortuguero). Single females come on land in many areas a Central American Pacific and coasts to nest. During one nesting season, which ean last from August until January (Pacific side) or April until October (Caribbean side), a female can produce 2 — 5 clutches (rarely up to 8) at intervals of 12 — 14 days, each with 38 ~ 120 (rarely up to 195) eggs, from which the young hatch after 50 — 75 days (MAGNUSON et al, 1990, Marquez 1990). The animals reach sexual maturity between eight and thirteen years of age. A female does not come on land every year for nesting, rather every 2-5 years. Green turtles subsist mainly on vegetarian fare (various sea algae and sea grass), but also feed upon sponges, mol- luscs, jellyfish, and fish, ‘The hawksbill sea turtle (Zretmochelys imbricata) is found around the world in all tropical seas and on both the Pacific, as well as the Caribbean side of Central America. Important Central American nesting grounds for this species are found on the coast of the Yucatan Peninsula, as 49Cheloniidae, Dermochelyidae Fig. 65. Kemp’s ridley sea turtle (Lepidochelys kempii) swimming in shallow water. Photo: R. D. Bartlett well as near Tortuguero (Costa Rica), while nesting specimens have only been sporadically reported elsewhere. The egg- laying season is from April until November, but it varies according to re- gion: Mexico and Belize — April to August; Guatemala ~ May to November; Honduras — June to September; Nicaragua — May to October; Costa Rica - May to November (Marquez 1990). During one season, each female comes on land once or twice to bury, clutches consisting of 93 - 223 eggs cach. A female only nests every 2 — 3 years. The incubation period lasts 47 — 75 days and the young have a shell-length of 38 — 46 mm (MARQUEZ 1990). The hawksbill sea turtle is mainly carnivorous, and its food ranges from crabs, sponges, sea anemones, jellyfish, molluscs, squid, and fish to sea algae. However, there are apparently great geographical differences with regard to its range of foods. According to the in- vestigations by Carr & SrancyK (1975), the hawksbill sea turtles in the Tortu- guero region (Costa Rica) feed primarily upon certain sponges. Both species of the genus Lepidochelys (ridley sea turtle) are found on the coasts of Central America, with the Kemp’s ridley sea turtle (L. kempii) only found on the Yucatan Peninsula in this region, and the olive ridley sea turtle (L. olivacea) along the entire Pacific coast of Central America. The latter is the most common species of sea turtle on the Pacific coast of Central America, with its most important nesting sites in Nicaragua (Masachapa, Pochomil, Boquita) and Costa Rica (Nancite, Ostional) (CORNELIUS 1995). Playa Nancite is a beach in the Santa Rosa National Park in Costa Rica that is naturally protected by the sharp rocks and thorny bushes that surround it: ‘There, one can experience the spectacular “arribadas” from July until September, in which up to 8000 L. olivacea females crawl up onto the beach to lay their eggs within a span of a few days. Important nesting grounds for L. kempii are located along the Gulf Coast, with the beaches near Rancho Nuevo (Tamaulipas, Mexico) of by far the greatest importance. In the 1940's, the amateur filmmaker Andrés Herrera docu- mented a mass landing of approximately 40,000 L. kempii near Rancho Nuevo (Macnuson et al, 1990). This type of mass landing is known only in the case of the species of the genus Lepidochelys. It is assumed that the simultaneous hatching of many young increases the chances that the necessary number of turtles survive, in spite of numerous predators. Lepidochelys kempii also differs from the other sea turt- Jes in that it comes onto the beach during the day to lay its eggs, while the other spe- cies nest at night. Also remarkable is the fact that the females of both these species nest nearly every year, producing 2 - 7 clutches, each with 50 — 140 eggs in one season, After an incubation period of 45 — 65 days, the young hatch with a shell length of 35 - 45 mm. (MARQUEZ 1990) Ridley sea turtles feed on crabs and other invertebrates (sponges, sea anemones, jel- lyfish, and squid).Fig. 66, Leatherback sea turtle (Dermochelys coriacea) nesting at Playa Naranjo, Costa Rica. Photo: M. Franzen The largest turtle living today is the leatherback sea turtle (Dermochelys coriacea), with the largest specimen ever measured having a shell length of 256.5 cm, and a mass of 916 kg. (MARQUEZ 1990). Most specimens, however, have shells with a length of 150 — 170 cm. This species can be found all over the world, as they are able to populate cooler seas than other sea turtle species. Leatherback sea turtles usually stay in the open ocean and are observed only occasionally near the coast. They can submerge to great depths. Important beaches for nesting in Central ‘America are located in Costa Rica (Playa Naranjo, Playa Tamarindo, between Tortuguero and Lim6n) and Panama. While. the nesting season on the Caribbean side lasts from April to October, the nesting season on the Pacific side lasts from October to March. The females come on land to nest every 2-3 years so that they can bury 4-5 clutches of 61-126 eggs each in a single season. The incubation period is 50-78 days and the young have a shell length of 51-68 mm (Marquez 1990). Leatherback sea turtles are mainly carni- vorous and feed on jellyfish and pelagic crustaceans, and occasionally also on sea algae. Cheloniidae, Dermochelyidae Fig. 67. Chelonia mydas. Photo: J, Pichler Cheloniidae Caretta caretta (LINNABUS 1758), Systema Naturae, ed, 10: 197; type locality: “insulas Americanas.” Carapace length to 105 cm. Worldwide; in Central America mostly in the Caribbean. Chelonia mydas (LinNasus 1758), Systema Naturae, ed. 10: 197; type locality: “insulas Pelagi: insulam Adscensionis.” Carapace length to 140 cm. Worldwide. Eretmochelys imbricata (LINNakus 1766), Systema Naturae, ed. 12: 350; type locality: “Mari Americano, Asiatico.” Carapace length to 114 cm, Worldwide. Lepidochelys_kempii (GakMAN 1880), Bull. Mus. Comp. Zool. 6: 123; type locality: Gulf of Mexico. Carapace length to 75 em. Gulf of Mexico. Lepidochelys olivacea (ESCHSCHOLT 1829), Zool. Atlas 1: 3; type locality: Manila Bay, Philippines. Garapace length to 75 em. World- wide; in Central America only in the Pacific. Dermochelyidae Dermochelys coriacea (LINNABUS 1766), Systema Naturae, ed. 12: 350; type locality: “maris Tyrrheni oram in agro Laurentiano.” Carapace length to 256,5 em, Worldwide.Cheloniidae, Dermochelyidae Fig. 68. Eretmochelys imbricata Photo: J. Pichler x ay ey ff wen the " a “J 7 a a 4 Caretta caretla 4 Eretmochelys inva WFR © Lepidochelys kempiiKey to sea turtles 1 a Shell covered with a continuous layer of undivided skin (Fig. 70a) . sss, Dermochelys coriacea b Shell covered with distinct horny scutes (Fig. T0b-f) a. S, rosaurae Abb. 134. Dorsal scales in Sphaerodactylus.b Dorsal scales weekly keeled, parietal scales usually smooth; subcaudals arranged in sequences of one small scale, one large seale, and a pair of small scales; 62-81 dorsals between levels of axilla and groin ...... Sphaerodactylus lineolatus 7 a Dorsal scales smooth .. b Dorsal seales keeled .. ocellated blotches... .. Sphaei Sphaerodactylus glaucus 8 8 a 38-50 dorsals between levels of axilla and groin; dorsum with longitudinal rows of jactylus argus bb 47-74 dorsals between levels of axilla and groin; dorsal pattern variable, but without longitudinal rows of ocellated blotches . Sphaerodactylus millepunctatus Further Reading Harxis & KLuGE '1984 (Sphaerodactylus of Central America), KOHLER 2001c. Thecadactylus rapicauda (Quintana Roo, Mexico). Photo: H. Bahena B. & Thocadactyus rapicouda CAGE Wo Gekkonidae Fig. 136, Theeadactylus rapicauda (Bartola, Rio San Juan, Nicaragua) Thecadactylus Thecadactylus rapicauda is a nocturnal, arboreal inhabitant of the tropical rain forest. At night, these large geckos can be observed on the trunks of large trees some 3 to 6 m above the ground, while during the daytime they retreat into bark crevices and hollow branches. Other typical day- time hiding places include piles of leaf litter among the buttress roots of large trees. Turnip-tailed geckos are also com- monly encountered on house walls. They feed on invertebrates (mostly crickets, moths, grasshoppers, and spiders) and small lizards. Like most geckos, 7. rapi- cauda is capable of vocalization and its call sounds like “chack-chack-chack” Bi 1944), During the reproductive season, female T) rapicauda lay several clutches each consisting of a single egg. Thecadactylus rapicauda (HouTtuyN 1782), Verh, Zeeuw. Genootsch. Wet. Vlissingen 9: 323; type locality: West Indies. SVL to 121 mm. Mexico through Central America to tropical South America; also on the Lesser Antilles, sea level to 1050 m elevation Further Reading TayLor 1956, AVILA-P1 Bauer 2002. 1995, RUSSEL & 7980 Gymnophthalmidae Gymnophthalmidae Prescn (1983) suggested separating the so-called “microteiids” into a separate family (Gymnophthalmidae) from the so- called “macroteiids” (Teiidac). This view has been accepted by the majority of her- petologists working in Central America (IBANEZ et al. 1996, SavacE 2002, WiLson pers. comm., McCRani# pers. comm.), whereas others preferred not to recognize the Gymnophthalmidae as a distinct family (Harris 1985, Myers & DONNELLY 1996). The lizards of the family Gymnoph- thalmidae, including about 30 genera with approximately 150 species, are distributed in temperate, subtropical, and tropical areas of southern Mexico, Central America, and South America. Limb reduc- tion has occurred several times within Gymnophthalmidae, and in many of these it is accompanied by body elongation. All gymnophthalmids are oviparous, usually producing two eggs per clutch. Key to genera of Gymnophthalmidae 1 a Without frontoparieta Is, frontal, therefore, in contact with parietals (Fig. 140a); no outer ear opening present; extremities greatly reduced .. Bachia Fig. 187. Neusticurus apodemus (Palmar, Costa Rica). Photo: R. W. Van Devender b Frontoparietals present, frontal, therefore, separate from parietals (Fig. 140b); outer ear opening present; extremities normal 2 2 a Without eyelids ....... Gymnophthalmus b Eyelids present seonenseeniee B 3 a Posterior edges of the parietal scales and the interparietal form a straight line (Fig. 141a); upper surface of tongue covered with plicae Ptychoglossus b Middle parietal seale clearly protrudes past the posterior edges of the parietal scales (Fig. 141b); Upper surface of tongue covered with papillae .. ner | 4 a Dorsal scalation heterogeneous (arge ubereular scales sprinkled in with the fer ground scalation) .. asics b Dorsal sealation homogeneous .... 6 5 a Prefrontals and frontal fragmented into small irregular scales (Fig, 142b); a clearly enlarged scale between first finger and wrist .. : Neusticurus b Prefrontals and frontal form large shields (Fig. 142); no clearly enlarged scale be- tween first finger and WriSt sn. su. Eehinosaura Anadia ist 6 a Dorsal scales smooth ... sal scales keeled ble longitudinal row of widened gular scales; a vertebral pale stripe usu- ally progent .......:0000- Prionodactylus b No double longitudinal row of widened gular scales; no vertebral pale stripe .... an sous Leposoma Fig, 138, Leposoma southi (Nusagandi, Panama).frontoparietals interparietal supfacculars pariéals — superciliaries Fig. 189. Scalation of dorsal surface of head in Prionodactylus. Fig. 140 Dorsum of head (frontal \ brown; parietals orange). a. Bachia b. Gymnophthaimus Fig. 141 Dorsum of rietal brown; parietals orange). a. Plychogiossus _b. Leposoma Fig. 142. Dorsum of tals brown; frontals orange), b. Echinosaura —_b. Neusticurus head (interpa- head (prefron- Gymnophthalmidae Fig. 143. Anadia ocellata (Rara Avis, osta Rica). Photo: T, Leenders Anadia In Central America, two species of the genus Anadia, A. ocellata and A. vittata, are found (OFTEDAL 1974). Both are very slender diurnal tree dwellers with long prehensile tails. The habitat of these lizards is the rain forest where they prefer trees with a plentiful supply of bromel- iads. Large bromeliads also serve these animals as hiding places. Their diet con- sists primarily of insects and spiders. 81Gymnophthalmidae Fig. 144, Anadia ocellata (Provincia Limén, Costa Rica). Photo: R. W. Van Devender ‘Anadia ocellata Gray 1845, Cat. Lizards Brit. Mus.: 58; type locality: Tropical America. SVL. to 75 mm, Costa Rica and western Panama; 500-1310 m elevation in premontane rain forest. Anadia vittata BOULENGER 1913, Proc. Zool. Soe, London 1913: 1033; type locality: Pea Lisa, 300 ft, Condoto, Choco, Colombia. SVL. to 67 mm. Eastern Panama to southern Colombia; 0-100 m elevation in lowland rain forest. Key to Anadia 1 a Pale dorsolateral stripes maximum of two ‘ales wide; obvious ocelli on body sides; 12 scales at midbody; males with a total of 11 femoral pores ..... Anadia ocellata b Pale dorsolateral stripes at least three scales wide; ocelli, if present, not obvious; 30-37 seales at midbody; males with a total of 11-14 femoral pores ..... Anadia vittata Further Reading OrreDAL 1974. 4 Anadia ocellata @ Anadia vittata 82 Bachia ‘The small lizards of the genus Bachia are slender ground dwellers. Two species of the predominantly South American genus reach the southern portion of Central America (Dixon 1973, McDiarmMip & DEWEESE 1977). Bachia blairi was consid- ered by Dixon (1973) to be synonymous with B. pallidiceps, a decision for which there was no evidence. McDiarmip & se (1977) were able to show that Bachia blairi was a separate species, which differs from B. pallidiceps in coloration, build, and several scale characters. Representatives of this genus ‘are characterized by the absence of an external ear opening and by the tendency to reduced limb size. On the front limbs, B. blairi and B. pallidiceps have four toes, whereas the rear limbs have only three. They are highly secretive ground dwellers in the rain forest. These lizards are usual- ly found in loose dirt or under trees or logs (Dunn 1940a). Bachia blairi (DUNN 1940a): Proc. Acad. Nat. Sci. Philadelphia 9: type locality Puerto Armuelles, Chiriqui, Panama. SVL to 70 mm. Southeastern Costa Rica and adja- cent areas in Panama, sea level to 40 m ele- vation. Bachia pallidiceps (Core 1862c), Proc. Acad. Nat. Sci. Philadelphia 14: 356, type locality: Rio Truando region, Colombia, SVL to 73 mm. Eastern Panama to northwestern Colombia, sea level to 100 m elevation. > 4 Bachia blairi © Bachia paliidicepsKey to Bachia 1 a Pale dorsolateral stripes present; 22-24 seales at midbody; 45-48 cross rows of dor- sal scales; 30-33 cross rows of ventral scales ... Bachia pallidiceps b Without pale dorsolateral stripes; 24-25 seales at midbody; 43-45 cross rows of dor- sal scales; 29-31’ cross rows of ventral les .... Bachia blairi Further Reading Dixon 1973, McDIARMID & DEWE 1977. . 145, Bachia pallidiceps (southeastern ta Rica). Photo: Rt. W. Van Devender Echinosaura The lizards of the genus Echinosaura are small terrestrial ‘microteiids” that are dis- tributed from western Panama across northwestern Colombia to central-western Ecuador, In his revision of the genus, Uva (1965a) recognized a single species (E. horrida) divided into three subspecies. Recently, Frirrs et al. (2002) described a new species of Echinosaura and elevated the former subspecies to species. Thus, as currently understood, the genus Echino- saura contains four species, two of which (, palmeri and E. panamensis) occur in southern Central America. Gymnophthalmidae Although usually found in the vicinity of streams, these lizards live somewhat more terrestrially than the semiaquatic Neusti- curus (UzzELL 1965a, b). According to Dunn (1944), BREDER (1946), and UzzeLL (1965a), active Echinosaura can be observed during the day as well as at night; how- ever, they appear to be predominantly noc- turnal (Uzzwi1. 1965a). The animals are active among the leaves of the ground cover or hide under rocks and in rotting plant material. Several reports state that Echinosaura will become absolutely rigid when handled (UzzELL 1965a, LEVITON & ANDERSON 1966), The animals will stiffen their bodies and press their limbs against their sides. If they are turned on their backs they will quickly turn over and freeze again. As with most microteiids, Echinosaura produces clutches, each with two soft-shelled eggs (U2zELL 1965a). Echinosaura palmeri BOULENGER 1911, Ann. Mag. Nat. Hist. (8) 7: 23; type locality: Noananod, Rio San Juan, Choco, Colombia SVL to 70 mm. Colombia and eastern Panama, near sea level to 1400 m elevation. Echinosaura panamensis BARBOUR 1924, Proc. New England Zool. Club 9: 8; type lo- cality: La Loma, Bocas del Toro Province, Panama. SVL to 71 mm. Western Panama, 300-1500 m elevation. Fig. 146, Echinosaura panamensis (El Copé, Panama), Photo: R, W. Van Devender 83Gymnophthalmidae =~ ances Fig. 147, Echinosaura panamensis (E] Copé, Panama). Photo: R. W. Van Devender Key to Echinosaura 1a Frontal plate transversely divided (Fig. 148a)... Echinosaura panamensis b Frontal plate single Fig, 148b) .. san Eehinosaura palmeri Further Reading DuNNW 1944, Uzze. 1965a, Frirrs et al. 2002. Gymnophthalmus Gymnophthalmus speciosus is the sole representative of the microteiids to extend as far north as Mexico. Whereas this sp cies has, at times, been divided into a num- ber of subspecies (MERTENS 1952c, PeTers & DoNoso-Barros 1970), more recent in- ‘be paimen — a. E. panamensis Fig. 148, Dorsal head sealation in Echinosaura (frontal orange). 84 Fig. 149. Gymnophthalmus speciosus (near Guanagasapa, Escuintla, Guatemala). vestigations indicate that such a division is not justified (KOHLER 1996c). The usually intensely red tail is particularly conspic- uous, and gives this small lizard the local name “Cola Roja” (“red tail”). These are typical undergrowth dwellers that spend their days searching for prey (mostly inver- tebrates) under the wilted leaves. The population density is apparently seasonal, with an increase during the dry season (TELFORD 1971). According to the investiga- tions of TELFORD (1971) the female of G. speciosus in Panama produces clutches of 1-2, sometimes as many as 3 or 4 eggs. The egg-laying season, during which the fe- males produce up to three clutches, lasts from October to March, with hatchlings being observed in the months from §@ Echinosaura panamensis ( i Echinosaura palmeri AFig. 150. Gymnophthalmus speciosus (Isla de Roatan, Honduras), February to May (TeLrorp 1971, ALVAREZ DEL Toro 1983), The young measure 17-19 mm at hatching. Gymnophthalmus speciosus (HALLOWELL 1861), Proc. Acad. Nat. Sci. Philadelphia 12: 484; type locality: Nicaragua. SVL to 40 mm. Isthmus of Tehuantepec on the Pacific side and Guatemala on the Caribbean side south- wards into northern South America, as well as in the Lesser Antilles, sea level to 1220 m elevation. No subspecies are recognized (KOHLER 1996a). Further Reading CoLe et al, 1990 (evolution and systematics of the genus); TeuroRD 1971 (ecology, reprodue- tion). === Gymnophthalmidae Leposoma Like most of the microteiids, the genus Leposoma is also most heavily distributed in South America. Only Leposoma rugiceps and L. southi occur in southern Central America. The males of L. southi generally display an intensive orange coloration on the underside of the body. Similar to spe- cies of the genera Ptychoglossus and Gym- nophthalmus, the Leposoma species are also predominantly undergrowth dwellers that are found between the wilting leaves on the rain forest floor. The following details on the reproduction of Leposoma rugiceps are based on the investigations of TeLrorp (1971) in Panama. During the egg-laying season, which lasts for several months (May to November) during the rainy season, the females produce clutches of two eggs each. The young hatch with a SVL of 16-19 mm. They are sexually ma- ture within six to eight months. Leposoma rugiceps (Core 1869), Proc. Acad. Nat. Sci. Philadelphia 20: 305; type locality: Rfo Magdalena, Colombia. SVL to 44 mm. Eastern Panama to Colombia, sea level to 500 ‘m elevation. Leposoma southi RutTHvEN & GAIGE 1924, Pap. Mus. Zool., Univ, Michigan 147: 1; type locality: Progreso, Chiriqui, Panama. SVL to ‘55 mm. Southern Costa Rica to Colombia, sea level to 700 m elevation Fig. 151, Leposoma rugiceps (Panama), Photo: M, A. Guerra 85Gymnophthalmidae Fig. 152. Leposoma rugiceps (Cordoba, Fig. 154. Leposoma southi (southeastern Costa Colombia). Photo: M. Lundberg Rica). Photo: R. W. Van Devender Key to Leposoma 1 a Frontonasal divided ....... Leposoma southi b Frontonasal undivided Leposoma rugiceps Further Reading Ruat 1952, UzzeLL & Barry 1971 (taxonomy), ‘TeLFoRD 1971 (ecology). Abb, 153. Leposoma rugiceps (Pearl Islands, Panama). Foto: C. W. Myers a. L. southii b. L. rugiceps ig. 155, Dorsal head sealation in Leposoma (frontonasals orange). ] 4 Neusticurus apodemus © Prionodactylus vertebralis 1B Leposoma southi = Leposoma rugiceps 86Fig. 156. Neusticurus apodemus (Palmar, Costa Rica’ Photo: R. W. Van Devender Neusticurus Water tegus of the genus Neusticurus have their primary distribution in South America, with only one species, Neusticurus apodemus, reaching southern Central America in Costa Rica. As with all species of this genus, N. apodemus is semiaquatic, found along fast flowing streams in the rain forest. Usually water tegus are found on rocks or roots by the edge of the stream, and they flee into the water at the slightest disturbance. Some active N. apodemus can also be found in leaf litter on the forest floor (UzzELL 1965b). Although representatives of the genus Neusticurus are active primarily during the day, N. apodemus has been observed swimming in the water at night, as well as during the day (UzzELL 1965b). While swimming, Neusticurus will fold in their legs alongside the body and hold only the head above the water's surface while making snake-like movements with the body and tail, Nothing is known about their reproduction, but most likely N. apodemus, as with other Neusticurus spe- cies, produces two eggs per clutch, which are deposited by the females under rotting logs (UzzeLL 1965b). Gymnophthalmidae Neusticurus apodemus Uzzett. 1965b, Bull ‘Amer, Mus. Nat. Hist. 132: 298; type locality: 15 km SW San Isidro del General, 865 m ele- vation, San José, Costa Rica. SVL to 47 mm. Southern portion of the Puntarenas Province, Costa Riea, near sea level to 900 m elevation, Further Reading Uzzeu1 1965b. Prionodactylus Only one species of this predominantly South American genus Prionodactylus reaches the border area between Panama and Colombia in the extreme southern portion of Central America. These agile lizards are ground dwellers in the rain forest, where they search for prey (mostly insects and spiders) during the day. To reproduce, the female P. vertebralis pro- duces clutches of two eggs each, which are only superficially buried (UzzeLL 1973). Prionodactylus vertebralis (O'SHAUGHNESSY 1879), Ann, Mag. Nat. Hist. (5) 4: 298; type locality: Intac, Ecuador, SVL to 68 mm. Extreme eastern Panama (bordering Colombia) to northwestern Peru, 700-2500 m elevation. Further Reading Ure. 1973. Fig. 157. Prionodactylus vertebralis (Cerro Mali, Darién, Panama), —_ Photo: C. W. Myers 8788 Gymnophthalmidac seececsaem scene snememcene em atm ER aL SN Fig. 158. Ptychoglossus festae (Nusagandi, Panama). Ptychoglossus ‘The primary distribution of the genus Ptychoglossus is in northwestern South America, with only a few species reaching Central America. Whereas P. plicatus reaches at least as far as northern Costa Rica, P. festae and P. myersi are restricted in their distribution to eastern Panama. Ptychoglossus festae is most easily distin- guished externally by the presence of small lateral scales between the dorsal and ven- tral scales. However, the only definitive dif- ference between the other two species can only be seen in the hemipenial morphology (Harris 1994). The species of the genus Ptychoglossus are small diurnal under- growth dwellers. Thus, they occupy an eco- logical niche similar to that of members of the genera Leposoma and Gymnophthalmus. Ptychoglossus plicatus vy ¥ Fig. 159. Ptychoglossus plicatus in Costa Rica. Photo: R. W. Van Devender Ptychoglossus festae (PrRAcca 1896), Boll Mus. Zool. Comp. Anat. Univ. Torino'11: 2; type locality: Rio Cianati, Darién, Panama. SVL to 65 mm. Eastern Panama and north- ern Colombia; sea level to 800 m elevation in rain forest. Ptychoglossus myersi Hannis 1994, Herpetol. Monogr. 8: 250; type locality: cloud forest on the SE slope of Cerro Pirre, 1000 m, 7°50'N, 77°43'W, Serrania de Pirre, Darién, Panama, SVL to 52 mm. Border area of Panama and Colombia; 800-1300 m elevation in cloud forest. Ptychoglossus plicatus (TavLox 1949b), Univ, Kansas Sci. Bull, 38: 272; type locality: Morehouse Finca, 5 miles SW Turrialba, Cartago, Costa Riea. SVL to 66 mm. Costa Rica to northern Colombia; 400-1890 m ele- vation in rain forest and cloud forest. Ptychoglossus festae 4 Plychoglossus myersiFig. 160. Ptychoglossus myersi (between Rio Jagué and Rio Imamadé, Darién, Panama). Photo: ©, W. Myers Key to Ptychoglossus 1 a A longitudinal fold separates dorsal scales from ventral scales (Fig. 161a) ...... Ptychoglossus festae b Without a longitudinal fold between dorsal seales and ventral scales (Fig. 161b) wv... 2a Hemipenis with 23-28 ring folds .. Plychoglossus plicatus b Hemipenis with 9-12 ring folds .. nouns Phyohoglossus myersi Further Reading RuIBAL 1952, Harris 1994. a. Plychoglossus festae . Ptychoglossus plicatus Abb, 161, Lateral body scales. ~==ees Helodermatidae Helodermatidae Of the two surviving species of the genus Heloderma, only one (H. horridum) is found in Central America. These are robust lizards that can reach a total length of over 70 cm, with massive heads and short, powerful limbs. They have fat tails, which are not capable of autotomy or regenera- tion. The back is covered with large ossified knobby scales. The tongue is fleshy, but forked at the tip. The Heloderma species are the only surviving venomous lizards. Elongated, multi-lobed poison glands are located along both sides of the lower jaw. A separate opening from each lobe of the gland leads to the base of the grooved teeth, which are up to 6 mm in length. When the teeth clamp down, the venom is squeezed out of the glands by the pressure of the jaw muscles and flows through the grooved teeth into the wound primarily by capillary action. Because Gila monsters are shy and live very secretively, they pose no threat to humans. Biting incidents occur only when the animals are provoked or captured. These otherwise calm lizards can launch themselves with unexpected speed and bite very suddenly. A Gila mon- ster will not readily release anything that comes between its powerful jaws, and it will bite down forcibly. Gila monster bites should be taken very seriously and can lead to localized as well as systemic symp- toms of envenomation, which absolutely require medical attention (MERS 1992). In Mexico and Central America, three sub- species of Heloderma horridum are recog- nized: H. h. horridum (WInGMANN 1829) along the Pacific side of Mexico from south- ern Sonora to Chiapas, H. h. alvarezi Bocert & MARTIN DEL CAMPO 1956 in the Rio Grijalva Basin, Chiapas, and H. h. charlesbogerti CAMPBELL & VANNINI 1988 in the Rio Motagua Basin, Guatemala. Gila monsters are crepuscular and nocturnal ground-dwellers in dry forest that become active just after dusk. On cloudy days, they can sometimes be observed during the day outside of their hiding places (burrows, 8990 Helodermatidae Fig. 162. Heloderma horridum charlesbogerti (E] Arenal, Zacapa, Guatemala), which they have either excavated them- selves or were pre-existing). They hunt down their prey, which consists of nestling rodents and birds, as well as bird and rep- tile eggs, with constant tongue flicking. Gila monsters can survive for several months without food by living off the fat stored in their tails. During the rainy sea- son, these animals are more active than during the drier months of the year. Females will produce one clutch per year (during the months from October to December), consisting of 4-15 elongated, thick-shelled eggs, which hatch after four to five months (ALVAREZ DEL TORO 1960, FircH 1970). Gila monsters are long-lived animals, reaching an age of over 20 years. Heloderma horridum (WixGMANN 1829), Isis von Oken 22: 421; type locality: Mexico. SVL to 325 mm. Southern Sonora to Chiapas, Mexico, also in Rio Motagua valley, Guatemala, sea level to 1000 m elevation. For subspecies sce text above, Further Reading Bocert & MARTIN DEL Campo 1956 (compre- hensive work on the biology of this genus). Fig. 163. Heloderma horridum horridum (Mixtequilla, Oaxaca, Mexico), Heloderma horridumIguanidae The phylogenetic relationships and taxon- omy of the pleurodont iguanians has been subject to considerable controversy in recent years, ETHERIDGE & Ds QUEIROZ (1988) demonstrated the existence of eight monophyletic groups within the family Iguanidae that were reclassified as eight families by Frost & Ersenipce (1989). The latter workers were unable to find any evi- dence for the monophyly of all pleurodont iguanians (the former family Iguanidae). Initially, this new classification was strongly criticized (BOHME 1990, LazeLi. 1992) and later rejected on the basis of new molecular data (MAczY et al. 1997, SCHULTE et al. 1998). Most recently, FRosT et al. (2001) have pres- ented a modified classification in which they recognized the pleurodont iguanians as a monophyletic group, based on the evi- dence provided by MAcey et al. (1997). Frost et al. (2001) proposed to maintain the subgroups as families and combine these in a higher level taxon (Pleurodonta). However, forthcoming work on this group of lizards is likely to result in additional changes (e.g. the former Tropiduridae and Polychrotidae were shown to be paraphyl- etic; see Frost et al. 2001), and there is a high probability that some of these groups will still be subject to change pending fur- ther studies. Because the current status of iguanian phylogenctics is still contentious, it appears best to combine all pleurodont iguanians in a single family (Iguanidae) and assign the status of subfamilies to the monophyletic subgroups. Key to Iguanidae genera 1 a Head with several horn-like projections; a lateral abdominal fringe with soft spines present (Fig. 325, p. 137)....... Phrynosoma b Head without horn-like projections, or, if head with horn-like projections, then ‘no lateral abdominal fringe with soft spines 2 2 Fingers and toes with widened smooth subdigital lamellae (Fig. 166d); males with an exiensile dewlap ...... anoles (genera Anolis, Ctenonotus, Dactyloa, Norops) Iguanidae Fig. 165, Adult male of Iguana i. rhinolopha (Caio Negro, Costa Rica). Photo: H, Maass b Lamellae beneath fingers and toes smooth or kecled but not widened (Fig. 166a-c); males without an extensile dewlap........3 3 a Head noticeably extended posteriorly, ¢ ther as vertical fin or horizontal shelf (hel- met, crown or fin; Fig. 168), most conspi uous in adult males i b Head not extended or drawn out posterio ly to form helmet, crown or fin 6 4 a Each subdigital lamella with a knob (Fig. 166a); horizontal fin (crown) on head Laemanctus 6. Anolis a, Laemanctus b. Basiliscus ¢. Scefoporus Fig. 166. Ventral surface of toes. 91Iguanidae 166b); vertical fin or helmet on head ......5 ‘ e ah 5 a Toes of hind foot with fringe of flat scales forming serrate margin (Fig. 166b); lateral b Subdigital lamellae without knobs (Fig. body scales homogeneous ........ Basiliseus ales zl | Besiiscus Corytophanes —_Laemanctus b Toes of hind foot without fringe of flat plumitrons enstatus serratus sales forming serrate margin; lateral body . scales heterogeneous Corytophanes Fig. 168. Lateral view of head. 6 a Tail with whorls of enlarged spinous scales | (Bigs. 169, 171) ses Tail without spines . e eos Bl 7 a A large, elongate subocular scale present, (Big. 169) srr nnr on Chenosaura b Orbit bordered below by an arc of several subequal scales . Morunasaurus 8 a Large pendulous dewlap present; one or two conspicuously enlarged rounded scales below tympanum; SVL to 50 em ... Iguana b No large pendulous dewlap; no conspic- uously enlarged scales below tympanum: small to medium-sized lizards Fig. 169. a. Scalation of tail in Ctenosaura quinquecarinata; b, Lateral head seales in Ctenosaura similis : . (enlarged subocular scale orange) 9 @ Very large interparictal plate present (Fig. WO b eantcsanctrss one, b Interparietal plate, if present, small (Fig. 170b).. : amma dt 10a A dorsolateral longitudinal fold alongside body present, usually with some enlarged scales; a complete transverse gular fold, differentiated from adjacent gular scales by very fine scales ... Urosaurus b No dorsolateral longitudinal fold alongside body; no transverse gular fold .... Sceloporus La Tail about three times SVL; no vertebral row of enlarged scales Polychrus b Tail less than twice SVL, a vertebral row of eo enlarged seales present ........ Enyalioides b. Polychrus Fig. 170. Dorsal head scalation (interparietal orange). Fig. 167. Urosaurus biearinatus (ventral view). Arrow points to gular fold Fig. 171. Tail of Morunasaurus annularis. 92Anoles (genera Anolis, Ctenonotus, Dactyloa, Norops) The anoles (Anolis sensu lato), with more than 300 valid species, represent the most diverse lizard genus, with more species being described almost every year. They are a conspicuous component of the fauna wherever they occur, and throughout their range it is common for several species of anoles to occur together. Despite the fact that anoles have been the focus of interest by herpetologists for more than a century, the phylogenetic relationships and taxon- omy of this group of lizards remains sub- ject to serious controversy in the recent literature. Indeed, our understanding of the systematics of these lizards is still poor, In the present work, I follow the clas- sification proposed by Guyer & Savace (1987, 1992) and SavaGE & GUYER (1989) in using the generic names Anolis, Cteno- notus, Dactyloa, and Norops for the anole species of Central America. This decision reflects the strong support for the assertion that Norops represents a monophyletic lineage. Norops is the beta section of anoles of ETHERIDGE (1959), whose species are characterized by autotomic caudal verte- brae with transverse processes that are directed anterolaterally; see figure 3A in EQHERIDGE (1967). However, the appro- priateness of GuYER and Savace's (1987, 1992) other proposed genera (the alpha group of EYHERIDGE 1959) appear more uncertain, The anole classification of Guyer & Savage (1987) was heavily criticized by CANNATELLA & DE QUEIROZ (1989) and by E. WILLIAMS (1989). For most “species groups” of anoles, as pro- posed by various rescarchers in the litera- ture, evidence that these actually repre- sent monophyletic groups is meager. Some of them probably are natural groups of spe- cies, while others represent artificial groupings of species that are superficially similar and are placed together because of particular character combinations, At this time, our knowledge of the phylogenetic Iguanidae relationships of the anoles is still too limited to accurately define the natural groupings. Most species are bush or tree-dwellers, al- though some are only found amongst the leaves on the ground. Even some semi- aquatic species that occur only in the viei- nity of water, such as Norops barkeri, N. lionontus, N. poecilopus, and N. aquaticus, are found in Central America. Typical leaf- dwellers in rain forest are some species of the ¢tropidonotus group (particularly N. humilis and N. uniformis), which in some areas reach a high population density (KOHLER 1998c), A more robust represent- ative of the genus is N. capito (SVL to 95 mm). These lizards are typical sit-and- wait predators, which are difficult to spot when sitting on a branch or tree trunk. Similarly large (to about 100 mm SVL), is the bright green colored N. biporcatus that inhabits the tropical rain forest, usually on trees at the forest edge, in clearings and at the shore of ponds and lakes. The true giants, however, amongst the Central American anoles are found in the genus Dactyloa, such as D. insignis (SVL to 160 mm). These are tree-dwellers that are dif- ficult to spot in the rain forest. With luck, they can be observed on the trunks of large trees where the animals usually sit with their heads pointed down. According to Scorr et al. (1976), these animals usual- ly remain on tree trunks at heights be- tween 40 and 200 cm above the ground, although specimens sitting up higher would be difficult to observe (Savace & TaLBor 1978). The species of the fuscoau- ratus group (e.g., Norops altae, N. exsul, N. fuscoauratus, N. kemptoni, N. limifrons, N. pandoensis, N. polylepis, N. rodriguezii, and N. zeus) are slenderly built anoles, which live mostly in bushes or low trees, where they are sometimes difficult to spot. They are predominantly lowland species, usually living in border areas and clear- ings in the rain forest, where they tend to be quite plentiful. The species of the cras- sulus group (e.g., N. amplisquamosus, N. anisolepis, N. crassulus, N. haguei, N. 9394 Iguanidae heteropholidotus, N. laeviventris, N. mural- la, N. rubribarbaris, N. sminthus, and N. wermuthi) and the schiedei group (c.g., N. cobanensis, N. hobartsmithi, N. johnmeyeri, N. pijolensis, and N. purpurgularis) are mostly highland species (above 1000 m ele- vation), usually with small areas of distri- bution in cloud and pine forest. It is be- lieved that the species of the schiedei group originated from the fragmentation of what was once the continuous distribution area of an ancestor species due to the uplift of mountain massifs during the Tertiary peri- od (McCRanIE et al, 1993b). All anole species reproduce by laying eggs, and, during the breeding season, will pro- duce several clutches, each consisting of a single egg, at intervals of one to two weeks. The ovary and oviduct on each side of the body are alternately active. As soon as one egg is laid from the right oviduct, for example, a new follicle is produced by the left ovary and passes through the left ovi- duct, where it is fertilized and shelled within a few days, before being laid. In the meantime, the right ovary has produced a new follicle, which is ready to ovulate. As long as there is enough food available and the weather conditions are suitable, this cycle will continue and eggs will be laid at regular intervals (FrrcH 1982). In the following list, information for body and dewlap coloration are for live speci- mens; except where otherwise indicated, dewlap coloration pertains only to adult males. Norops binotatus (PETERS 1863b) is excluded because there appears to be no valid record for this species in Central America. BARBOUR (1923) erroneously reported this taxon for Panama, based on misidentified N. vittigerus (KOHLER unpubl. data). Anolis allisoni Barbour 1928, Proc. New England Zool. Club 10: 58; type locality: Coxen Hole, Ruatan, Bay Islands, Honduras, SVL to 87 mm. Body green, dewlap pink. Islas Guanaja and Roatan, Cayos Cochinos, Honduras, as well as in Belize and on Cuba, sea level to 300 m elevation. Ctenonotus cristatellus (DuMéRIL & BIBRON 1837), Erp. Gén. 4: 143; type locality: Martinique [in error]. SVL to 70 mm. Body brown, dewlap greenish yellow with orange margin. Puerto Rico and Virgin Islands; introduced in Costa Rica, sea level to 750 m elevation; occurrence on Isla Cozumel doubt- fal (LEE 1996). Dactyloa casildae (AROSEMENA, IBANEZ & DE Sousa 1992, Rev. Biol. Trop. 89: 255-262; type locality: Margenes de Quebrada Frank, 1100 m, 8°44°N, §2°13°W, Reserva Forestal de Fortuna, Chiriqui, Panama, SVL to 108 mm, Body green with 4-6 oblique brown bands (not formed of adjacent ocelli) interspersed by blue-outlined yellow patches; some females have a broad, dark brown mid-dorsal stripe extending from nape to base of tail; dewlap dirty cream with broad yellow scale rows irregularly interspersed with smaller eme- rald green scales. Cordillera Central in the Provinces Chiriqui and Bocas del Toro, Panama, 1050-1400 m elevation. Dactyloa chloris (BOULENGER 1898), Proc. Zool. Soc. London 1898: 110; type locality: Paramba, Ecuador, SVL to 60 mm. Body green, dewlap yellowish white, pale blue anteriorly, Eastern Panama to northwestern Ecuador. Dactyloa chocorum (WiL1AMS & DUBLLMAN 1967), Breviora, Mus. Comp. Zool. 256: 2; type locality: Rio Tuira at Quebrada La Plata, 100 m, Darién, Panama. SVL to 79 mm in males, to 73 mm in females. Body green with or without darker green oblique bands; dewlap pale orange with green lines. Southern Costa Rica to Colombia, 100-1100 m elevation. Dactyloa frenata (Cope 1899), Sci, Bull. Philadelphia Mus. 1: 6; type locality: unknown. SVL to 143 mm in males, to 118 mm in fe- males. Body green with several dark green ocelli (most outlined with yellow) that are close- ly arranged and form dark oblique bands; dewlap uniformely cream-colored (occasional- ly with black shading). Costa Rica to north- ern Colombia, sea level to 820 m elevation in lowland and premontane rain forests. Dactyloa insignis (Core 1871), Proc. Acad. Nat. Sci. Philadelphia 24: 213; type locality: San José, Costa Rica. SVL to 160 mm in males, to 135 mm in females. Body yellowishFig. 172. Anolis allisoni (Isla de Roatan, Fig. 176. Dactyloa chocorum (Cerro Sapo, Honduras). Darién, Panama). Photo: C, W. Myers Fig. 173. Ctenonotus cristatellus Photo: R. D. Bartlett Photo: A. Plischendriger Fig. 174. Dactyloa casildae (Fortuna, Panama). Photo: K. Fig. 175. Dactyloa chloris (Rio Jaque, Darién, _Fig. 178. Dactyloa insignis (Las Cruces, Punta. Panama). Photo: C. W. Myers —_renas, Costa Rica). Photo: K.-H. Jungfer96 Iguanidae == gray to tan or greenish with a distinct large, pale-centered, dark-outlined blotch above shoulder; dewiap orange red, with or without several horizontal green or white bars and dark spotting and with a white to greenish free margin. Costa Rica and Panama, sea level to 1600 m elevation Dactyloa latifrons (BertHoLD 1845), Nachr. Georg-Augusts Univ. und K. Ges. Wiss. Gottingen 3: 39; type locality: western Colombia (see Myers & BouME 1996). SVL to 124 mm in males, to 97 mm in females. Body green with a distinct large, pale-centcred, dark-outlined blotch above shoulder; dewlap pale greenish gray with pale yellow scales. Eastern Panama to northwestern Ecuador. Dactyloa microtus (Core 1871), Proc. Acad. Nat. Sci. Philadelphia 24: 214; type locality: San José, Costa Rica. SVL to 111 mm in males, to 104 mm in females. Body yellowish gray to tan with several pale brown-centered dark bands that may change to green on flanks and a distinct postorbital dark-bor- dered pale stripe; dewlap salmon with white scales. Costa Rica and Panama, 425-1500 m elevation, Norops altae (Dunn 1930b), Proc. New England Zool, Club 12: 22; type locality: Finca ‘Acosta, 7000 ft, Volean Barba, Costa Rica. SVL to 52 mm. Body grayish brown, dewlap rust orange grading to yellow on free margin. Highlands of Costa Rica, 1220-2000 m eleva- tion. Fig. 179. Dactyloa latifrons (near Noanamé, Darién, Panama). Photo: C. W, Myers Norops alvarezdeltoroi (Niro MONTES DE Oca 1996), J. Herpetol. 80: 20; type locality: 19,5 km N, 8,1 km W Ocozocoautla, 940 m, 16°56'N, 97°27°W, Chiapas, Mexico. SVL to 66 mm, Body grayish brown, dewlap dark red with a suffusion of black pigment. Region of EI Ocoto, Chiapas, Mexico, 910-940 m eleva- tion Norops amplisquamosus McCrantr, WIsoN & WiLLiams 1992, Carib, J. Sci. 28: 209; type locality: Finca El Cusuco, 1550 m, 15 31'N, 88 12'W, 5,6 km WSW Buenos Aires, Sierra de Omoa, Cortés, Honduras. SVL to 46 mm. Body grayish brown, dewlap yellow-orange. Only known from the type locality, 1530-1720 m elevation. Norops anisolepis (Smrra, BuRLEY & FRITS 1968), J. Herpetol. 2: 147; type locality: 10 mi SE San Christébal Las Casas, Chiapas, Mexico. SVL to 47 mm. Body grayish brown, dewlap pinkish red. Meseta Central de Chiapas, Mexico, 1500-2500 m elevation. Norops aquaticus (TavLor 1956), Univ. Kansas Sci. Bull. 38: 141; type locality: Palmar, Puntarenas, Costa Rica. SVL to 71 mm, Body grayish yellowish green with broad transverse brown bands and a pale greenish yellow lateral longitudinal stripe; dewlap red-orange with yellow stripes. Costa Rica and Panama, 30-1170 m elevation. Norops auratus (DAUDIN 1802), Hist. Nat. Rept.: 89; type locality: unknown. SVL to 52 mm, Body grayish brown, dewlap blue. Central Panama and northern South America, sea level to 2000 m elevation. Norops barkeri (Scuntpr 1939), Field Mus. Nat. Hist. Publ. Zool. Ser. 24: 7; type locality: Cascajal, upper Uzpanapa River, Veracruz, Mexico, SVL to 101 mm. Body grayish brown, dewlap dark orange with yellowish orange margin. Caribbean versant of Isthmus of Tehuantepec (Veracruz, Oaxaca, Tabasco, and Chiapas, Mexico), 200-1100 m elevation. Norops bicaorum Konter 1996b, Sencken- bergiana biol. 76: 21; type locality: trail to Rock Harbour, about 3 km north of the town of Utila (16° 6,34’N; 86° 53,94°W), Isla de Utila, Islas de la Bahia, Honduras. SVL to 75 mm. Body grayish brown, dewlap orange with suffusion of black pigment centrally and with white scales, Endemic on Isla de Utila, Islas de la Bahia, Honduras, sea level to 100 m ele- vation. Norops biporcatus (WigoMaNN 1834), Herp. Mex. 47; type locality: Santa Rosa de Pansos [=Panzos], Alta Verapaz, Guatemala. SVL to 102 mm, Body green (capable of rapid colorFig. 184. Ne tus (Canjilones. Gualaca, Chiriqui, Pana Photo: M. Lundberg . 181. Norops amplisquamosus (Cusuco National Park, Honduras). Fig. 186. Norops bicaorum (Isla de Uti Hondur Norops aquaticus (Las C: Punta- Fig. 187 ‘osta Rica) Photo: K.-H. Jungfer — Niearagua).Iguanidae change to brown), dewlap blue with dark orange margin, Southern Mexico through Central Ameriea to northern South America, sea level to 2000 m elevation. The nominate subspecies occurs in Central America. KOHLER & Baur (2001) demonstrated that the holotype of Norops biporcatus (WiEGMANN 1834) is conspecific with the syntypes of N. petersii (BOCOURT 1873). In order to stabilize the current and long established usage of the names N. biporcatus and N. petersii, the International Commission on Zoological Nomenclature (ICZN) set aside the status of the original holotype of N. biporcatus and designated MNHM. 2426 (holotype of Anolis cope: BocouRT 1873) as the neotype of N. biporcatus (International Commission on Zoological Nomenclature 2002). Norops capito (Peres 1863b), Monats. Akad. Wiss, Berlin 1863: 142; type locality: Costa Rica. SVL to 96 mm. Body grayish brown with green elements, dewlap yellowish green, Caribbean versant of southern Mexico through Central America, sea level to 1300 m elevation. Norops carpenteri (ECHELLE, ECHELLE & Finch 1971), Herpetologica 27: 355; type lo- cality: Rio Reventazon, 7 km ESE Turrialba, Cartago, Costa Rica. SVL to 45 mm. Body greenish, dewlap orange. Nicaragua, Costa Rica, and Panama, sea level to 1100 m eleva- tion. Anolis procellaris MYERS 1971a_ is regarded to be a synonym of N. carpenteri according to Firci et al. (1976). Norops cobanensis (STUART 1942), Oce. Pap. Mus. Zool. Michigan 464: 6; type locality: 3 km S Finca Samac (6 airline km of Coban), 1350 m, Alta Verapaz, Guatemala, SVL to 50 mm. Body grayish brown, dewlap purple. Highlands on the Caribbean versant of Guatemala, 1300-1410 m elevation Norops compressicauda (Smurn & Kersver 1955), Herpetologica 11; 193; type locality: Near La Gloria, Oaxaca, Mexico. SVL to 55 Fig. 188. Norops eristifer (Depto. Retalhuleu, Guatemala). Photo: L. Melendez mm, Body grayish brown, dewlap uniform rose to purple. Disjunct populations in eastern Oaxaca and western Chiapas, Mexico, 500-1200 m elevation Norops concolor (Cope 1862b), Proc. Acad. Nat. Sci. Philadelphia 14: 180; type localit Nicaragua. SVL to 45 mm. Islas Providencia and San Andres, Colombia (off the Caribbean coast of Nicaragua). Norops crassulus (Copp. 1864), Proc. Acad. Nat. Sci. Philadelphia 16: 173; type locality: Cobiin, Alta Verapaz, Guatemala. SVL to 59 mm, Body grayish brown, dewlap red. Highlands of northern Central America (southern Mexico, Guatemala, Honduras, and El Salvador), 1300-3000 m elevation. Norops cristifer (SmitHt 1968a), Trans. Kansas Acad. Sci. 71: 195; type locality: near Acacoyagua, Chiapas, Mexico. SVL to 88 mm. Body grayish brown. Pacific versant of Chiapas, Mexico, and western Guatemala, Norops cupreus (HALLOWELL 1861), Proc. ‘Acad. Nat. Sci, Philadelphia 12: 481; type locality: Nicaragua, SVL to 57 mm. Body grayish brown, dewlap brown with orange margin. Honduras to Costa Rica, sea level to 1250 m elevation. Norops cuprinus (Surrit 1964), Herpetologica 20: 31; type locality: Vicinity of Zanatepec, Oaxaca, Mexico, SVL to 69 mm. Body grayish brown, dewlap red. Eastern Oaxaca and western Chiapas, Mexico, 200-500 m eleva- tion. Norops cusuco McCRanie, KOHLER & WILSON 2000, Senckenbergiana biol. 80: 214; type locality: Parque Nacional El Cusuco Centro de Visitantes, 15°29.92°N, 88°12.88°W, 1550 m elevation, Cortés, Honduras. SVL to 46 mm: Body grayish brown, dewlap unicolor dirty white. Known only from vicinity of type lo- cality, 1550-1935 m. Norops dollfusianus (BocourT 1873), Miss, Sci. Mex., Rept. 1873: 84; type locality: Voledn Atitlén, 1200 m, Guatemala. SVL to 44 mm. Body grayish brown, dewlap yellow. Pacific versant of eastern ‘Chiapas, Mexico, and western Guatemala, 200-1200 m elevation. Norops exsul (AROSEMENA & IbaNt2 D. 1994), Carib, 1. Sci. 80: 222; type locality: “orillas del Rio Chiriqui, a unos 100 m de la desemboca- dura de la Quebrada Bijau (8°45'11"N, 82°11'07"W), entre 1050-1075 m sobre nivel del mar” Reserva Forestal de Fortuna, Chiriqui, Panama. SVL to 48 mm. Body gray- ish brown, dewlap orange red with yellow seales. Chiriqui, Panama, 1050-1075 m eleva- tionFig. 189. Norops capito (Bartola Nicaragua). ig. 190. N Ju Fig. 191. Norop lorops compressicauda region, Chiapas, I Fig. 194, Norops cupreus (Selva Negra, Matagalpa, Nicaragua Fig. 196. Norops dollfusianus (Reserva Natui Suchitepéquez, Guatemala).100 Iguanidae Norops fortunensis (Anosema & Inanez, D. 1993), Rev. Biol. Trop. 41: 267; type locality: Reserva Forestal de Fortuna, along the Rio Chiriqui, in front of the estacion hidromete- reolégica del Instituto de Recursos Hidraulicos y Electrificacién (IRHE) in Bijau, 1050-1075 m elevation, Chiriqui Province, Panama. SVL to 48 mm. Body grayish brown, dewlap orange red with red ‘margin and yel- low scales. Cordillera Central, Chiriqué Province, Panama, 1050-1075 m elevation. Norops fungosus (Myeks 1971b), Amer. Mus. Novitat. 2471; 13; type locality: north slope Cerro Pando, 1450 m, Cordillera de Talamanea, Bocas del Toro, Panama. SVL to 47 mm. Body grayish brown, dewlap red. Cordillera de Talamanca in Panama and Costa Rica, 1200-1600 m elevation, Norops fuscoauratus (D'ORBIGNY 1837), in DUuMERIL & BIBRON: Erp. Gén. 4: 110; type locality: Chile [in error]. SVL to 50 mm. Body grayish brown, dewlap pink to pinkish- brown, Eastern’ Panama; widely distributed in the Amazon Basin, sea level to 1000 m ele- vation. Norops haguei (STUART 1942b), Oce. Pap. Mus. Zool. Univ. Michigan 464: 3; type local- ity: 2 km south of Finea Chichén, 1750 m, Alta Verapaz, Guatemala. SVL to 53 mm. Body grayish brown, dewlap red, Only known from type. Norops heteropholidotus (Mextens 1952a), Zool. Anz, 148: 89; type locality: Hacienda Los Planes, 2000 m, Miramundo, Santa Ana, El Salvador. SVL to 59 mm. Body grayish brown, dewlap red with pale to dark brown scales. Highlands of northern El Salvador and south- western Honduras, 1870-2200 m elevation. Norops hobartsmithi (NieT0-MONTES DE OCA 1995), Publ. Espec. Mus. Zool. Facultad de Ciencias, UNAM, 10: 160; type locality: Parador Selva Negra on México Highway 195 (approx. 0.5 km W Pinabeto), 1992 m, Fig. 197. Norops fuscoauratus (Arutam, Pastaza, Ecuador), 17°12'58"N, 92°57'47"W, Municipality of Rayon, at the border with Municipality of Pueblo Nuevo Solistahuacén, Chiapas, Mexico. SVL to 49 mm. Body grayish brown, dewlap rose. Highlands of northwestern Chiapas, Mexico, 1525-2000 m elevation. Norops humilis (Pevexs 1863b), Monats. Akad. Wiss, Berlin 1863: 138; type localit Veragua, Panama. SVL to 44 mm. Body gray- ish brown, dewlap red-orange with yellow margin or dewlap uniform red orange (popu- lations in eastern Panama). Northern Costa Rica along both versants to central Panama east of the Canal Zone, sea level to 1600 m elevation, Norops johnmeyeri (WILSON & McCRANIB 1982), Trans. Kansas Acad. Sci. 85: 133; type locality: 5.6 km WSW Buenos Aires, 1580 m, E] Cusuco, Cortes, Honduras. SVL to 70 mm. Body grayish brown, dewlap orange red with large central blue blotch. Females with well developed dewlap (yellow with large central blue blotch). Sierra de Omoa (Dptos. Copan and Cortés), Honduras, 1410-1995 m eleva- tion. Norops kemptoni (DuNN 1940a), Proe. Acad. Nat. Sci. Philadelphia 92: 111; type locality: Finea Lérida, Chiriqui, Panama. SVL to 55 mm, Body grayish brown, dewlap red with orange blotch anteriorly, and with white scales, Highlands of Chiriqui, Panama, 1000- 1800 m elevation. Norops kreutzi McCrante, Kouter & Wison 2000, Senckenbergiana biol. 80: 218; type locality: 2.5 airline km NNE La Fortuna, 15°26'N, 87°18'W, 1670-1690 m elevation, ‘Yoro, Honduras. SVL to 51 mm, Body grayish brown, dewlap pale yellow with purple scales, Known only from the type locality. Norops laeviventris (WIRGMANN — 1834), Herpetologica Mexicana: 47; type locality: Mexico. SVL to 52 mm. Body grayish brown, dewlap dirty white. Southern Mexico to western Panama, 1160-2000 m. elevation. Based on the studies of Borrcnes (1999) and KONLER (2001e) I consider Anolis intermedius PETERS 1863b, Anolis nannodes Cops 1864, and Anolis bourgeaei BOcoURT 1873 to be synonyms of N. laeviventris. Norops lemurinus (Cope 18614), J. Acad. Nat. Sci. Philadelphia 12: 213; type locality: Veragua, Panama, SVL to 75mm, Body gray- ish brown, dewlap red (often with black scales). Central Veracruz, Mexico, along Caribbean versant of Central America to cen- tral Panama; also on Pacific versant in Costa Rica and Panama, sea level to 1080 m eleva- tion,Fig. 202. Norops laevive Honduras). pa, Nici Fig, 200. Norops johnmeyeri (Cusuco National 8 Park, Cortés, Honduras). Atlantieo Norte, Nica ee eid Se att Yorops hemptoni (north of Boquete, : cops lion hiriqui, Panama). Photo: T. Bille pga, Nicaragua). “rons (Cerro Sa gua). us (near Pueblo Wiso,102 Iguanidae Norops limifrons (Core 1862b), Proc. Acad. Nat. Sci, Philadelphia 13: 178; type locality: Veragua, Panama. SVL to 48 mm. Body gray- ish brown, dewlap dirty white with a basal orange-yellow spot or (populations in central and eastern Panama) dewlap almost com- pletely orange-yellow. Eastern Honduras to Panama, sea level to 1350 m elevation. Anolis biscutiger TavLor 1956 and Anolis godmani BOULENGER 1885 are both regarded to be synonyms of N. limifrons according to SAVAGE (2002). Norops lionotus (Cops 18614), J. Acad. Nat. Sci. Philadelphia 12: 210; type locality: Cucuyas de Veragua, Panama. SVL to 78 mm. Body grayish brown, dewlap yellow to yellow- orange. Eastern Honduras to central Panama, sea level to 1200 m elevation. Based on the studies of OnERMEMER (1998), I consider Norops oxylophus CPE 1876 to be a synonym of N. lionotus. Norops loveridgei (ScuMIDT 1936a), Proc. Biol. Soc. Washington 49: 47; type locality: Portillo Grande, 4100 ft, Yoro, Honduras. SVL to 118 mm, Body grayish brown, dewlap or- ange with pink streaks. Mountainous regions in the Dptos. de Yoro and Atlantida, Honduras, 550-1100 m elevation. Norops macrophallus (Wennex 1917), Mitt. Zool. Mus. Hamburg 34: 31; type locality: San José, Guatemala. SVL to 45 mm. Body gray- ish brown, dewlap rose with a orange yellow basal bloteh. Pacific versant of northern Central America (Guatemala and El Salvador), 40-1320 m elevation. Norops matudai (Sr 1956), Herpetologica 12: 1; type locality: region ‘of Soconuseo, Chiapas, Mexico, SVL to 42 mm, Body gray- brown, dewlap purple. Sierra Madre de Chiapas, Chiapas, Mexico, and adjacent high- lands in ‘Guatemala, 200-1000 m elevation. Norops muralla Koutrn, McCRanin & Wisow 1999, Amphibia—Reptilia 20: 285; type locality: along trail to Cerro de Enmedio, 1500 m, Parque Nacional La Muralla, Olancho, Honduras. SVL to 56 mm. Body grayish brown, dewlap red. Parque Nacional La Muralla, Honduras, 1440-1740 m elevation. Norops ocelloscapularis Konuir, McCRANI® & WILSON 2001a, Herpetologica 57: 248; type locality: near Quebrada Grande off a trail to Laguna del Cerro, 15°04.82'N, 88°55.45°W, 1200 m elevation, Copan, Honduras. SVL to 47 mm, Body grayish brown, dewlap orange. Known only from vicinity of type locality, 1150-1370 m elevation. Norops pachypus (Core 1876), J. Acad. Nat. Sci. Philadelphia (2) 8: 122; type locality: Slopes of Pico Blanco, Costa Rica, SVL to 50 mm, Body grayish brown, dewlap red-orange Highlands of Costa Rica and Voleén Chiriqui, Panama, 1370-1560 m elevation. Norops pandoensis SAVAGE & GUYER 1998, Rey. Biol. ‘Trop. 46: 805; type locality: La Changuinolita, 1950 m elevation, Cant6n Coto Brus, Puntarenas Province, Costa Rica. SVL to 60 mm. Body grayish brown to olive green, dewlap pink with orange anterior mar- gin. Cordillera Talamanca on both sides near the border between Costa Rica and Panama, 1350 to 1960 m elevation. Norops parvicireulatus (ALVAREZ DEL TORO & Sari 1956), Herpetologica 12: 5; type local- ity: El Suspiro, 1200 m, Chiapas, Mexieo. SVL to 50 mm. Body grayish brown, dewlap yellowish orange with a red central blotch. Central-eastern Chiapas, México, 500-1200 m elevation. Norops pentaprion (Core, 1862b), Proc. Acad. ‘Nat. Sci. Philadelphia 14: 178; type locality: ‘Truando River, Colombia, SVL to 79 mm. Body grayish brown, dewlap red. Along the Caribbean versant of Central America from Tamaulipas, Mexico, to Panama, including Yucatan Peninsula; also on Pacific versant in Costa Rica and Panama, sea level to 900 m elevation (to 1780 m in Veracruz, Mexico, according to Voor et al. 1997). Two subspe- cies are recognized: Norops p. pentaprion (Caribbean versant of Central America from eastern Honduras to Panama) and Norops p. beckeri (BOULENGER 1881) (Caribbean ver- sant from Tamaulipas, Mexico, to Honduras; including Yueatan Peninsula). Norops.petersii (BocouRT 1873), Miss. Sei Mex., Rept.: 79; type locality: Alta Verapaz, Guatemala. SVL to 102 mm. Body greenish, dewlap purple with pale yellow margin and pale yellow gorgetal scales. San Luis Potosi, Mexico, to Honduras, 200-2130 m elevation. Norops pijolensis McCraxis, Wuson & WILLIAMS 1993b, J. Herpetol. 27: 393; type locality: Bast slope of Pico Pijol, 2050 m, 15°10'N, 87°33'W, Montafa de Pijol north- west of Tegucigalpa, Yoro, Honduras, 1860- 2050 m elevation. SVL to 60 mm. Body gray- ish brown, dewlap rose with pink scales. Only known from the type locality. Note that. the species name has been emended from the original spelling ‘pijolense” to “pijolensis” to correspond to the masculine gender of the genus name Norops (following H. Sura & LARSEN 1974a).104 Iguanidae Norops poecilopus (Cope 1862b), Proc. Acad. Nat. Sei Philadelphia 14: 179; type locality: near Cartagena and on Rio Truando, Colombia. SVL to 68 mm. Body grayish brown, dewlap yellow to yellowish orange. Panama east of the Canal Zone and Colombia, sea level to 1000 m elevation. Norops polylepis (Pevers 1873), Monats. Akad, Wiss. Berlin 1873: 738; type locality: Chiriqui, Panama. SVL to 57 mm. Body gray- ish brown, dewlap orange. Southwestern Costa Rica and Panama, sea level to 1330 m elevation. Norops purpurgularis McCrante, Cruz & HoLM 1993a, J. Herpetol. 27: 386; type local- ity: 2.5 km airline NNE La Fortuna, 1690 m, 15 26'N, 87 18'W, Cordillera Nombre de Dios, Yoro, Honduras. SVL to 60 mm, Body grayish brown, dewlap pink. Mountainous regions in the Dptos, de Yoro and Atlantida, Honduras, 1690-2040 m elevation. Norops pygmacus (ALVAREZ DEL ToRO & SMITH 1956), Herpetologica 12: 7; type locality: El Ocote, Chiapas, Mexico, 600 m. SVL to 35 mm, Body grayish brown, dewlap pale red. Region of El Ocote, Chiapas, and south- eastern Oaxaca, Mexico, Norops quaggulus (Cope 1885), Proc. Amer. Phil. Soc, 22: 391; type locality: Rio San Juan, Niearagua, Body grayish brown, dewlap red” orange with yellow margin, On the Caribbean versant from eastern Honduras through Nicaragua to extreme northern Costa Rica, sea level to 1350 m elevation. Norops roatanensis Kowen & McCranip 2001, Senckenbergiana biol. 81: 240; type loeality: between West End Point and Flowers Bay, 16°17.98'N, 86°34.82'W, 30 m elevation, Abb. 214, Norops purpurgularis (2.5 airline km NNE of La Fortuna, Yoro, Honduras). Photo: J. R. MeCranie Isla de Roatan, Islas de la Bahia, Honduras. SVL to 63 mm. Body grayish brown, dewlap pink red with suffusion of black pigment cen- trally and with white scales. Endemic to Isla de Roatan, Islas de la Bahia, Honduras. Norops rodriguezii (BocoURT 1873), Miss. Sei. Mex., Rept.: 62; type locality: Panzos, Guatemala. SVL to 49 mm. Body grayish brown, dewlap orange with darker’ basal area. Isthmus of Tehuantepec, Mexico, to western Honduras, sea level to 2000 m eleva- tion Norops rubribarbaris Konner, McCrante & Wison 1999, Amphibia-Reptilia 20: 280; type locality: 4 km S of San Luis de los Planes, 1700 m elevation, N slope of Montafa de Sania Barbara, Santa’ Barbara, Honduras. SVL to 48 mm. Body grayish brown, dewlap red. Only known from the type locality. Norops sagrei (Duméntt. & Biskon 1837), Exp. Gén. 4: 149; type locality: Cuba. SVL to 57 mm. Body grayish brown, dewlap yellow- orange to red-orange with yellow margin. Southeastern Mexico to Honduras, sea level to 200 m elevation. In Central America, the subspecies Norops s. mayensis Smith & BURGER 1949 occurs. Norops sericeus (HALLOWELL 1856), Proc. Acad. Nat. Sci. Philadelphia 8: 227; type local- ity: Jalapa [= Xalapal, Veracruz, Mexico. SVL to 52 mm. Body grayish brown, dewlap yel- low-orange with central blue blotch. Isthmus of Tehuantepec on the Pacific versant and Tamaulipas, Mexico, on the Caribbean ver- sant along both versants to Costa Rica, sea level to 1500 m elevation. Norops serranoi KOHLER 1999a, Salamandra 35: 39; type locality: forest in the vicinity of butterfly farm of Dr. Francisco Serrano (13°49,46'N, 89°59,98'W), 225 m elevation, Departamento Ahuachapan, El Salvador. SVL to 85 mm. Body grayish’ brown, dewlap brick red with suffusion of black pigment cen- trally and with white scales. Pacific versant of northern Central Ameriea from Chiapas, Mexico, to eastern El Salvador, sea level to 950 m elevation. Norops sminthus (DUNN & EMLEN 1932), Proc. Acad. Nat. Sei, Philadelphia 84: 26; type locality: San Juancito, 6900 ft, Honduras. SVL to 55 mm. Body grayish brown, dewlap red with pale to dark brown scales. South- central Honduras, 1840-2100 m elevation. Norops townsendi (StBINEGER 1900), Bull. Mus. Comp. Zool. 36: 163; Isla del Coco, Costa Rica. SVL to 49 mm, Body grayish brown with’ a lateral stripe bordered above andFig. 219. Norops sericeus (Rancho Alegre, Chiap; Fig. 216. N Hondur ande, Copén, Hondur Mexi Norops sminthus (La Tigra, Francisco Photo: M. Lundberg ter (Cordoba, Photo: M. Lundberg106 Iguanidae below by a black stripe; dewlap dark brown. Isla del Coco, Costa Rica, sea level to 630 m elevation, Norops tropidogaster (HALLOWBLI. 1856), Proc. Acad. Nat. Sci. Philadelphia 8: 224; type locality: Colombia, SVL to 65 mm. Body gray- ish brown, dewlap red-orange with white seales, Eastern Panama and northern South ‘America, sca level to 700 m elevation. Norops tropidolepia (BOULENGER 1885), Cat. Lizards Brit. Mus. 2"4 ed. 2: 63; type locality: Voledn Irazu, Costa Rica. SVL to 59 mm, Body grayish brown, dewlap purplish red to pink. Voledn Irazu ‘and Monteverde, Costa Rica, 1220-2500 m elevation. Norops tropidonotus (PeteRs 1863b), Monats. Akad. Wiss. Berlin 1863: 135; type locality: Huanusco, Veracruz, Mexico. SVL to 60 mm. Body grayish brown, dewlap dark orange with yellow to yellowish orange margin and a central dark streak. Southern Mexico to northern Nicaragua, 90-1740 m elevation. Norops uniformis (Cope 1885), Proc. Amer, Phil. Soc. 22: 392; type locality: Yucatan, Mexico. SVL to 40 mm. Body grayish brown, dewlap rose with purple spots. Southern Mexico to western Honduras, sea level to 1300 m elevation. Norops utilensis KOHLER 19962, Sencken- bergiana biol. 75: 24; type locality: 2 km NNE of the town of Utila, Isla de Utila, Islas de la Bahia, Honduras, SVL to 59 mm. Body gray- ish brown, dewlap red with white scales. Isla de Utila, Honduras. Norops villai (Frrcn & Hennerson 1976), Milwaukee Pub. Mus, Contrib. Biol. Geol. 9:1; type locality: Great Corn Island, Nicaragua. SVL to 55 mm. Body grayish brown, dewlap poem elk bas? Fig. 223. Norops fungosus (north slope of Cerro Pando, 1450 m, Bocas del Toro, Panama), Photo: C. W. Myers chocolate Great Corn Island, Nicaragua, brown. Norops vittigerus (Core 1862b), Proc. Acad. Nat. Sci. Philadelphia 14: 179; type locality: ‘Truando region, Colombia, SVL to 72 mm. Body grayish brown, dewlap red-orange with a dark blotch. Eastern Panama and northern South America, sea level to 900 m elevation. Norops vociferans (MYERS 1971b), Amer. Mus. Novitat. 2471: 10; type locality: 4 km W Cerro Punta, 1829 m, Chiriqui, Panama. SVL to 64 mm. Body grayish brown, dewlap red with white scales. Costa Rica and Panama, 1400- 1830 m elevation. Norops wampuensis McCraxie & Komen 2001, Senekenbergiana biol., Frankfurt a. M., 81 (1/2): 228; type locality: confluence of Rios Aner and Wampd, 15°03'N, 85°07'W, 110 m elevation, Olancho, Honduras. SVL to 51 mm. Body grayish brown, dewlap orange-red with yellow margin. Known only from the type locality, 95-110 m elevation Norops wermuthi Konuer & OBERMErER 1998, Senckenbergiana biol. 77: 129; type locality: road from Matagalpa to Jinotega at km 146 (13°01,995'N, 85°55.848'W), 1400 m elevati- on, Departamento Jinotega, Nicaragua. SVL to 54 mm. Body grayish brown, dewlap red with pale to dark brown scales. Northern Nicaragua, 1230-1500 m elevation Norops woodi (DUNN 1940a), Proc. Acad. Nat. Sci. Philadelphia 92: 110; type locality: El Voledn, Chiriqui, Panama. SVL to 100 mm. Body grayish brown, dewlap dark-orange, brownish olive or black (capable of color change). Puntarenas Province, Costa Rica, to Chiriqui, Panama, 1150-2500 m_ elevation. Anolis aitenuatus Tavor 1956 is regarded to be a synonym of N. woodi according to SAvace (2002). Norops yoroensis McCranir, NICHOLSON & KOnLeR 2002, Amphibia-Reptilia 22: 466; type locality: 2.5 airline km NNE La Fortuna, 1600 m elevation, 15°26'N, 87°18'W, Cordillera Nombre de Dios, Yoro, Honduras. SVL to 47 mm. Body grayish brown, dewlap unicolor orange. Northwestern Honduras (Yoro), 1180-1600 m elevation Norops zeus Kouuer & McCranié 2001, Senckenbergiana biol. 81; 236; type locality Liberia, Parque Nacional Pico Bonito, 90 m elevation, Atlantida, Honduras. SVL to 44 mm, Body grayish brown, dewlap unicolor dirty white. Caribbean yersant of the Cordillera Nombre de Dios in northern Honduras, 90-900 m elevation,is (Reserva St Fig, Norops vittigerus (Cordoba ‘osta Rica), Photo: T. Bille Colombia) Photo: M. tropidono all p iropidonotus (female; Rancho Mexico), Alegre, Chiapas, Mexico), s Norops uniformis ( R Fig. 230. Ni vermuthi (Cerro Bl Toro. Chiapas, Mexico). ua), pera at sea Tega eeIguanidae Norops barkeri MiNorops capito 4 Anolis allisoni © Cienonotus cristatellus Dactyloa insignis © Dactyloa microtus © Dactyioa latirons © Dactyloa chioris 4 Dactyloa chocorum ¥ Dactyloa casildae Norops 4/ Nv, alvarezdettoroi 1 N. hobartsmithi Y NV, cobanensis © 1. johnmeyeri 4. pijotensis VN. purpurgularis ON. overidget GN. sagrei Norops 1 N. bicaorum Y/N. roatanensis, (EN. lemurinus EN. serranoi EN. vittigerus 108 [BNorops biporcatus IB Dactyloa frenata ‘© ‘Norops exsul 4 Norops fortunensis: 1B Norops kemptoni 4 Norops pandoensis Y Norops altae © Norops auratus Norops 4.N. crassulus NN. haguei YN. anisolepis ©. N. amplisquamosus AN. rubribarbaris ON. parvicirculatus ON. matudai IBN. rodriquezit ON, yoroensis © N. cuprinus YN. villai 4 N. dolifusianus WN. cupreus IBN, macrophallusIguanidae Norops Norops © W. zeus | YN. carpenteri EN. uniformis N limifrons TEN. fuscoauratus © N. concolor aquaticus MN. sericeus Norops © N. fungosus YN. vociferans BN. polylepis © N. pachypus 4 Ni tropidolepis, BN. wood © N. kreutzi sens 4 N.cusuco © N.utilensis YN. laeviventris, EN. pentaprion EN. cristifer BEN. quagguius N. humilis TEN. tropidonotus ¥ N. compressicauda @ N. wampuensis Norops © N. ocelloscapularis © N. wermuthi YN. muraila & N. sminthus WN. heterophotidotus EN. tropidogaster © 1. petersit YN. pygmaeus EN. lionotus IBN. poscilopus Norops 109Iguanidae Anolis allisoni Ctenonotus cristatellus Norops amplisquamosus Norops anisolepis Norops barkeri Norops bicaorum Norops capito Norops carpenteri Norops cobanensis Norops compressicauda 's have no or very are shown, Fi small dew bis 273, Anoles with extended dewlaps. Because in most species fen: ps (one exception in this respect: is Norops johnieye 110Iguanidae Norops crassulus Norops eupreus Norops dollfusianus Norops humilis (eastern Panama) Norops johnmeyeri (female) Norops humilis (central Panama) Norops limifrons Norops laeviventris Norops lemurinus (Nicaragua) litIguanidae Norops limifrons (eastern Norops lionotus Norops macrophallus Panama) Norops pentaprion Norops petersii Norops poecilopus Norops polylepis (normal) Norops rodriguezii Norops sagrei Norops sericeus 112Iguanidae Norops serranoi Norops utilensis Norops wermuthi Norops woodi Norops zeus 113114 Iguanidae Key to anoles (genera Anolis, Dacty- loa, Ctenonotus, Norops) Elaborating an identification key to all Central American anoles is a difficult task, partly because some species are known from a single specimen or a few individuals. In these cases, the variation of supposely diagnostic character- istics is insufficiently documented. Some species are extremely similar in their external morpho- logy and coloration, but can be distinguished readily by their very different hemipenial morphology, which, in some cases, is interpreted as evidence for only distant relationships of these eryptie species (e.g. Norops cupreus and NV, macrophallus; Kontrr & KRetT2 1999). The fol- lowing information (including SVL) refers to adults only, Juveniles of many anole species are extremely difficult to identify. Also, there is con- siderable intraspecific (particularly sexual) variation in some species. Therefore, a few spe- cies appear more than once in the following key. Statements regarding coloration (of body and dewlap) refer to specimens in life; dewlap color refers to adult males, if not otherwise specified (females of most anole species have no or only a rudimentary dewlap). Subdigital lamellae counts are given for phalanges ii to iv of the fourth toe (not including distal phalanx), if not otherwise specified 1 a Mostly green in life (capable of rapid color change to olive brown); in adult males head conspicuously large and elongate; dewlap pink; 2-3 scales between nasal ang rostral; SVL 65-87 mm; short-legged (4"° toe of adpressed hindlimb reaches usually to ear opening, in some individuals to point be- tween ear opening and eye); dewlap pink x . Anolis allisoni b Combination of characters different from that above 2 Fig. 274, Norops tropidonotus; note tube- axillary pocket (arrow). Fig. 275. alation anteri- or and posterior of tympanum b. Norops humilis a. Norops biporcatus c. Norops fimifrons e. Norops pjolensis 1, Dactyloa latitrons Fig. 276. Ventral scales in anoles: a. imbricate, strongly keeled, mucronate b. imbricate, weakly keeled, non-mucronate c. smooth, non-imbricate d. granular, not keeled e. imbricate, smooth £ slightly imbricate, smooth Fig. 277. Norops wermuthi. a (left), Mid-dorsum; note several rows of enlarged dorsal scales, b (right), Heterogeneous lateral body scales.2a ‘A deep tube-like axillary pocket present (Fig. 274) rasan No deep tube-like axillary pocket present, at most, a shallow axillary depression .... 8 Seales anterior to ear opening flat and kee- led and about twice as large as scales posterior to ear opening — latter are usually granular; Fig. 275a) .. .4 Scales anterior to ear opening about same size as scales posterior to ear opening (all granular; Fig. 275b) Dewlap uniform rose to purple; tail in males conspicuously compressed... sruistunn Norops compressicauda Dewlap orange red with yellow margin with or without a central dark streak); tail not conspicuously compressed 5 Adult males 50-60 mm SVL, adult females 45-55 mm SVL; dewlap orange red with a yellow margin and a distinct central dark Streak .. Norops tropidonotus Adult males 40-50 mm SVL, adult females 42-51 mm SVL; dewlap orange red with a yellow margin, but without a central dark Streak ... .. Norops wampuensis Devilap rose with purple spots ..... vu. Norops uniformis Dewlap red orange with yellow margin or dewlap uniform red orange without yellow margin (populations in eastern Panama) 7 Hemipenis relatively large with well- developed elongate lobes and with a strongly calyculate surface on both the truncus and the lobes; maximum SVL 46.2 mm in males and 50.0 mm in females Norops humilis (in part) Hemipenis relatively small with short and stout lobes and without a strongly calycu- late surface on either the truncus or the lobes; maximum SVL 37.0 mm in males and 41,0 mm in females ..... a " Norops quaggulus Tail distinctly compressed laterally and with a crest of enlarged scales (especially visible in males) wd Tail not distinctly compressed laterally, although a slightly carinate crest ean be present ... 12 Ventrals smooth; tail with a crest of supra- caudal scales of unequal size (sequences of smaller seales that increase in size to the following lateral whorl; Fig. 280); dewlap greenish yellow with orange margin .. = Ctenonotus cristatellus 12a 13a dda 15a Iguanidae Ventrals faintly to distinctly keeled; supra caudal scales of equal size (Fig. 280b- dewlap dark orange, orange-red or orange with pink streaks .. 10 3 seales between supraorbital semi circles; supraoculars smooth -Norops barkeri Bither 0-1 or 46 scales between supra- orbital semicircles; supraoculars keeled 11 0-1 scales between supraorbital semi- circles; males with enlarged postanal scales . vosnnnn NOrOps sagret 4-6 scales between supraorbital semi- circles; males without enlarged postanal scales .....0.:. Norops loveridgei (in part) Head conspicuously broad and stout, snout rather blunt in lateral aspect; long legged toe of adpressed hindlimb reaches at least to anterior border of eye); usually with a pale band across chin (most distinet in juveniles; Fig. 279);dorsal seales smooth, juxtaposed,’ mostly pentagonal or hexa- gonal; tip of tail enlarged, not tapering; no enlarged postanal seales in males; SVL 80- 96 mm; dewlap uniform greenish yellow rm Norops capito Combination of characters different from that above assests 13 Short-legged (4h toe of ites hindlimb reaches usually to ear opening, in some individuals to point between ear opening and eye); large adult size (SVL to 100 mm); 9-11 supralabials to level below center of eye; males without enlarged postanal scales; dorsals small, not much larger than flank seales .. 4 Combination of characters different from that above ....... 15 Dorsal head seales faintly keeled; ventrals only slighty keeled with rounded posterior margins (Fig. 276b), 62-78 ventrals be- tween levels of axilla and groin; 120-140 scales around midbody; throat region with dark streaks; dewlap’ purple with pale yellow margin and pale yellow gorgetal Seales . ...Norops petersi Dorsal scales strongly keeled; head ventrals strongly keeled and mucronate (Pig. 276a), 40-52 between levels of axilla and groin; 100-112 scales around midbody; no dark streaks in throat region; dewlap blue with dark orange margin . ‘Norops biporcatus Iris metallic blue in life; large adult size (SVL to 100 mm); body often with dark green or brown bands with numerous small dots; 2-3 scales between supraorbital 115116 Iguanidae Fig. 278. Enlarged post- anal scales in Norops tropidogaster. a, Ctenonotus cristatellus b. Norops sagroi . Norops capito 0. Norops poiylepis Fig. 280. Lateral scalation of tail (scales of whorls shaded) a. Norops utifensis , Norops pentaprion Fig. 281. Ventral surfaces of foot. a. Dactyloa lations b. Dactyloa frenata Fig. 282, Eye region (enlarged superciliaries orange). 18a 19a 20a 2a semicircles; 1 scale between suboculars and supralabials; males with enlarged postanal scales; dorsal ground color dull brick red or dark olive in life, pale yellow to tan with numerous small punetations in preservative; dewlap large, dark-orange, brownish olive or black (capable of color change) ... Norops woodi Combination of characters different from that above woe eee) 17 22 Ventrals distinctly keeled; 1-2 scales be- tween suboculars and supralabials; males without enlarged postanal scales -Norops loveridgei (in part) Ventrals smooth; suboculars and suprala- bials in contact; males with enlarged postanal scales .vs.secssssssesssessestone 1B, Short-legged (44 toe of adpressed hind- limb reaches usually to ear opening, in some individuals to point between ear open- ing and eye); brown and cream / white in coloration, often with a greenish east in life 19 Long-legged (4th toe of adpressed hindlimb reaches at least to center of eye; mostly green in life «..seseeseneeeeesnses 20 Interpgrietal plate not distinet, no parietal eye; 4°" toe with fewer than 50 subdigital lamellae (from base of digit to elaw); adult size to 111 mm (males) or 104 mm (fe- males) Dactyloa microtus A well developed interparietal plate pres- ent, distinctly larger thag adjacent scales, parietal eye present; 4° toe with more than 50 subdigital lamellae (from base of digit to claw); adult size to 153 mm (males) or 135 mm (females)... enna . Dactyloa insignis Adult size more than 100 mm... Adult size less than 90 mm .. All of upper margin of eye with enlarged superciliaries forming a’ stiff ridge (Fig. 2820) so Dactyloa latifrons Enlarged superciliaries restricted to anter- ior half of eye, followed by small granular Seales posteriorly, not forming a stiff ridge (Fig. 282) a 21 Dorsum with 4-6 oblique brown bands (not. formed of adjacent ocelli) interspersed by blue-outlined yellow patches; dewlap dirty cream with broad yellow scale rows irregu- larly interspersed with smaller emerald green scales; nasal in contact with rostral or separated by one scale .. sass Dactyloa easildae Doraum with several datk green, anell (most outlined with yellow) that are close-22a 25a 27a ly arranged and form dark oblique bands; dewlap dull, cream-colored (occasionally with black shading); nasal separated from rostral by one or two scales Daetyloa frenata 6-9 scales between interparietal and supraorbital semicircles (Fig. 283b); body yellowish green, with broad transverse brown bands and a pale greenish yellow lateral longitudinal stripe; dewlap red- orange with oblique yellow stripes, . Norops aquaticus 0-4 seales between interparietal and supraorbital semicircles (Fig. 283a); body pattern different than above; dewlap color different than above ..... sean 88 Bright green in life ... a 24 Mostly gray or brown (at most dull olive green) in life oe : 225 Vaptrale distinetly keeled: short-legged toc of adpressed hindlimb does not ate eye); SVL to 60 mm .. be ‘sun Dactyloa chloris Vontrale snspoth; lotgelegwed (44 toe: of adpressed hindlimb reaches at least to anterior border of eye); SVL to 73 mm ....... Dactyloa chocorum Eyes blue in life; very long-legged (4th toe of adpressed hindlimb reaches beyond anterior border of eye); distal phalanx of fingers and toes conspicuously long, about the same length as combined length of penultimate and antepenultimate phalan- ges; dorsal ground color olive brown with a greenish cast; lateral body with 3-5 oblique or vertieal’ pale lines - Norops alvarezdeltoroi Combination of charseters different, than that above, particularly, distal phalanx of fingers and toes not conspicuously long, much shorter than combined length of penultimate and antepenultimate phalan- ges seme Ventrals amie or smooth and subimbri- cate; flank seales homogeneous; very short- legged (44 toe of adpressed hindlimb rea- ches not much further than shoulder); 5 or fewer horizontal rows of loreals; supraocu- lars smooth; lining of throat black (pentaprion group .. Combination of characters different from that above ... 31 A pair of small bony parietal protuber- ances posterior and lateral to interparictal plate; seales along dorsal surface of tail carinate, but not enlarged .. - ‘Norops fu Tguanidae b No bony parietal protuberances; seales along dorsal surface of tail carinate (en- larged oF not)... sonussssse 28 28a Ventrals keeled; a dark interorbital bar present; males with enlarged postanal SCAIES .erneeissrnies NorOps vociferans b Ventrals smooth; no dark interorbital bar; males without enlarged postanal scales 29 29a Proximal subdigital scales of toes granular (Big. 28a); scales along dorsal surface of tail carinate, but not enlarged; SVL to 59 mm Norops utilensis bb Proximal subdigitel acales of toes differen tiated as slightly broadened lamellae (Fig 281b); scales along dorsal surface of tail b. Norops aquaticus Fig. 283. Dorsal view of head. Norops wermuthi Norops polylepis Fig. 284. Chin region (postmental seales orange). 117118 Iguanidae 30a Bla 32a 38a 35a carinate and distinctly enlarged; SVL to 79 mm .... . 80 Adult males with a well-developed serra- ted mid-dorsal crest (Fig. 285a), SVL to 88 mm oi " Norops cristifer A double-row of slightly enlarged mid-dor- sal scales, but no serrated crest present, (Fig. 285b); SVL to 79 mm .. ‘ssi Norops pentaprion Most dorsal head and body seales strongly keeled; lateral body scales homogeneous; ventral scales strongly keeled and mucronate; 7-12 distinetly enlarged dorsal seale rows; males without enlarged post- anal scales; dewlap red orange with yellow margin or dewlap uniform red orange without. yellow margin (populations in eastern Panama); maximum SVL 46.2 mm. in males and 50.0 mm in females saves .. Norops humilis (in part) Combination of characters different from that above ... ti 82 Usually a conspicuous lyriform marking in occipital region, can be fragmented (pre- sent in about 80% of individuals); ventrals strongly keeled and mucronate; relatively Jong-legged (4*® toe of adpressed hindlimb reaches at least to posterior border of eye); males without enlarged postanal scales; dewlap red, red-orange or brown-red (emurinus group) .. 33 Combination of characters different from that above ... ST 10-20 dorsal seale rows irregularly en- larged; flank scales usually heterogeneous (enlarged scales scattered among smaller scales); SVL 50-60 mm (males) or 55-72 mm (females); dewlap red-orange with a dark blotch Norops vittigerus Only the two medial dorsal scales rows slightly enlarged; flank scales homoge- neous; dewlap without a dark blotch, al- though often with black gorgetal scales 33 Dewlap red to red-orange without suffu- sion of black pigment, often with black gor- getal scales; males, on the average, smaller than females -Norops temurinus Dewlap red to pink-red, with suffusion of black pigment centrally; gorgetal scales white; males, on the average, larger than females, or both sexes about the same size 0 85 5-8 horizontal rows of loreals; hemipenis unilobate; Pacific versant of northern Central America .......... Noreps serranoi 8-10 horizontal rows of loreals; hemipenis hilobate; endemic on Islas de la Babi Honduras ... oe 36 a. Norops cristifer b. Norops pentaprion Abb, 285. Mid-dorsal scales in Norops eristifer and N, pentaprion 36a 37a 38a 39a Dewlap orange red, with suffusion of black pigment centrally; males average 65 mm SVL (maximum 75 mm), females 64 mm SVL (maximum 67 mm); endemic on Isla de Utila .... Norops bicaorum Dewlap pink red, with suffusion of black pigment centrally; males average 55 mm. SVL (maximum 62 mm), females 57 mm SVL (maximum 63 mm); endemic on Isla de Roatan ....s.n Norops roatanensis Lateral body scales heterogencous, solitary enlarged keeled or elevated scales (often whitish) scattered among smaller (mostly granular) laterals (Fig. 277b); males with enlarged postanal scales; suboculars and supraiabials mostly in contact (erassulus group) .. . 88 Combination of characters different from that above, particularly, lateral body scales homogeneous ........ 49 Enlarged dorsal scales abruptly different in size from much smaller lateral body scales; dewlap orange-yellow ..... . Norops amplisquamosus Enlarged dorsal scales grade into smaller lateral body scales (Fig. 277a); dewlap color different from that above ....00e00--e000 BO Dewlap dirty white, pale yellow or pale gray; ratio tail length /SVL'1.9 or less .. 4043a 45a 46a 4a 48a, Dewlap color different; SVL greater than 1.8 Dewlap extending to about level of axillac Norops laeviventris Dewlap extending well onto chest posteri- or to level of axillae ....... ar Dewlap pale yellow with purple gorgetal scales; 6-7 scales between second canthals ‘Norops kreutzi Dewlap dirty white or pale gray with gor- getal scales of same color; 7-10 scales be- tween second canthals ..... Norops cusuco Ventrals strongly keeled; 8 or more rows of distinctly enlarged dorsal scales ........ 43 Ventrals smooth or only faintly keeled; 6-8 rows of distinctly enlarged dorsal scales Ratio tail length / SVL 2.5 or greater; 4 scales between second canthals; 8 rows of distinetly enlarged dorsal scales; yentrals in one head length ' Norops rubribarbaris Ratio tail length / SVL smaller than 2.4; 5- 9 scales between second canthals; 10-23 (mostly 12-15) rows of distinctly enlarged dorsal scales; 13-27 ventrals in one head length . a Ad Usually fewer than 27 dorsals in one head length (17-33, mean 24); no nuchal crest with enlarged scales ... Norops crassulus Usually more than 27 dorsals in one head length (21-43, mean 30); a nuchal crest: with enlarged scales usually present .... 45 1-2 seales between supraorbital semi- circles; 22-26 lamellae beneath 4. toe; SVL. to 47 mm... Norops anisolepis 0-1 scales between supraorbital semi- circles; 23-30 lamellae beneath 4. toe; SVL to 53 mm ... Norops haguei 47 48 Medial dorsal scales uniform in size, with- out interspersed small scales sinus Norops heteropholidotus Small scales irregularly interspersed among the enlarged medial dorsal scales ‘Norops muralla Ventrals perfectly smooth Ventrals faintly keeled ....... Hemipenis with one asulcate-side process; dewlap of adult males of moderate size with relatively numerous gorgetal scales ... Norops sminthus Hemipenis with two asuleate-side proces- ses; dewlap of adult males small, with rela- 49a 5la 52a 53a 55a ~ewwwwe Iguanidae tively few gorgetal _seales e . Norops wermuthi Ventrals perfectly smooth, flat and distinetly imbricate (Fig. 276e); 2 medial dorsal scale rows enlarged; canthus rostra- lis distinct; supraoculars distinetly keeled (schiedei group) ......... BO Combination of characters different from that above ... Bist 56 Dewlap of males orange-red with a large central blue blotch; dewlap of females well developed, yellow with a large central blue blotch; SVL to 70 mm (males) or 65 mm (emales) Norops johnmeyeri Dewlap color different; females without a well developed dewlap; SVL to 60 mm .. 51 Parietal ridges usually well developed, a distinct parietal depression present; males usually with slightly enlarged postanal scales anaes O8 Parietal ridges absent to only slighty developed, parietal depression absent or only slightly developed; males usually without enlarged postanal scales... 58 6-8 scales between second canthals; snout scales multicarinate, 52-62 dorsal’ scales between levels of axilla and groin; 7-9 (rarely 6) horizontal rows of loreal dewlap purple .... Norops purpurgularis 10-11 seales between second canthals; snout scales unicarinate; 43-54 dorsal seales between levels of axilla and groin; 6- 7 horizontal rows of loreals; dewlap rose with purple spot Norops pijolensis Seales of supraorbital semicircles distinet- ly smaller than seales between semicircles; dewlap yellowish orange with a red central blotch .. ... Norops parvicireulatus Scales of supraorbital semicircles about equal in size to, or larger than seales be- tween semicircles; dewlap color different BA Suboculars and supralabials usually in contact; usually 3-4 faint pale oblique lines on flank (especially in males); ratio shank. length / head length 1.04-1.18; dewlap pur- ple eas ... Narops cobanensis Usually one complete scale row separating suboculars and supralabials; no faint pale oblique lines on flank; ratio shank length / head, length variable; dewlap color variable... ssh LE Nasal and posiscatel seales fused; 6 or fewer medial rows of dorsal scales slightly enlarged (distinetly larger than flank 119120 Iguanidae b 56a 57a 62a scales); dewlap rose ... ‘Norops hobartsmithi Nasal and postrosital scales distinct; 4-20 medial rows of dorsal scales slightly en: larged (distinctly larger than flank scales) dewlap purple . Norops matudai 8-10 medial dorsal seale rows slightly enlarged; anterior supereiliary conspic- uuoysly large and elongate; short legged (4"2 {oe of adpressed hindlimb reaches to a point between shoulder and tympanum, rarely beyond tympanum); tympanum very small (less than 1/2 the size of interpa- rietal plate); no enlarged postanal scales; dewlap yellowish orange with large blue blotch .. Norops sericeus Combination of characters different from that above : 57 Usually 4 (rarely only 2) medial dorsal scale rows slightly enlarged; 0-2 scales be- tween supraorbital semicircles; supraocu- lars distinctly keeled; ventrals faintly to distinctly keeled, but not mucronate; males without enlarged postanal scales; 4* toe of adpressed hindlimb reaches to a point be- tween shoulder and eye; dewlap of males large 58 Combination ot chersaesrs dittavant tigi that above .. 62 Ventrals faintly keeled; dewlap wegaly chocolate brown; endemic on Great Corn Island, Nicaragua .. Norops villai Ventrals distinctly keeled; dewlap color different; distributed on mainland Central America .... 59 Dewlap yellow; SVL to 44 mm ..... saunas Norops dolifusianus Dewlag cole dittarent Geom! that slow: maximum SVL greater ........ 60 Interparietal plate smaller than ear open- ing; dewlap uniformly red; SVL to 69 mm vo ... Norops cuprinus Interparictal plate larger than ear open- ing; dewlap bicolor with darker basal por- tion; SVL to 55 mm... 61 Surface of hemipenis covered with numer- ous small spines; dewlap rose with a orange-yellow basal blotch Norops macrophallus Surface of hemipenis calyculate; dewlap brown or purple with darker brown or orange brown basal blotch 8... : ..Norops cupreus Ventrals distinely keeled, imbricate and usually mucronate; anterior nasal usually divided i sssnis OB b 63a 64a 68a Ventrals smooth, or, if faintly keeled, then with rounded posterior margins; anterior nasal usually single 68 Very long-legged (4') toe of adpressed hindlimb reaches beyond eye); without enlarged postanal scales; 4-6, rarely 3 scales between supraorbital semicircles; pads under 4" (oe only slightly expanded, at most about twice as wide as those on base of digit; flanks have pearly (nacreous) reflectance in bright light Norops tropidolepis Lage shorter; males with enlarged posta- nal scales or not; rarely 4 scales he- tycen supraorbital semicircles; pads under toc slightly or distinetly expanded; flanks with or without pearly (nacreous) reflectance in bright light ......:.:.000.000. 64 Short legged (4th toe of adpressed hindlimb reaches to a point between shoul- der and tympanum, rarely beyond tym- panum); dewlap blue; 0-1 scales between supraorbital semicircles; dorsals large with their keels arranged in longitudinal rows; males with enlarged postanal scales Norops auratus Combination of characters different from that above ... 65. A conspicuous pale lateral stripe from neck to at least mid-trunk present; 18-24 rows of medial dorsal scales distinctly enlarged, dorsals flattened, penta- or hexagonal; 1 seale between suboculars and supralabials; 2.4 scales between supraorbital semi- circles; dewlap uniformly yellow or orange (lionotus group) conn OB Combination of characters different trom that above 67 6-17 scales between second canthals ... Norops lionotus 20-22 scales between second canthels ... Norops poecilopus Males with strongly enlarged postanal seales; no lateral stripe bordered above and helow by a black stripe along each side of the body ............ Nerops tropidogaster ‘Males without enlarged postanal scales; & lateral stripe bordered above and below by a black stripe along each side ofthe body - _ Norops townsendi 4-6 seales between supraorbital semi- circles; first and second canthals with more than one keel; 8-10 horizontal rows of lore- als; 1-2 scales between subogplars and supralabials; long-legged (4° toe of adpressed hindlimb reaches to a point beyond eye) ... Nerops pachypusWa Vda ‘Ta 76a, Combination of characters different from that above .... .. 69 8-10 rows of dorsal scales distinctly en- larged and abruptly larger than much smaller laterals; males without enlarged Postanal scales; dewlap pale red Norops pygmaeus thd rows of doreal scales slightly enlarged, usually grading into smalll lateral scales; males with enlarged postanal scales or not; dewlap color different 70 Long-legged (4¢} toe of adpressed hindlimb reaches at least to center of eye, usually beyond eye) . 71 Short-legged (4th toe of adpressed hindlimb does not reach to eye, usually not beyond ear opening) ....e.eee-- 76 Ventrals distinctly keeled and lightly imbricate .. ae Ventrals usually smooth on ni ares 73 An ocellated shoulder spot present .. _ Norops ocelloscapularis No ocellated shoulder spot . avandia ‘Norops yoroensis 2 elongate, overlapping superciliaries; SVL to 57 mm; dewlap orange with a darker basal portion or dirty white with an orange basal portion ... Norops polylepis 1 elongate superciliary; SVL to 48 mm; dewlap color vaiable 4 4th toe of adpressed hindlimb reaches to center of eye; dewlap orange with a darker basal portion ...........Vorops rodriguezti 4th toe of adpressed hindlimb reaches beyond eye; dewlap color different ....... 75 Dewlap dirty white with a basal orange- yellow spot or (populations in central and eastern Panama) dewlap almost. complete- ly orange-yellow; females with 34-49 dor- sals in one head length Norops timifrons Dewlap uniformly dirty white, no basal orange-yellow spot; females with 49-59 dorsais in one head length ... Norops zeus Body greenish in life (in preservative yel- low, with numerous small dark punctua- tions, resulting in a fine speckled appear- ance); venter usually with a lichenous re- ticulum: dewlap uniformly orange « ..... Norops carpenteri Body in life gray or brown (in preservative, with no fine speckled appearanee): dewlap variable .. ant Iguanidae T7a 2-4 rows of slightly enlarged medial dorsal scales; males with enlarged postanal scales evens 8 b Without rowa of enlarged medial doreal scales; males without enlarged postanal scales . 719 78a 2-3 scales between supraorbital semi- circles; dewlap of males very large (poster- ior insertion at mid-venter), orange-red with yellow gorgetal scales and a red mai BIN cece seeesssesssseerr Norops fortunensis: b 1-2 scales between supraorbital semi- circles; dewlap of males smaller (posterior insertion at level of chest), pink with white gorgetal seales and orange margin ne Norops pandoensis 79a Dewlap of males small (posterior insertion about level of shoulder), rust orange gra- ding to yellow on free margin ". Norops altae b Dewlap of.siales larger (posterior insertion about level of chest or beyond), color dit: ferent . 80 Dewlap pink to pinkish-brown Norops fuscoauratus b Dewlap color different from that above 81 Bla 2 scales between supraorbital semicircles; dewlap of males orange with red margin, gorgetal scales yellow; SVL to 47 mm rcncigece ecco Norops exsul b 1 scale between supraorbital semicircles; dewlap of males red with orange spot_at anterior margin, gorgetal scales white; SVL to 54 mM... Norops kemptoni Further Reading DUNN 1930b, Baxwour 1932, 1934, H. Sir & Kerster 1955, Stuarr 1955, ETHERIDGE 1959, Mever & W1SON 1971, E. WILLIAMS 1976, Savack & TaLnor 1978, Guyer & Savacr 1987, 1992, Savace & Guvek 1989, E. WiLLiams 1989, McCranib et al. 1992, 1993a,b, Anosema & Inaxkz 1994, KOHLER 1996a-c, 1999a, KOHLER & McCRanté 1998, KOHLER & OBERMEIER 1998, Savace & Guyer 1998, Konner et al, 1999, KouLen & McCRant: 2001, KouLer et al. 2001a, McCraniz & Kouer 2001, McCrante et al 2000, 2001, NicHOLSON et al. 2001 (morphology, systematics); SEXTON et al. 1971, Frc 1973a, b, Fircu et al. 1976, Fircit 1982 (ecology, ethology); FLASCHENDRAGER & WIJFFELS 1997 (eaptive hus- bandry and breeding). 121Iguanidae Basiliscus All four known species of basilisk are found in Central America. Earlier claims that the distribution of the Ecuador basilisk (Basiliscus galeritus) reached as far as Costa Rica (LANG 1989) proved to be erroneous, probably due to misidentifica- tion of a juvenile B. basiliseus (J.M. SAVAGE pers. comm. 1996). However, this species has been collected in the Darién region of Panama (R. IBANbz pers. comm. 2000). Basilisks, with their head and dorsal crests, which give them a truly dragon-like are among the most impres- sive and bizarre lizards of Central America. Their ability to run on their hind legs even across rapidly flowing water has earned them the local name “Iguano Jesucristo”, Two important adaptations that allow these animals to walk on the water's sur- face are broad fringes of skin on the toes, which distribute their body weight over a larger area and the relatively heavy tail, which acts as a counter-balance to the upraised body. These attractive lizards are found in a number of habitats, wherever water in the form of streams, ponds, or swamps is present, and are quite numer- ous in many areas. On the Islas de la Bahia, Honduras, Basiliscus vittatus is vir- tually ubiquitous. Basilisks feed on insects, spiders, and small vertebrates, as well as the occasional bit of vegetable matter. Basiliscus basiliscus Wi Basiliscus plumifrons ¥ Basliscus galeritus Fig. 286, Adult male of Basiliscus vittatus (Isla de Utila, Honduras) Female basilisks produce three to seven clutches of eggs per season, each with 4-18 eggs, which hatch in 2-3 months (KOHLER 1993b). @ Basiliscus vittatusIguanidae Fig. 287. Juvenile Basiliscus basiliscus Fig. 290. Adult male of Basiliscus basiliscus (Manuel Antonio, Puntarenas, Costa Rica). (Manuel Antonio, Puntarenas, Costa Rica). Basiliscus basiliscus (LINNAEUS 1758), Systema Naturae, ed. 10: 206; type locality “America australi”, SVL to 240 mm Southwestern Nicaragua to northwestern South America, sea level to 1200 m elevation. Basiliseus galeritus Duméru. 1851, Cat. Méth. Coll. Rept. Mus. Paris: 61; type locality “N.-Grenade” [= Colombial. SVL to 240 mm. Darién region of Panama to western Ecuador, sea level to 1000 m elevation Basiliscus plumifrons Cove 1876, J. Acad. Nat. Sci. Philadelphia (2) 8: 125; type locality purio, Costa Rica. SVL to 240 mm, Bastern Honduras to Panama, sea level to 780 m ele- vation. Basiliscus vittatus WIEGMANN 1828, Isis von Oken 21: 873; type locality: Mexico. SVL to 170 mm. Jalisco and Tamaulipas, Mexico, to Panama, sea level to 1500 m elevation. Fig. 288. Adult male of Basiliscus plumifrons (near Pueblo Wiso, Jinotega, Nicaragua). Fig. 289. Adult male of Basiliscus galeritus (Ecuador) 123Iguanidae Key to Basiliscus la Ventral scales keeled; 1-2 mental shields in contact with infralabials (Fig. 292); head crest of adult males single, triangular in outline Basiliseus vittatus Ventral scales smooth; 3-4 mental shields on each side in contact with infralabials (Fig, 292b); head crest of adult males dif- ferent than above .. 2 A series of enlarged, raised middorsal seales, regularly separated from each other by 2-4 small scales (Fig. 293); head crest of adult males single, fleshily thiekened at base; no dorsal fin .. Basiliseus galeritus Middorsum with slightly enlarged scales showing no evidence of regular spacing; head crest of adult males different; a large dorsal fin present .. seen B Ground color green; one or two longitudi- nal rows of pale spots along sides of body; head crest of adult males comprising two lobes; chin region of juveniles uniform Basiliseus plumifrons Ground color brown; continuous pale longi- tudinal stripes along sides of body; head crest of adult males single, often extending asa thin process posteriorly; chin region of juveniles with distinct longitudinal dark streaks Further Reading Marurana 1962, Lanc 1989, KouLER 19980, 1999b. B.vitatus B galeritus aeH A . = ees B. plumifrons B. basiliscus Fig. 291. Shape of helmet in Basiliseus. ( P a. Basiliscus vittatus b. Basiliscus basiliscus Fig. 292. Arrangement of chin shields (orange) Fig. 293. Mid-dorsal scales in Basiliseus galeritus. Corytophanes ‘The helmeted iguanas are among the rela- tives of the basilisks and their distribution is restricted to Mexico and Central America. All helmeted iguanas have a later- ally flattened body and delicate proporti- ons, Even at single locality, the coloration of the C. cristatus can vary from dark brown to olive green and yellow with irreg- ular blackish brown spots or crossbands. The ventral side is pale brown in color with a few dark spots, Males are easily disti guished from the females by the hemipenial bulge at the base of the tail, larger helmets and somewhat larger heads. The helmeted iguana is a peaceful tree- dweller that spends most of its time in par- tial shade, lying in wait for its predomi- nantly insect prey. Analysis of stomach contents reveals that C. cristatus in the wild rarely, if ever, eat during the dayIguanidae KOA Fig. 294. Corytophanes perearinatus (El Imposible National Park, Ahuacha Salvador Fig, 296. Corytophanes hernandesii (El Ocote region, Chiapas, Mexico). Axprews 1979), Helmeted iguanas pro- duce 4-11 eggs per clutch, each weighing 2- 4 g and measuring 10-15 x 18-30 mm (Ream 1965, Bock 1987). Depending on temperature, the eggs will hatch after an incubation period of 60-155 days (KOHLER 1999). Hatchling C. cristatus total Fig. 295. Corytophanes cristatus (Sasl length of 57-75 mm and a SVL of 25-30 mm National Park, Nicaragua). (REAM 1965, KOHLER et al. 1994),126 Iguanidae Corytophanes cristatus (Merrem 1822), ‘Tentamen Systematis Amphibiorum: 50; type locality: *Ceylon’” [in error]. SVL to 120 mm. Veracruz, Mexico, through Central America as far as northwestern Colombia, sea level to 1640 m elevation in tropical and subtropical wet and rain forest. Corytophanes hernandesii (WrecMaNN 1831), in Gray, Synopsis Species Class Reptilia, in Guirer7H, Cuviers Animal Kingdom 9: 45; type locality: Mexico. SVL to 100 mm. From Central Veracruz, Mexico, along the base of the Yucatdn Peninsula into northwestern Honduras, sea level to 1100 m elevation in rain forest and dry forest. Corytophanes percarinatus DUMERIL 1856, Arch, Mus. Hist. Nat. Paris 8: 518; type loca- lity: “Ascuintla, dans l’Amer. centrale” = Escuintla, Guatemala]. SVL to 100 mm, Isthmus of Tehuantepec, Mexico, along the Pacific coast as far as El Salvador and into southwestern Honduras, 200-2500 m eleva- tion, Key to Corytophanes 1 a Crest of helmet continues without inter- ruption into the dorsal crest sasiital b Crest of helmet not continuous with dorsal Corytophanes hernandesii es on dorsal surface of head keeled or at least clearly lined .. .. Corytophanes percarinatus b Helmet extends past bony skull process; scales on dorsal surface of head smooth Corytophanes cristatus Further Reading LANG 1989, KOHLER et al. 1994, KOHLER 1999b, B Corytophanes cristatus Fig. 297. Corytophanes cristatus (Pico Bonito, Atlantida, Honduras Nationalpark). x S\ b. ©. percarinatus Fig. 298. Shape of head (lateral view) in \ the species of Corytophanes yy i § Corytophanes jsinendest 4 Hi Corytophanes percarinatus ryCtenosaura Depending upon species, spiny-tailed iguan- as can reach a total length of anywhere from 25 to over 100 cm. These striking uanas are found in the lowlands of Mexico and Central America from the south- eastern Baja California and the southern Sonora in western Mexico, also more or less above the Tropic of Cancer in eastern Mexico as far as Panama, and on certain islands in the eastern Pacific, the western Caribbean and the Sea of Cortez. Of the larger species (maximum total length over 100 cm), C. acanthura and C. pectinata reach only the north of Central America in the area of the Isthmus of Tehuantepec, Mexico, whereas C. similis is distributed all _acro: the Central American land bridge. These three species are predominantly ground dwellers, but they will also climb in trees. The smaller ctenosaur species, as a rule, are tree-dwel- lers. When threatened, they will retreat into knotholes, and block the entrance with their spiny tails. The beautifully colored Yucatan Spiny-tailed Iguana (Ctenosaura defensor) is a ground dweller in the rocky terrain of the northern Yucatan Peninsula, Limestone crevices serve as hiding places. All spiny-tailed iguanas lay eggs, with clutch sizes ranging from 5-15 (small spe- cies) and 20-88 (C. similis), Suitable nesting sights are open, sun exposed areas where up to five females will use a common system of holes to lay their eggs (FItcH & HACKFORTH- JoNss 1983). Three nest tunnels examined by Fitch & HACKFORTH-JONES (1983) were 11-22 m long and reached 20- 36 cm under the surface of the ground. The eggs require an incubation of approximately three months (KOHLER 1993a). The hatchlings of most spiny-tailed iguana species are predominantly green in color with green markings. One exception in this regard is the Utila Iguana (C. bake- ri), whose hatchlings are grey-brown with Iguanidae virtually no markings, possibly due to the habitat (border zones of mangrove swamp) in which the hatchlings of this species are found. Spiny-tailed iguanas reach sexual maturity at two to three years of age. Based on research of the geographic varia- bility of the spiny-tailed iguana formerly designated as Ctenosaura palearis, Buckiey & AXTELL (1997) have described the Honduran populations as a separate species, C. melanosterna. Thus C. palearis is restricted in its distribution to the Motagua Basin of Guatemala. Similarly, the populations of what was formerly known as C. quinquecarinata have been split into two species. Because of the holo- type’s identity, the name quinquecarinata has to apply to the Nicaraguan and Costa Rican populations and the Mexican popu- lations received a new name, C. oaxacana (Haspun & KOHLER 2001, KonLer & Hassun 2001). Recently, the distributional range of the three large ctenosaur species (C. acanthura, C. pectinata, and C. similis) in the area of the Isthmus of Tehuantepec and in the Central Depression of Chiapas was studied (KOuLER & MENDOZA manuscript). Contrary to earlier views that considered the ctenosaur species in the Central Depression of Chiapas to be C. pec- tinata (ALVAREZ DEL Toro 1960, 1983; JouNson 1989, 1990, StarFoRD 1990), our field work provided evidence that it is actually C. acanthura. Fig, 299. Juvenile Ctenosaura similis. 127128 Iguanidae Ctenosaura acanthura (SHAW 1802), Gen. Zool., London 3: 216, type locality: unknown SVL to 315 mm. Along the Caribbean versant from central Tamaulipas to southern Veracruz, Mexico; also in the Central Depression of Chiapas, Mexico, sea level to 1000 m elevation. Ctenosaura alfredschmidti KOHLER 1995a, Salamandra 31: 5; type locality: 70 km E Escareega on road to Chetumal, Campache, Mexico. SVL to 170 mm, Southern Campeche, Mexico. Ctenosaura bakeri St@iNEGER 1901, Proc. US. Natl. Mus, 23: 467; type locality: Isla de Utila, Islas de la Bahia, Honduras. SVL to 300 mm. Endemic to Isla de Utila, Islas de la Bahia, Honduras, sea level to 30 m elevation. Ctenosaura defensor (Core 1866), Proc. Acad. Nat. Sci. Philadelphia 18: 124; type locality: Yucatan, Mexico. SVL to 145 mm. Northern portion of the Yucatan Peninsula, Mexico, sea level to 100 m elevation. Fig. 300. Ctenosaura acanthura (Monte Bonito, Chiapas, Mexico). Fig. 301, Cienosaura alfredschmidti (southern Campeche, Mexico) Ctenosaura flavidorsalis KOHLER & KLeMA 1994, Salamandra 30: 197; type locality: 1 km S La Paz, 750 m elevation, 14°16'N, 87°40°W, La Paz, Honduras. SVL to 170 mm. Disjunet populations from eastern Guatemala to eastern El Salvador and southwestern Honduras to the Comayagua Valley in south- central Honduras; 35 to 1010 m elevation, Ctenosaura melanosterna BUCKLEY & AXTELL 1997, Copeia 1997: 139; type locality: 2 km S Coyoles Central, Yoro, Honduras. SVL to 310 mm, Valle de Rio Aguan and Cayos Cochinos, Honduras, sea level to 250 m elevation. Ctenosaura oaxacana Konner & Hasbun 2001, Senckenbergiana biologica 81: 260; type locality: Tehuantepec, Oaxaca, Mexico, SVL to 170 mm, Restricted to the Pacific ver- sant of the Isthmus of Tehuantepec, Oaxaca, ‘Mexico; sea level to 250 m elevation. Ctenosaura_vedirhina Dr Queiroz 1987a, Copeia 1987: 892; type locality: approx. 4.8 km W Roatan on path to Flowers Bay, Isla de Roaléin, Islas de la Bahia, Honduras. SVL to 270 mm. Isla de Roatén and its satellite islands (Santa Elena, Barbareta, Big Pigeon Cay), Islas de la Bahia, Honduras, sea level to 100 m elevation. Ctenosaura palearis SEINE US. Natl. Mus, 21: 381, type locality: Gualan, Guatemala. SVL to 300 mm, Valle de Rio Motagua, Guatemala, 150-700 m elevation. Ctenosaura pectinata (WIRGMANN 1834), Herpetologica Mexicana: 42; type locality: unknown. SVL to 353 mm. Southern Sinaloa, Mexico, along the Pacific versant to western Chiapas, sea level to 2000 m elevation. Ctenosaura quinquecarinata (Gray 1842), Zool. Misc, 1842: 59; type locality: “Demerara?” [based on morphological and molecular genetic data, HASBUN & KOHLER 2001 allocated the C. quinguecarinata holo- type to Nicaraguan and Costa Rican popula- tions}. SVL to 200 mm, Isolated populations in Nicaragua and northwestern Costa Rica, sea level to 250 m elevation. Ctenosaura similis (GRAY 1831), Synopsis Species Class Reptilia, in Grirrirn, Cuviers Animal Kingdom 9: 38; type locality: un- known, SVL to 489 mm, Isthmus of ‘Tehuantepec (central Tabasco on Caribbean versant and central Oaxaca on Pacific ver- sant), Mexico, across all of Central America (except highlands and very humid regions) to Panama, sea level to 1320 m elevation.ra bakeri (Isla de Utila Fig. 306. Ctenosaura oaxacana (Oi Photo: W. § Mexico), 307. Ctenosa , Hondur Fig. 304. Ctenosaura flavidorsalis ( Paz, Hondur Fig. 806. Juvenile of Ctenosaura melanosternaIguanidae Key to Ctenosaura 1 a Tail at the point of maximum cireumfer- ence wider than high (width-height ratio 1,66 — 1.76); ratio of tail length/SVL less than 1,5; maximum SVL to 200 mm; distal dark tail bands not much longer than 2 proximal ones b Tail at the point of maximum circum! ence approximately as wide as high (width-height. ratio 0.90-1.11); ratio of tail length/SVL greater than 1.6; maximum SVL greater than 250 mm; posterior dark tail bands twice as long as anterior ones 6 2 a Only about. 2/8 of tail covered with whorls of enlarged spinous scales; parietal eye clearly visible with naked eye; keels at the base of the 3°4 toe not fused together ..... 3 b Entire tail covered with whorls of en- larged spinous scales; parietal eye barely visible with naked eye; keels at the base of the 3°4 toe fused together werner 5 3 a Adult males with middorsal crest spines <2 mm high; usually gaps present along the crest row in nuchal and anterior tho- racic region; 2 or 4 postmentals; dorsal coloration usually with some yellow ele- MENLS «se Clenosaura flavidorsalis b Adult males with middorsal crest spines usually >2 mm high; nuchal and thoracic crests continuous; 2 postmentals (Fig. 31a); absence of yellow elements in dor- sal coloration vo CC. oaxacana ©. quinquecarinata Fig. 310. Dorsal view of anterior tail. 130 Fig. 311. Ctenosaura quinquecarinata (near Fig. 312, Ctenosaura similis (Isla de Utila, Honduras), Fig. 313. Mid-dorsal seales. Teustepe, Boaco, Niearagua). Photo: F. Schmidt4a b Ta Tail less spinous (paramedian caudal scales spinous on proximal 3-7 [mean value 4.5] caudal whorls); usually body pattern of a broad pale middorsal band, which may be fragmented into large blotches; SVL to 170 mm in males, 124 mm in females ... Ctenosaura oaxacana ‘Tail more spinous (paramedian caudal scales spinous on proximal 4-13 [mean value 6.4) caudal whorls); usually no broad pale middorsal band evident; SVL to 195 mm in males, 145 mm in females sessisin Ctenosaura quinquecarinata 27-29 tail whorls; ratio of tail length/SVL 0.79-0.85; rows of medial dorsals uniform and reaching to pelvic region; parietals well formed with small parietal eye; only the first 0-3 interspaces between tail whorls without complete rows of small flat scales; SVL to 170 mm sone Ctenosaura alfredsel 22-24 tail whorls; ratio of tail length/SVL 0.65-0.74; rows of enlarged medial dorsals irregular and reduced; parietals reduced, parietal eye not distinguishable; usually no complete rows of enlarged spiky scales between the first eight tail whorls; SVL to 145 mm . Ctenosaura defensor 31-47 dorsal spines; dewlap more or less strongly developed 7 66-106 dorsal spines; no dewlap, rather just a transverse fold across the throat .. 9 ‘Two complete rows of small interealary scales between whorls of enlarged spinous scales distally (starting after 7! to the 17*® tail whorls); markings and coloration of dorsal spines conform to that of adja- cent dorsal scales; dorsum predominantly unicolor grey-brown Ctenosaura bakeri One complete row of small intercalary scales between all whorls of enlarged spinous scales (two rows of intercalary scales present between most proximal 3 whorls in some specimens); markings and coloration of dorsal spines do not conform to that of adjacent dorsal scales; several wide grey to black cross bands on dorsum, separated by rows of small pale spots Ventral area predominantly black in color- ation; 9-12 supralabials a - .. Clenosaura palearis Ventral area brown in coloration; 8-10 supralabials Ctenosaura melanosterna Enlarged heavily keeled scales on the dor- sal surface of shank (Fig. 316a); snout. of older individuals blunt and rounded off (Fig. 817); 9-15, femoral pores; in the area between the 4) and 6 tail whorls, only 1. Ctenosaura quinquecarinata _b. Ctenosaura similis Fig. 314. Chin scalation (postmentals orange). a. Ctenosaura similis —_b, Ctenosaura quinquecaninata Fig. 315. Dorsal view of anterior tail (inter- calary scales orange) a. Ctenosaura alfredschmiati —_b, Ctenosaura similis Fig. 316. Scalation on dorsal surface of shank. Fig. 317, In older individuals of C. oedirhina the snout is blunt and rounded off. 131132 Iguanidae one complete row of small intercalary scales .. ... Ctenosaura oedirhina b Scales on the dorsal surface of shank not enlarged and only slightly keeled; snout not blunt and rounded off; 4-19 femoral pores; in the area between the 4 and eth tail whorls, at least two ae rows of small interealary scales son 10 10a Scales of tail whorls very spiky (angle of keels greater than 30 degrees); dorsal crest usually interrupted in pelvic region; in some specinyens, rows of scales between the 5! and 8° tail whorls reduced to one complete row .... Ctenosaura acanthura b Scales of tail whorls moderately spiky (angle of keels less than 20 degrees) with keels running diagonally (Fig. 318¢); dorsal crest in pelvic region interrupted or not; always at least two complete rows of seales between tail whorls .....c.ssee0 LL 11a Several dark crossbands on dorsum, which usually have a pale center along the dor- somedial line; dorsal crest continuous with c. C, acanthura 18. Dorsal view of anterior tail tail crest; in area between 3°4 and 5th tail whorls, two complete rows of small inter- calary Scales .. Ctenosaura similis b No dark crossbands on dorsum; dorsal crest interrupted in pelvic region er pot; in the area between the 3° and 5% tail whorls, 8 complete rows of small inter- calary scales Ctenosaura pectinata Further Reading BAILEY 1928, HENDERSON 1973, Fitch & HENDERSON 1977, 1978, VAN DBVENDER 1982, Frrcn & HackrortH-Jongs 1983, Gicca 1983, De QuEIROZ 1987a,b, 1990a,b, BuckLey & AXTELL 1990, 1997, KOHLER 1991a, 1998a,¢, 1994a,b, 1995a-e, 1996e, 1998a,b,e,g, KOHLER & KLeMMER 1994, KouLer & Srrect 1996, KOHLER & Vese.y 1996, Kouter & RITrMann 1998, Koner et al, 2000, Hassun & Konier 2001, HasBuN et al, 2001, KOHLER & HaSBUN 2001, KOHLER 2002 Clenosaura (BC. defensor C. quinquecarinata 4 ©. alfredschmidl YC. bakeri IC flevidorsalis C. o@xacana © ©. melanosterna AC. oedirhina YG. palearisIguanidae Fig. 319. Enyalioides heterolepis (Lita - San Lorenzo, Esmeraldas, Ecuador). Photo: K.-H. Jungfer Enyalioides From the primarily South American genus Enyalioides, only one species, E. hetero- lepis, reaches as far as southern Central America. So far, the existence of this spe- s has only been established at a few in the Pa Bocas del Toro, Coldn, Coclé, Panamé, and San Blas (NICHOLSON 1998, Lips 1999, KOHLER unpubl. data). The species of this genus are sit-and-wait predators in the rain forest that primarily feed on insects and spiders. At night, these lizards will sleep at a height of 0.5-1.5 m on tree trunks or lianas, sometimes even in the rosettes of the aerial roots of palm trees. amanian provinces of nyalioides heterolepis (Bocourr 1874), Ann. Sci. Nat. Paris (5) 19: 1; type locality: Vera- gua, Panama. SVL to 138 mm. Panama to northwestern Ecuador, 100-600 m elevation. Further Reading NicHOIsoN 1998, Lips 133Iguanidac Iguana ‘The green iguana (/guana iguana) is cer- tainly the best known lizard in Central and South America. The preferred habitat of this magnificent giant lizard is lowland forest, near streams, rivers, or lakes. Whereas adult iguanas spend a lot of their time in the treetops, juveniles are often found closer to the ground in bushes However, I have also observed adult green iguanas basking in the vegetation along the shoreline in Tortuguero, Costa Rica Green iguanas are impressive swimmers and divers, as they have demonstrated by leaping into the water from as much as a 10 m height when I approached within a few meters of them. In the wild, green ig- uanas consume an almost exclusively vege- tarian diet, consisting mostly of leaves. At the beginning of the dry season, the female iguana will excavate a nesting site, which, depending on how hard the ground is, will be from 30 to 200 em long (RAND 1968, RAND & DuGaN 1983). Clutch size varies from 20 to 60 (seldom to 80) eggs, primarily depending upon the size of the female. The young will hatch following an incubation period of approximately three months, which usually corresponds with the onset of the rainy season. Thus, the hatchling iguanas are ensured a ready supply of tender leaves as nourishment. Iguana iguana (LiNNAbUS 1758), Systema Naturae, ed. 10: 206; type locality: “Indiis” SVL to 550 n. Sinaloa and Veracruz, Mexico, across all of Central America as well as large parts of South America, sea level to 1000 m elevation. Two subspec Iguana i. iguana (Costa Rica southwards) and I. i. rhi- nolopha (WiEGMANN 1834) (Mexico to Costa Rica) Further Reading BurGuarpr & RAND 1982, KOHLER 1998d. Fig, 320. Iguana iguana (near Pueblo Wiso, Jinotega, Nicaragua) Fig. 821. [gwana iguana rhinolopha (Isla de Utila, Honduras). Iguana iquanaLaemanctus The casquehead iguanas, which are found only in Central America and Mexico, are remarkably well adapted for life in the trees, which they leave only to lay eggs. Predictably, casquehead iguanas are difficult for the human observer to spot amongst the green leaves. Several times a year, Laemanctus females will produce a clutch of 4-9 eggs, which hatch after only two months at an incubation temperature of 28-30 °C. shly hatched Laemanctus longipes, on average, have a SVL of 50 mm and a tail length of about 170 mm Laemanctus longipes WIEGMANN 1834, Reptilia and Batrachia. Biologia Centrali- Americana: 46; type locality: Jalapa alapal, Mexico. SVL to 150 mm. Central feracruz, Mexico, to Nicaragua, near sea level to 1200 m elevation in moist forest and rain forest. In Central America, the subspe- cies Laemanetus 1. deborrei BOULENGER 1877 (Isthmus of Tehuantepec to Nicaragua) and L. L. waltersi Sckupt 1983b (northwestern Honduras) occur. Laemanctus serratus Core 1864, Proc. Acad. Nat. Sci. Philadelphia 16: 176; type locality: Orizaba Valley, Mexico. SVL to190 mm. ‘Tamaulipas, Mexico, along the Atlantic coast as far as northwestern Honduras, near sea level to 1500 m elevation in dry forest. In Central America, the subspecies L. s. alticoro- natus Core 1864 (Yucatsin Peninsula) and L s. mecoyi_ PéRE2-Hicarepa & Voor 1985 (southern Veracruz) occur. Iguanidae Fig. 322. Laemanctus longipes. Photo: F. Riedel Fig. 323. Laemanetus serratus (female), Laemanctus serratusIguanidae Key to Laemanctus 1 a Posterior of head with spiky scales; dis- tinet dorsal crest present; seales on dorsal surface of snout large and regular .. Laemancetus serratus b Posterior of head without spiky seales; no dorsal crest present; scales on dorsal sur- face of snout small and irregular. ... Laemanctus longipes Further Reading McCoy 1968, LANG 1989, KOHLER 1999b. Morunasaurus Two species make up the genus Morunasaurus: M. annularis in premon- tane areas on the Amazon side of Ecuador and Peru and M. groi, which is distributed as far north as southern Central Americ They are diurnal ground-dwellers of stocky build that live in the shady areas of the rain forest. Both species have a very heterogeneous scalation on the dorsal and lateral surfaces of the trunk, with large conical scales scattered among the granu- Jar scales. The tail is equipped with regu- larly arranged whorls made up of enlarged spiky scales, which become increasingly smaller distally and are barely distinguish- able at the tail tip. Morunasaurus groi is predominantly reddish brown in color. The body has dark brown crossbands, which reach to the midpoint of the sides and then break into small irregular spots According to the observations of CorkiDoR et al, (1985), the animals live in burrows, which they excavate in the floor of the rain forest. During the day, these lizards will sit motionless in front of the entrances to their burrows attentively watching the surrounding area. If they feel threatened, they will disappear with lightning speed into their burrows, blocking the entrance with their spiky tails. Predators, such as snakes, are thus prevented from forcing their way into the burrows. Morunasaurus annularis reproduces by laying eggs. Morunasaurus groi DUNN 1933a, Occ. Pap. Boston Soc. Nat. Hist. 8: 76; type locality: Valle de San Anton, Panama. SVL to 110 mm. Central Panama and northwestern Colombia in tropical rain forest at 700-800 m elevation. Further Reading DUNN 1933a, CorRepor et al. 1985, KOHLER 1991b. 4 Morunasaurus grof Fig. 324, Female of Morunasaurus groi (El Valle de Anton, Panama). Photo: W. E. DuellmanPhrynosoma Of the twelve species of horned lizard that are distributed across Mexico and the USA, only the giant horned lizard (Phrynosoma asio) reaches as far as the north of Central America (REEVE 1952). Their presence in Guatemala has not been verified by any finds in this century. In Oaxaca and Chiapas, the giant horned lizard is found in sparse forest and rocky semi-desert (Baur & Monranucci 1998, KOHLER pers. obs. 1993). Phrynosoma asio feeds on insects and spiders, with the larg- est portion of their diet, as with all horned lizard species, consisting of ants (Monta- Nucct 1989). This is one of the egg-laying species of the genus, with clutch size rang- ing from 15 to 24 eggs. Young measuring 32-37 mm SVL will hatch after an incuba- tion period of 93 to 106 days. (BAUR & MONTANUCCI 1998). Phrynosoma asio Cok 1864, Proc. Acad, Nat. Sci. Philadelphia 16: 178; type locality: Colima, Mexico, SVL to 124 mm. Colima, Mexico, along the Pacific side of Mexico ta Chiapas, near sea level to 750 m elevation in savanna and dry forest Further Reading Reeve 1952, Montanucct 1987, 1989, Baur & Monranucci 1998. Iguanidae Fig. 325. Phrynosoma asio Polychrus Polychrus gutturosus, the only bush anole species in Central America, lives in the crown region of the lowland rain forest and, apparently, only occurs in low popula- tion densities. Because they move very little, bush anoles are difficult to detect in the leafy tangle of the rain forest trees. On the 9th of May, Roberts (1997) observed a copulating pair of P. gutturosus about 2 meters up a tree in La Selva, Costa Rica. A second female with palpable eggs was examined on the 24h of July (Roberts 1997). i 1 Polychrus gutturosus 137Iguanidae Fig. 926. Adult male of Polychrus gutturosus (Rio San Juan, Nicaragua). 27, Adult female of Polychrus gutturosus (Rio San Juan, Nicaragua). Polychrus gutturosus BertuoLp 1845, Nachr. Georg-Augusts Univ. und K. Ges. Wiss. Got tingen 3: 38; type locality: western Colombia (sce Myers & Borne 1996). SVL to 165 mm Northern Honduras to northwestern Eeuador in rain forest, sea level to 700 m elevation. Further Reading Tavior 1956, 138 Sceloporus In the dry regions of Mexico and Central America, next to spiny-tailed iguanas, spiny lizards are the dominant iguanine group. The genus contains over 70 species, with increasingly fewer varieties occur- ring towards the south of Central America (only three species occur south of Guatemala). Spiny lizards are diurnal small to mid-sized lizards, most of which are covered with extremely spiky scales. ‘They feed on various arthropods, as well as smaller vertebrates. Among the Sceloporus species are ground-dwellers and saxicolous species, as well as primari- ly arboreal species. Most of the Sceloporus species reproduce by laying eggs whereas the species of the formosus group, as well as S. serrifer (torquatus group), are live- bearers. The species of the variabilis group are ground dwellers in the lowlands and, in some areas, occur in high densities. Frequently these animals can be seen on tree stumps or rocks. The males, as with all species of the genus, are highly territori al and attempt to intimidate their rivals with push up-like movements of the upper body. All species of this group lay eggs (2-7 eggs per clutch depending on species). ‘The systematics of the species of the for- mosus group have yet to be satisfactorily resolved, as the morphological variation, as well as geographic distribution of cer- tain species, is as yet unknown. Thus, Sruart (1971) claimed that comparisons between populations of Sceloporus mala- chiticus produce “...) only a hopeless jum- ble of random variations”. In his generic revision, SMITH (1939a) considered mala- chiticus and smaragdinus as subspecies of formosus. Subsequently, both taxa have been designated as full species (STUART 1971, ViLLA et al. 1988, KOHLER 1990) Also, the exact distribution of the various species of the formosus group remains to be well documented. SMITH et al. (2001)stated that S. acanthinus ranges into El Salvador, whereas KOHLER & HEIMES (2002), partly based on advice from Eric N Smith (pers. comm. to P, Heimes), referred all malachite Sceloporus in El Salvador to S. malachiticus, and I have used the latter name for all El Salvadoran populations herein. With a vertical distribution of 600-3800 m above sea level, S. malachiticus and 8. smaragdinus are typical highland dwel- lers (Marion & Sexron 1971, KOHLER 1990), They are found on trees, fence posts, and on the roofs of houses (MERTENS 1952c, Ropinson 1983), Usually only a single male will inhabit a tree, which it will defend from rivals (MERTENS 1952c), Animals found at higher elevations tend more towards a terrestrial lifestyle (Ropinson 1983). On Cerro de la Muerte (Cordillera de Talamanca, Costa Rica, 3000 m above sea level), S. malachiticus will inhabit natural stone walls and rocks covered with mosses and ferns. At this ele- vation, S. malachiticus is active only when the sun shines. The air temperature mea- sured 150 cm above the ground barely reaches to 20 °C despite the sun. These lizards, therefore, have adapted to optimal use of the microclimate. They flatten their bodies and turn very dark while basking on rocks protected from the wind. The females of S. malachiticus produce one brood of 3-10 young per year (MARION & SExTON 1971, KOHLER 1990, 1998d), while at lower clevations, births occur more or less regularly throughout the year. In populations above 2000 m elevation, however, there is a highly seasonal repro- ductive cycle, with births occurring prim- arily in late January and February (Marion & Sexton 1971). The siniferus group is represented in Central America by three species, all of which are ground dwellers in hot, dry environments. In the Valle de Comayagua, Honduras, 8. squamosus is quite common- ly seen (EisenserG & KOHLER 1996). The spiny lizards in this area are generally Iguanidae Fig, 328. Adult male of Sceloporus acanthinus (near Guanagasapa, Escuintla, Guatemala) Fig. 329, Sceloporus taeniocnemis (male; near San Cristébal de las Casas, Chiapas, Mexico). found on individual stone blocks or on piles of rocks. This species lays 4-5 eggs, which hatch after 50-53 days at an incu bation temperature of 29 °C. The young measure from 17.0-18.5 mm SVL. As the sole representative of the forquatus group in Central America, Sceloporus ser- rifer reaches Guatemala and Belize. This species can be seen in great numbers at the Mayan ruins in Yucatan, although the animals are quite shy and will flee at the slightest disturbance. These attractive lizards bask on trees, as well as on rocks and brick walls, and will disappear into the nearest crack or opening when threat- ened, 139Iguanidae Sceloporus acanthinus BocourT 1873, Ann. Sci. Nat., Zool. (5) 17 (6): 24; type loeality: San Agustin, near Voledn de Atitlin, Guatemala. SVL to'99 mm. Pacifie versant of Chiapas, Mexico, and Guatemala, 400-1500 m cleva- tion. Sceloporus carinatus H, SuitH 1936b, Proc. Biol. Soe. Washington 49: 89; type locality: near Tuxtla Gutiérrez, Chiapas, Mexico. SVL to 55 mm. Grijalva Valley of Chiapas, Mexico, and adjacent Guatemala, 500-1000 m elevation. Sceloporus chrysostictus Core 1866, Proc. Acad. Nat. Sci. Philadelphia 18: 125; type locality: Yueatin, Mexico, SVL to 60 mm. Yucatén Peninsula from central Belize and north Guatemala northwards in dry forest, sea level to 200 m elevation. Sceloporus cozumelae JONES 1927, Oce. Pap. Mus. Zool., Univ, Michigan 186: 1; type locali- ; Isla Cozumel, Quintana Roo, Mexico. SVL to 54 mm, Coastal area of the northern Yucatdn Peninsula, including the offshore islands Cozumel, Isla Contoy, and Isla Mujeres in dry forest and open rocky areas, sea level to 50 m elevation. Sceloporus edwardtaylori H. Sux 1936a, Herpetologica 1: 6; type locality: Ixtepec (San Gerénimo), Oaxaca, Mexico, SVL to 107 mm. Pacific coast of the Isthmus of Tehuantepec (@axaca, Mexico) in dry forest, sea level to 750 m elevation. Sceloporus internasalis H. Swim & BuMzaHeM 1955, Herpetologica 11: 118; type locality: La Gloria, Oaxaca, Mexico, SVL to 100 mm. Los Tuxtlas region in southern Vera- cruz, Mexico, southward through extreme eastern Oaxaca to northwestern Chiapas, sea level to 2000 m elevation in oak forest. Sceloporus lunaci Bocourt 1873, Ann. Sci. Nat., Zool. (5) 17 (10): 1; type locality: Plateau of Guatemala. SVL to 95 mm. Motagua Basin and Baja Verapaz, Guatemala, 150-1000 ele- vation in dry forest. Sceloporus lundelli H. SMTi 1939, Field Mus. Publ. Zool. Ser. 26: 66; type locality: Cohune Ridge, 20 miles SE Benque Viejo, Belize. SVL to 100 mm. Yucatan Peninsula, sea level to 300 m elevation in dry forest. Two subspecies: Sceloporus 1. lundelli (base of the ‘Yucatan Peninsula) and Seeloporus |. gaigeae H. SMrri 1939 (northern part of the Yucatan Peninsula). Seeloporus malachiticus Core 1864, Proc. ‘Acad. Nat. Sci. Philadelphia 16: 178; type locality: “Arriba” [“this name apparently was applied loosely to the Meseta Central” accord- ing to Savace 1974:77], Costa Rica. SVL to 98, mm, El Salvador and Honduras across Nicaragua and Costa Rica to Panama, 600- 3800 m elevation in pine and cloud forest. Seeloporus_melanorhinus Bocourt 1876a, Ann, Sci. Nat, Zool., Paris, (6) 3, art. 12: 2; type locality: Isthmus of Tehuantepec, Oaxaca, Mexico. SVL to 105 mm, Southern Nayarit, Mexico, along the Pacific side of Mexico to the Isthmus of Tehuantepec, as well as in the Rio Grijalva Basin, Guatemala, sea level to 2000 m elevation in dry forest. In Central America, the subspecies Sceloporus m. melanorhinus (Mexico) and S. m. stuarti Swit 1948 (Guatemala) occur. Sceloporus serrifer CoPE 1866, Proc. Acad. Nat. Sci. Philadelphia 18: 124; type locality: Yucatén, Mexico. SVL to 100 mm. Several iso- lated populations from southern Tamaulipas, Mexico, as far as Guatemala and Belize, as well as in the north of the Yucatan Peninsula, 50-2300 m elevation in dry forest. In Central America, the subspecies Seeloporus s. serrifer (Yucatan Peninsula) and Sceloporus s. prezy- gus H, Sur 1942b (southwestern Guate- mala) occur. Sceloporus siniferus Cope 1870, Proc. Amer. Phil. Soc. 11: 159; type locality: Pacific ver- sant of Isthmus of ‘Tehuantepec. SVL to 71 mm, From western Guerrero, Mexico, along the Pacific Coast to eastern Chiapas, sea level to 2000 m elevation in dry forest. Fig. 380, Sceloporus carinatus (Rancho Alegre, Chiapas, Mexico).ig. 335, Sceloporus lundelli gaigeae (Coba, imes Quintana Roo, Mexic Photo: J. C. Lee 37. Sceloporus melanorhir Morelo Photo: P. Heimes Sceloporus internasalis (Catemaco, Fig celoporus serrifer (Ruinas de Veracruz, Mexico) Photo: P. Heimes — Mayapan, Yucatan, Mexico).Iguanidae Sceloporus smaragdinus Bocourt 1873, Ann. Sci. Nat., Zool. (5) 17 (10): 1; type locality Guatemala, in the vicinity of Solola, Totonicapam, and Quetzaltenango. SVL to 85 mm, Highlands of Guatemala and Chiapas, 2000-4000 m elevation in pine and cloud forest. Sceloporus smithi HARTWEG & OLIVER 1937, ce. Papers Mus. Zool. Univ. Michigan 356: type locality: Quiengola Mountain, about miles NW of the town of Tehuantepec, Oaxaca, Mexico. SVL to'71 mm, Pacific side of the Isthmus of Tehuantepec (Oaxaca Mexico), sea level to 600 m elevation in dry forest. celoporus squamosus Bocourt 1874, Miss Sci. Mex., Rept.: 212; type locality: “Environs de Guatemala et de ’Antigua” (=Voledn Antigua, Guatemala). SVL to 59 mm. From western’ Chiapas, Mexico, along the Pacific side of Central America ‘into northwestern Costa Rica, sea level to 1000 m elevation in dry forest. Sceloporus taeniocnemis Cops 1885, Proc Am. Phil. Soc. 22: 399; type locality: Guatemala. SVL to 80 mm. Central plateau of Chiapas, Mexico, to central Guatemala, 1200-2500 m elevation in pine and cloud forest. Sceloporus teapensis GONrHER 1890, Reptilia and Batrachia. Biologia Centrali-Americana: 75; type locality: Teapa, Tabasco, Mexico. SVL to 70 mm, From Xalapa, Veracruz and north- eastern Oaxaca, Mexico, across northern Guatemala as far as Belize, sea level to 1000 m elevation in savanna and dry forest. Seeloporus variabilis WigGMANN 1834, Herpetologica Mexicana: 51; type locality: Mexico. SVL to 74 mm, From Texas, USA along the Caribbean side of Mexico into northwestern Costa Rica, sea level to 1700 m elevation (to 2500 m elevation according to JOHNSON 1989) in savanna and dry forest. In Central America, only the nominate subspe- cies occurs (of which olloporus H. Swrri 1937 is a synonym). Fig. 341. Sceloporus smithi (near Mixtequilla, Oaxaca, Mexico). 142 Fig. 889. Seeloporus siniferus (near Mixte- quilla, Oaxaca, Mexiko). Fig. 340. Sceloporus smaragdinus (btw. Soloma and Santa Eulalia, Guatemala), Photo: T. BilleIguanidae Sceloporus A S.carinatus 4 S. edwardtaylori S. chrysostictus IS. squamosus BS siniferus Fig. 342. Sceloporus squamosus (Santa Lucia, Intibuca, Honduras) Sceloporus 4 S. cozumelae S. lundeti S. variabilis IS. teapensis BS smithi HS. acanthinus S. internasalis WS. lunaei Ws. malachiticus S. smaragdinus & MES. taeniocnemis 343, Sceloporus teapensis (Palenque, japas, Mexico). Photo: P, Heimes | Sceloporus serrifer | Sceloporus melanorhinus Fig, 344, Sceloporus variabilis (Cafton de Sumidero, Chiapas, Mexico) 143144 Iguanidae Key to Seeloporus al pocket present group) b Without postfemoral pocket... 2a (Fig. 347) aeniousB i) Males uniform pale below, no colorful belly patches; endemic to the north of the Yuca- tin Peningula ..... Seeloporus cozumelae b Males with obvious ventral markings, con- sisting of two large purple-colored belly patches, usually with black borders 3 49 or fewer dorsal scales from posterior of head to base of tail; 7-9 rows of enlarged dorsal scales in the pelvic area; seales in the inguinal region relatively large saan ‘Sceloporus teapensis b 50 or moro dorsal seales from posterior of head to base of tail; 10 or more rows of enlarged dorsal scales in the pelvic area; scales in inguinal region relatively small 4 Ground color very dark, nearly black, with very obvious yeliow or white longitudinal stripes (Fig. 348a), which are 2-2.5 scales wide; without obvious dark diagonal bars on dorsum and body sides; keels of the dor- sal scales form distinct fine combs; hatch- lings with pink colored tail .... 3 sesnenneeernsses Seeloporus smithi b Ground color grey to brown, with less distinct pale longitudinal stripes (Fig. 348b), which are 1,5-2 scales wide; distinct dark diagonal bars on dorsum and body sides present; keels of the dorsal scales do not form distinct fine combs; hatchlings with grey brown tail . Sceloporus variabilis Males uniform pale below, no colorful belly patches (siniferus and chrysostictus groups) .. 6 b Males with conspicuous colored belly pat- ches; if without colorful belly patches (Seeloporusedwardtaylori, only on the Isthmus of Tehuantepec), then large, very spiky-sealed species . 9 1 canthal (Fig. 349b) .. b 2canthals (Fig. 3498) 9-12 femoral pores; 38-44 dorsal scales from posterior of head to base of tail 5 . Sceloporus carinatus b 3-7 femoral pores; 28-37 dorsal scales from posterior of head to base of tail ............ . sso Seeloporus squamosus 3-11 femoral pores; 31-46 dorsal scales from posterior of head to base of tail . . Seeloporus siniferus superciliaries Fig. 345. Lateral view of head in Sceloporus malachiticus. frontonasals _supragculars interparietal frontoparietals posterior frontal anterior frontal (divided) prefrontals: Fig, 346. Dorsal view of head in Sceloporus chrysostictus. b 12-17 femoral pores; 42-57 dorsal scales from pesterior of head to base of tail zi Sceloporus chrysostictus 9 a A distinct dark neckband present ........10 b No dark neckband present (spinosus group) ssiatincoatvaibwi 16 Dark neckband on both sides pale grey or blue bordered (torquatus-group) seein: Seeloporus serrifer b Dark neckband not pale bordered (formo- SHE-GTOUD) soninisitioniveensamisaurcns LL In the snout area (between the nasals and between rostral and median frontonasal) many small scales (Fig. 350a); usually two seales between nasal and rostral . .. Sceloporus internasalis b In the snout region, a few large seales (Fig. 350b); usually 1 scale between nasal and rostral .. 7 12 12a Usually, only a single canthal 13. b Usually two canthals 16 ——aN 7) Fig. 347, Post- fermoral pocket. After SmiTH (19394). a. S. smithi Fig. 348. Dorsal coloration. a. S. siniferus b. S. squamosus Fig, 349, Lateral view of head (canthals orange). a. S. internasalis b. S. malachiticus Fig. 350. Dorsal view of head in Sceloporus. = Iguanidae 18a Supraoculars very large and arranged in a row, usually one or more in contact with medial head scales; 31-38 dorsal scales from posterior of head to base of tail; low- Jand species «0 Seeloporus lunaei b Supraoculars smaller and arranged in two rows, seldom in contact with medial head scales; 30-51 dorsal scales from posterior of head to base of tail; highland species a os 14 1da_ 30-39 dorsal scales from posterior of head to base of tail Sceloporus malachiticus b 39-51 dorsal scales from posterior of head to base of tail . Sceloporus smaragdinus 15a 34-39 dorsal scales from posterior of head to base of tail .... Sceloporus acanthinus b 39-47 dorsal scales from posterior of head to base of tail Sceloporus taeniocnemis 16a Males without colored ventral spots = Sceloporus edwardtaylori b Males with conspicuous colored Spot cae 17-24 femoral pores ee Sceloporus melanorhinus b 9-12 femoral pores Sceloporus lundelli ventral Further Reading H. Smrrat 1939a (Revision of Mexican and Central American Sceloporus species); Sires & Dixon 1982, H. SMITH et al. 1993, KOHLER 1994e, H. SMITH et al. 1995b (Sceloporus varia~ bilis group); H. Sarr & Buwzanem 1955, Suan? 1971, KOHLER 1990, 1993d, H. Surrn & PéREZ-HIGAREDA 1992 (Sceloporus formosus group); OLSON 1987, KOHLER et al. 1998; Kouter & Hemes 2002 (summary of biology, care and breeding). 145146 Iguanidae m= Urosaurus ‘The delicate, small lizards of the genus Urosaurus are distributed predominantly in the US and Mexico. Of the nine Urosaurus species, only one reaches the northern portion of Central America, namely U. bicarinatus (MITTLEMAN 1942, Wizns 1993). Phylogenetically, the Urosaurus species are closely related to the Spiny Lizards of the genus Sceloporus (BrHeRIDGE & De Queiroz 1988). Urosaurus bicarinatus is an inconspicuous pale grey colored bush and tree dweller in dry areas, usually found on the outer, thin- ner twigs. Thus, thorny bushes are a pre- ferred habitat. The chest of the male is adorned with two bright blue spots, while the throat is yellowish in color. The threat behavior of these territorial lizards con- sists of pushup-like movements, in which the whole body is pushed vertically upright and back down again. The diurnal Urosaurus feeds on small invertebrates (insects and spiders, among others). For egg laying, which, in Chiapas, occurs pri- marily in June, the females come down from the trees in order to bury their clutches of around ten eggs (approx, 10x7 mm in size) (ALVAREZ DEL TORO 1960). Urosaurus bicarinatus (Dumern, 1858), Arch. Mus. Hist. Nat. Paris 8: 549; type locality: Mexico. SVL to 58 mm. Along the Pacific Coast of Mexico from southern Sonora as far as Chiapas, sea level to 1200 m elevation. The subspecies Urosaurus 6. anonymorphus MrrrLeMan 1940 (eastern Oaxaca northward) and Urosaurus b. spinosus BUMZAHEM & SmrH 1954 (Chiapas) occur in Central America. Further Reading MrrrLeman 1942, Wiens 1993. Fig. 352, Urosaurus bicarinatus (near Mixtequilla, Oaxaca, Mexiko). Fig. 351. Urosaurus bicarinatus (near Mixtequilla, Oaxaca, Mexiko). VS. Urosaurus bicarinatusScincidae From the large family of skinks, only five genera with a total of nine species occur in Central America. Most of the remaining species are found in the Old World. Skinks typically have an elongated head, body and tail, with relatively short legs and very smooth scales, Immediately under the scales are smal] dermal bones (osteo- derms). Key to Skink Genera 1 a Without supranasals, therefore rostral in complete contact with first unpaired scale on dorsal surface of snout (Fig. 366a,b) .. 2 b Supranasals present, therefore rostral entirely or partially separated from first unpaired scale on dorsal surface of snout (Big. 855¢,4) eer.) 2 a One frontopai B55b) ..... a ‘Sphenomorphus b Two frontoparietals(Fig. 355a) 3a Scales in temporal region same size as body seales (Fig. 353b) .. Mabuya b Scales in temporal region conspicuously enlarged (Fig. 353a) .... 4 a Scales along the dorsomedial line not obviously larger than paramedian dorsals; with three pale longitudinal stripes, the middle of which branches on the middle of the head (Fig. 356a) . Eumeces b Scales along the dorsomedial line obvious- ly larger than paramedian dorsals; mark- ings different than those above (Fig. 356b,¢) Mesoscincus a, Eumeces b. Mabuya Fig. 358. Lateral view of head (temporals pale brown). Seincidae Fig. 354, Mabuya unimarginata (Isla de Guanaja, Honduras). Photo: E. Kohler ©. Mabuya Fig. 355. Dorsal head scalation in skinks (supranasals orange, frontoparietals brown). Z 1 aE. sumichrasti_b. M-managuae cM. schwartzei Fig. 356, Dorsal coloration in Eumeces and Mesoscincus.148 Scincidae Eumeces Based on a phylogenetic analysis of spe- cies traditionally grouped under the gen- eric name Eumeces, GRIFFITH et al. (2000) provided evidence that this assemblage was not a natural grouping, Consequently, they split the Eumeces sensu lato into several genera. According to this new classification, one Central American spe- cies (E. sumichrasti) remains in the genus Eumeces, whereas two species (M. mana- guae and M. schwartzei) are assigned to the genus Mesoscincus. A clutch of 11 eggs from E. sumichrasti was discovered in a freshly fallen bread- fruit tree in Belize (MILLER 1997), The eggs (15x11 mm, a bit less than 1 g in weight) lay in a knothole, which would have been about 20 m above the ground before the tree had collapsed. They were buried in a substrate mix of fallen bark and other organic material. A female E. sumichrasti (92 mm SVL) stayed by the clutch, possibly indicating that the species practices brood care, as is the case with other species of this genus (FitcH 1954, Virt & Cooper 1985, SomMA & FawceTT 1989). Freshly hatched E. sumichrasti have a total length of 70 mm with a SVL of 26 mm and a weight of 0.7 g (MILLER 1997), While adult Z. swmichrasti have a pale brown ground color with black longi- tudinal stripes, the young are somewhat. more brightly colored. The latter have a bright blue tail, and several orange spots on the head, as well as three orange to yel- low longitudinal stripes on the body (MILLER 1997). Eumeces sumichrasti (Core 1867), Proc. Acad. Nat. Sci. Philadelphia 1866: 321; type locality: Orizaba, Veracruz, Mexico. SVL to 96 mm. Central Veracruz, Mexico, to northern Honduras; also northeastern portion of Yucatan Peninsula, sea level to 1000 m elevation. Further Reading ‘TayLor 1935, GRIFFITH et al. 2000. Fig. 357. Juvenile Eumeces sumichrasti from Honduras. Photo: J. R. MeCranieMabuya While earlier authors (e.g. TAYLOR 1956) proposed that, in Costa Rica alone, three species (alliacea, brachypoda, unimargi- nata) of the genus Mabuya occurred, recent authors conclude that only one spe- cies, M. unimarginata, is distributed all across Central America (SAVAGE & VILLA 1986, VILLA et al. 1988). However, the geo- graphic variation of Central and South American representatives of this genus has hardly been examined. Therefore, the current taxonomic division can only be considered temporary. This insectivorous skink occurs commonly in many location: in the lowlands of Central America. These animals are generally found on the ground, but they will climb tree trunks in. order to bask. Mabuya unimarginata is a livebearer, with litter size varying from 2 to 7 young. Mabuya unimarginata Cope 1862b, Proc. Acad. Nat. Sci. Philadelphia 14: 187; type locality; Panama. SVL to 90 mm. Mexico to Panama, sea level to 1500 m elevation. Further Reading ‘TAYLOR 1956, Webb 1958. Fig. 358. Mabuya unimarginata (near Guanagasapa, Escuintla, Guatemala) is =. hg. SOAS a Fig. 359. Mabuya unimarginata (Bartola, Rio San Juan, Nicaragua) Fig. 860, Mabuya unimarginata (Voleén Maderas, Isla Ometepe, Nicaragua), Fig. 361, Mabuya unimarginata (Ruinas de Mayapan, Yucatén, Mexico) 149150 Scincidae Fig. 362. Me: Guanacaste, Costa Rica), sseincus managuae (Portegolpe, Photo: H. Bringsoe Mesoscincus The genus Mesoscincus was recently pro- posed for three species of skinks formerly assigned to the diverse genus Eumeces (Grirerri et al. 2000). Two of the species included in Mesoscincus occur in Central America, M. managuae and M. schwartzei. These skinks are ground dwellers in dry and seasonal rain forest, where they can be seen basking on roots and fallen logs. In July 1999, I discovered an adult specimen of M. managuae at the foot of the Volcén Cosiguina, Nicaragua, under a musty tree trunk. Sometimes these lizards are also Mesoscincus sthwartzei i Mesoscincus managuae found under loose bark on upright tree trunks. The Mesoscincus species are shy, secretive lizards that are rarely encounter- ed in nature. Mesoscincus managuae (DUNN 1933¢), Proc. Biol. Soc. Washington 46: 67; type locality: Aviation Field, Managua, Nicaragua. SVL to 117 mm, El Salvador to northwestern Costa, near sea level to 530 m elevation in dry forest. Mesoscincus schwartzei (FISCHER 1885), Abh. Naturwiss. Ver, Hamburg 8: 3; type locality: small island in Laguna de Términos, Campeche; Mexico. SVL to 120 mm. Yucatan Peninsula and bordering Tabasco, Mexico, into central El Petén in Guatemala, sea level to 250 m elevation. Key to Mesoscincus 1 a With two wide pale longitudinal stripes, which begin at the tip of snout, and fuse into a broad band in the anterior dorsal area (Fig. 356c); 21 scales at midbody; front and rear limbs touch, i.c., overlap, when laid at the side of the body oe Mesoscincus schwartzei b Dorsum brown with dark brown longitudi- nal lines, which in some individuals break up into spots (Fig, 356b); 17-19 seales at midbody; front and rear limbs do not touch when laid at the side of the body ....... soossssninninm Mesoseincus managuae Further Reading ‘TayLon 1935, 1956, Cruz et al, 1979, Reeper 1990, Grirrrrii et al, 2000. Fig. 363. Mesoscincus managuae (Rio Sandillal, Costa Rica). Photo: R. W. Van Devenderam ss Scincidae Fig. 364. Mesoscineus schwartzei (19 km N ‘Temoz6n, Yucatan, Mexico). Photo: Section of Herpetology, Natural History Museum of Los Angeles County. Fig, 365. Scincella gemmingeri (Hidalgo, Mexico). Seineella According to the taxonomic division by Greer (1974), one Central American skink species, S. gemmingeri from the subfamily = Lygosominac, is included in the genus Scincella, whereas the species assatus, cherriei, incertus and rarus are placed in the genus Sphenomorphus (Myers & DONNELLY 1991). Scincella gemmingeri has 60-68 dorsal scales from the parietal | Ml Scincella gemmingeri — to the base of the tail (at the level of the posterior border of the thighs) and 25-28 longitudinal rows of dorsal scales (H. Sarr 1949). Scincella gemmingeri (Corr 1864), Proc. ‘Acad. Nat. Sci. Philadelphia 16: 180; type locality: Orizaba, Veracruz, Mexico, SVL to 45 mm. Hidalgo, Veracruz and Oaxaca, Mexico, 200-2000 m elevation in pine forest. Further Reading ‘TayLor 1937a, H. SMH 1949. Fig. 366. Scincella gemmingeri (Hidalgo, Mexico).152 Scincidae a= Sphenomorphus The Central American herpetofauna in- cludes three apparently closely-related, small skink species from the subfamily Lygosominae (assatus, cherriei, and incer- tus) that, until the beginning of the 1970's, were variously included in the genera Lygosoma (e.g. STUART 1940), Scincella (eg. H. Smith & TAayLOR 1950) or Leiolopisma (e.g. TAYLOR 1956, STUART 1963, Perers & Donoso-Barros 1970). In his revision of genera related to Leiolopisma, GREER (1974) placed the three above-named species in the genus. Sphenomorphus, a view which is widely held today (FitcH 1983, VILLA et al. 1988). Recently, Myers & DONNBLLY (1991) described a fourth species from this genus in Panama, S. rarws, that is known only from a few specimens. Whereas Sphenomorphus cherriei are diurnal ground dwellers in the unde growth of the rain forest, S. assatus is mostly an inhabitant of subhumid area: The short legs are assisted in forward loc motion by snake-like movements of the body, especially when increasing speed. At air temperatures between 23 and 27 °C, the animals are active primarily in the morning, while they will hide from the midday sun in cooler places. In the after- noon, the agile Sphenomorphus will undertake another period of activity (FitcH 1983). Diet consists of small arthro- pods (insects and spiders), which they track primarily with their olfactory senses. Large prey animals are preferred, and even cannibalism is known among these skinks (FITCH 1983). The species of the genus Sphenomorphus reproduce ovipar- ously, with females producing several clutches per season, each with one to three (seldom four) eggs. Whereas Sphenomorphus in the constantly moist Caribbean low- lands areas reproduce throughout the year, populations at higher altitudes or areas with distinct dry periods are highly seasonal in their egg production. Thus, Fig. 367. Sphenomorphus cherriei (Cerro Saslaya, Atlantico Norte, Nicaragua). Fig. 369, Hatehlings of Sphenomorphus cherriei (Quintana Roo, Mexico). Photo: H, Bahena B.Fire (1983) reported that the reprodue- tive activities of S. cherriei in the region of Turrialba, Costa Rica, stagnate in the months from November to April. With an SVL of approx. 22 mm, the hatchlings of S. cherriei are already 40 % of their adult size, Within seven months, the young are sexually mature at an SVL of 48 mm. The average duration of a generation of S. cherriei is approximately one year. Countless predators (primarily snakes, birds and mammals) prey upon these small skinks. All Sphenomorphus species will discard their tails if they are seized Sphenomorphus assatus (Cope 1864), Proc. Acad. Nat. Sci. Philadelphia 16: 179; type locality: Volcan Izalco, El Salvador, SVL to 50 mm, Colima, Mexico, to El Salvador and southern Honduras (McCrante & KOHLER 1999a), sea level to 1500 m elevation (to 2500 m elevation according to JOHNSON 1989). Sphenomorphus cherriei (COPE. 1893), Proc. Amer. Phil. Soc. 81: 340; type locality: El Palmar, Costa Rica. SVL to 66 mm, Tabasco, Mexico, to western Panama, from sea level to 1300 m elevation in lowland and premontane rain forest. Sphenomorphus incertus (Stuart 1940), Oce. Pap. Mus. Zool., Univ. Michigan 421: 10; type locality: Voleén’Tajumuleo, Guatemala. SVL, to 67 mm. Guatemala and Honduras. Sphenomorphus rarus Myers & DONNELLY 1991, Amer. Mus. Nov. 3027: 2; type locality: SW headwaters of Rio Guabo, 780 m eleva- tion, 8°47'N, 82°11'W, Bocas del Toro, Panama, SVL to 52 mm. Known only from the type locality. Fig. 370 Sphenomorphus rarus (holotype). Photo: C. W. Myers Scincidae Key to Sphenomorphus 1a Without prefrontal scale, therefore, contact occurs between frontonasal and frontal .. Sphenomorphus rarus b Prefrontal scale present, frontonasal not in contact with frontal ... eB 2 a 30-36 scales at midbody; front and rear extremities touch when laid at side of body; tail brown in adult specimens, blue in juveniles .... Sphenomorphus cherriei b 24-31 Scales at midbody; front and rear extremities do not touch when laid at side of body; tail red brown in adult specimens, intense red in juveniles . 3 3 a 67-77 dorsal scales between parietal and base of tail (at level of posterior border of thighs); 27-31 scales at midbody ... seyioa . Sphenomorphus assatus b Fewer than 65 dorsal scales between parietal and base of tail (at level of poster- ior border of thighs); 26 or fewer scales at midbody ........ Sphenomorphus incertus Further Reading TayLor 1937a, Sruarr 1940, H. Smrrn 1946, 1951, Greer 1974, Witson & McCrantg 1994, McCranié & KOHLER 1999a. IG Sphenomorphus assatus © Sphenomorphus incertus ‘A Sphenomorphus rarus 1 Sphenomorphus cherriei 153Teiidae Teiidae ‘The lizards of the family Teiidae, including ten genera with approximately 110 species, are widespread in temperate, subtropical, and tropical areas of the USA, Mexico, Central America, South America, and the Antilles. Most species are active terrestrial lizards, and all are diurnal. All teiids are oviparous; parthenogenesis is prevalent among Aspidoscelis. Key to genera of Teiidae 1 a One row of greatly enlarged scales on the outside of the upper arm (Fig. 372a); dor- sal pattern with five or fewer longitudinal stripes; a fleshy sheath enclosing base of tongue (Fig. 371a) . ‘Ameiva b Three or more rows of moderately en- larged scales on the outside of the upper arm (Fig. 372b), dorsal pattern usually with six or more pale longitudinal stripes, or with pale spots; no fl closing base of tongue (Fig. 371b) 2 a One interparietal and 4 parietals (Fig. 386a) Cnemidophorus b One interparietal and 2 parictals (Fig. 386b,c) ... Aspidoscelis Cc a. Ameiva , Aspidoscelis Fig. 371, Floor of mouth. Note tongue sheath (arrow) in Ameiva. — se q SD a Amon ) Fig, 372. Foreleg (enlarged scales shaded). 154 Fig. 373. Ameiva undulata (El Imposible National Park, Ahuachapan, El Salvador).Fig. 376. Ameiva undulata (San Julia Suchitepéquez, Guatemala) Fig. 377. Ameiva ameiva. Photo: R. D. Bartlett 8. Ameiva festiva (Nusagant San Blas, Panama). Teiidae Ameiva In Central America, there are six ameiva species, of which A. festiva and A. undula- ta are the most widely distributed (ECHTERNACHT 1971). Ameivas are diurnal, very agile lizards that are common in many places, primarily in open secondary forests along the paths and in clearings. ‘The sunlight and heat loving ameivas keep their body temperature around 37 °C during their active phase (HirTH 1963, HILLMAN 1969, FitcH 1973a). Cooler and rainy days are spent in burrows in the ground. Ameivas are tenacious hunters, who will wander around their territory looking for prey, scratching in the leaves, digging in the ground, and searching in the under- growth and under bark, consuming any creature they can overpower, primarily insects and arachnids. The lizards are con- stantly flicking their tongues, checking scents in their surroundings. Although these lizards are ground dwellers, they will also climb tree trunks and low vegeta- tion to bask or to reach prey. Important predators of the ameivas are the diurnal, quick colubrids, including Conophis linea- tus, Dryadophis melanolomus, Drymobius margaritiferus, and Masticophis mentova- rius. The Central American ameiva species pro- duce several clutches of eggs per year, with clutch size dependent upon species and the size of the female (A. ameiva 1-9, A. festiva 2-4, A. undulata 1-7, ECHTERNACHT 1983, Kou! 1997). At 28-30 °C, the incu- bation period for the eggs of A. ameiva lasts 129-137 days. The juveniles of A. fest- iva, A. quadrilineata, and A. undulata have pale blue-colored tails, just like the juvenile whiptail Aspidoscelis deppii. Up to three ameiva species can occur in a single area (sympatric) (e.g., the species festiva, leptophrys, and quadrilineata in southeastern Costa Rica), with each making use of a slightly different ecologi- cal niche to reduce the competition (HCHTERNACHT 1971, 1983, FrrcH 1973a).Cs Es Ameiva ameiva (LINNAEUS 1758), Systema Naturae, ed. 10: 202; type locality: Brazil. SVL to 197 mm (males) and 157 mm (fe- males). Costa Rica Panama as well as large parts of tropical South America, sea level to 700 m elevation. In Panama, the subspecies Ameiva a, praesignis (BAIRD & GIRARD 1852) occurs, Ameiva chaitzami Stuart 1942a, Proc. Biol. Soc. Washington 55: 143; type locality: along Cahabon-Languin trail about 2 km N Finca Canihor, about 38 km ENE Coban, Alta Verapaz, Guatemala, SVL to 85 mm (males) and 75’ mm (females). Eastern Chiapas, Mexico, as well as eastern and central Guatemala, 800-2000 m elevation. [Ameiva ameiva Ameiva festiva (LICHTENSTEIN & VON MARTENS 1856), Nomenclator Reptilium et Amphibiorum Musei Zoologici Berolinensis: 13; type locality: Veragua, Panama, SVL to 114 mm (ales) and 139 mm (females). Tabasco, Mexico, across all of Central America as far as Colombia, sea level to 1200 m elevation. In Central America, three sub- species occur: Ameiva f festiva (Panama and northern Colombia); Ameiva f° edwardsii Bocourt 1873 (Isthmus of Tehuantepec to | @ Nicaragua); Ameiva f. occidentalis TavLon Ameiva leptophrys 1956 (Costa Rica). 7 ——— Ameiva leptophrys Core 1893, Proc. Amer. — Phil. Soc. 31: 341; type locality: Buenos Aires, Costa Rica. SVL to 133 mm (males) and 129 mm (females), Southeastern Costa Rica to eastern Panama, sea level to 700 m elevation. @ Ameiva quadrilineat Ameiva quadrilineata (HALLOWELL 1861), Proc. Acad. Nat. Sci. Philadelphia 12: 483; type locality: Nicaragua. SVL to 88 mm (males) and 82 mm (females), Southeastern Nicaragua to western Panama, sea level to 1050 m elevation. Ameiva undulata (WikGMaNn 1834), Herpetologiea Mexicana: 27; type locality: Mexico. SVL to 129 mm (males) and 111 mm (females). Nayarit, Mexico, to Costa Rica, sea level to 1500 m elevation. In Central America, six subspecies occur: Ameiva u. gai- geai SMiTn & LAUFE 1946 (northern Yucatan Peninsula), Ameiva u. hartwegi Sori 1940 (Base of the Yucatan Peninsula and northern Honduras), Ameiva u. miadis BarBour & LoveripGe 1929a (Isla del Mafz Grande), Ameiva u. parva Barnour & NoBLE 1915 (Pacific side of Central America), Ameiva u. pulchra Hau.oweLt, 1861 (Honduras to Costa Rica along the Caribbean side), Ameiva w. thomasi SwrrH & Laure 1946 (upper Rio Grijalva Basin, Chiapas). IB Ameiva festiva 156Key to Ameiva 1 a 10-12 longitudinal rows of ventral seales; no obvious enlarged throat scales (Fig. 379a), rather gradual inerease in size posteriorly; scales on the underside of the neckband ' (mesoptychial scales) not obviously enlarged ........ Ameiva ameiva b 8 longitudinal rows of ventral scales; 2-5 throat scales enlarged; scales on the underside of the neckband (mesoptychial seales) obviously enlarged (Fig. 379b-d) .. 2 2 a Anterior throat seales smaller than pos- terior throat scales (Fig. 382c) me b Anterior and posterior throat scales of equal size (Fig. 3791.4) wssssssossesssereeseseeenns 3 a Parietals and frontoparietals separated by one or more scales (Fig. 380a); scales at mid-throat greatly enlarged . Ameiva leptophrys b Parietals in contact with frontoparietals (Fig. 380b); scales at mid-throat only slightly enlarged Ameiva quadrilineata 4 a Scales at mid-throat greatly enlarged and not arranged in a longitudinal row (Fig. 879b);, a pale dorsomedial longitudinal stripe present (may be absent in very large specimens) . Ameiva festiva b Scales at mid-throat greatly enlarged and arranged in a longitudinal row or only slightly enlarged and irregularly arranged (Fig. 379d); without pale dorsomedial longitudinal stripe 5 5 a SVL to 85 mm (males) and 75 mm (fe- males); paravertebral stripes very narrow; dorsolateral spots in males fused with pale dorsolateral stripes ... Ameiva chaitzami b SVL to 129 mm (males) and 111. mm (females); paravertebral stripes wide; dor- solateral ‘spots in males, if present, not fused with pale dorsolateral stripes .. Ameiva undulata | © Ameiva chaitzami a. A. ameiva b.A. festva ©. A. leptophrys dA quadritineate e. A. undulata aA leptophrys b. A. quadrilineata Fig. 380, Dorsal view of head in Ameiva (frontoparietals orange, parietals brown). Further Reading EcHYERNACHT 1971, 157158 Teiidae _ Aspidoscelis Whiptails are the most common lizards in the dry areas of Mexico and Central America, REEDER et al. (2002) proposed a new classification of the whiptails based on a phylogenetic analysis, Because the diverse assemblage of species that was traditionally grouped under the generic name Cnemidophorus was demonstrated to be paraphyletic, REEDER et al. (2002) split the group into several genera. The lemniscatus group retained the generic name Cnemidophorus, whereas the other North American and Central American species were transferred to the resurrec- ted genus Aspidoscelis. The genus Aspidoscelis contains at least 87 currently recognized bisexual and unisexual taxa and is distributed throughout most of North America (except Canada), south through Mexico into lower Central America. In Central America, the genus is represented by six species (DUELLMANN & WELLMAN 1960, VILLA et al. 1988, Lee. 1996). Whiptails are active hunters that will eat everything that they can overpower, most- ly inseets and other arthropods. To the human observer, these lizards scem tre- mendously restless, as they are virtually Fig. 881. Aspidoscelis motaguae Photo: R. D. Bartlett Fig. 382, Aspidoscelis deppii (male above, fe- male below) at Playa Tamarindo, Costa Rica. in constant motion. As heat-loving ani- mals, the whiptails are out in the open pri- marily on hot, sunny days, whereas they retreat to their burrows when the sky is cloudy. Most of the 189 Aspidoscelis deppii examined by Fitcu (1973b) had body tem- peratures from 29-42 °C. Aspidoscelis are most active in the morning. Most of the Aspidoscelis species produce 2-5 clutches per year, each with 1-5 eggs, which they bury superficially in the sand. In the dry season, reproductive activity comes almost. entirely to a standstill, whereas the egg- laying period begins with the onset of the rainfall. The incubation period lasts from 60-80 days. Aspidoscelis deppii becomes sexually mature at the age of 5-6 months (ECHTERNACHT 1983). Worthy of note is the fact that several Aspidoscelis species, including the Central American A. cozu- mela and A. rodecki, reproduce partheno- genetically. In these two species, only females, which produce fertile eggs with- out any previous copulation, are known.Aspidoscelis angusticeps (Cope 1877), Proc. Amer, Phil. Soc. 17: 95; type locality: Yucatén, ‘Mexico. SVL to 115 mm. Yueatén Peninsula, sea level to 100 m elevation. Two subspecies are recognized: Aspidoscelis a. angusticeps (northern part of the Yucatén Peninsula) and Aspidoscelis a. petenensis Brarcik & McCoy 1964 (base of the Yucatan Peninsula). Aspidoscelis cozumela (GAvOw 1906), Proc. Zool. Soc. London 1906: 316; type locality: Isla Cozumel, Quintana Roo, Mexico, SVL to 83 mm, Yueatan Peninsula, sea level to 100 m elevation. Two subspecies are recognized: Aspidoscelis ¢. cozumela (Isla Cozumel) and Aspidoscelis c. maslini (Farris 1969) (Yucatan Peninsula), Aspidoscelis deppii (WIEGMANN 1834), Herpetologica Mexicana: 28; type locality: Mexico. SVL to 93 mm, Northern Veracruz and Guerrero, Mexico to Costa Riea, sea level to 1200 m elevation. Only the nominate sub- species occurs in Central America, Aspidoscelia guttata (WIEGMANN 1834), Herpetologica Mexicana: 29; type locality: Mexico. SVL to 145 mm. Mexico, Veracruz on the Atlantic side and from Guerrero to western Chiapas on the Pacifie side, sea level to 1200 m elevation. Teiidae Aspidoscelis motaguae (SackRTt 1941), Notulae Naturae 77: 1; type locality: Motagua River, Zacapa, Guatemala. SVL to 145 mm, Oaxaca, Mexico, to El Salvador and Honduras, 500-1200 m elevation. Aspidoscelis rodecki (McCoy & Mastin 1962), Copeia 1962: 620; type locality: Isla Mujeres, Quintana Roo, Mexico. SVL to 70 mm. The northeastern portion of the Yucatan Peninsula, sea level to 50 m elevation. Fig. 384. Aspidoscelis angusticeps Photo: R. Cedeito V. Fig. 383. Aspidoscelis cozumelae mastini (Banco Chinchorro Reef, Quintana Roo, Mexico). Photo: H. Bahena B. Fig. 885. Aspidoscelis guttata (male right, fe- male left) near Mixtequilla, Oaxaca, Mexico. 159160 Teiidae Key to Aspidoscelis 1 a 4 supraoculars b 3 supravculars ... 2a Dorsum of juveniles with six pale longitudinal siripes on a dark brown back- ground; in adults (SVL > 100 mm), dorsal stripes usually completely replaced by spots Aspidoscelis motaguae b_Dorsum of juveniles with six or seven pale longitudinal stripes on a black back- ground; adult pattern variable, but some trace of striping usually retained _ Aspidoseelis angusticeps 3 a Frontoparietals divided from parietals by one or more Seales wss.o . 2 b Frontoparictals in contact with ‘osual 4 a Dorsal pattern with indistinct para- vertebral stripes, discontinuous or absent on the posterior body hal... sessre Aspidoscelis rodecki b Dorsal pattern with four distinct contin- uous paravertebral stripes. . snninmne Aspidoscelis cozumela 5 a Fewer than 140 scales around midbody, not counting ventral scales sesons sues Aspidoscelis guttata b More than 165 scales around midbody, not counting ventral scales... Aspidoscelis deppit aC. lenmiscatus b. A. motaguae Fig, 386, Dorsal head scalation (supraoculars orange; parietals brown) in Aspidoscelis and Cnemidophorus. Further Reading BurT 1931, DUELLMAN & WELLMAN 1960, DUELLMAN & ZWEIFEL 1962, Mastin & Skcoy 1986, WrichT & Virr 1993, Reever et al, 2002 Fig. 387. Aspidoscelis c. cozumelae (Isla Cozumel, Mexico). Photo: R. Cedeno V. Aspidoscelis 4A. cozumela WA. ceppii Aspidoscelis IBA. angusticeps BA. motaguae 1 Aspidoscelis guttataFig. 888, Adult male of Cnemidophorus lemniscatus (Isla de Utila, Honduras). Cnemidophorus REEDER et al. (2000) proposed a new classification for the whiptail lizards and transferred the majority of species from Cnemidophorus to Aspidoscelis. As cur- rently recognized, C. Jemniscatus is the only Central American representative of the genus Cnemidophorus. Cnemidophorus lemniscatus is a typical diurnal inhabitant of the open coastal areas, but will also advance inland along large rivers, hardly ever penetrating the forest. These active lizards are often seen in great numbers. They run a bit, stop ab- ruptly and execute rapid turning motions with their forelegs, which has earned them the name “Shaky-Paw” amongst the inha- bitants of the Islas de la Bahia, Honduras (Wison & Hann 1973, KOHLER 1996d, 19981). Although Cnemidophorus lemnis- catus feed primarily on insects and other arthropods, juveniles have also been observed eating cactus blossoms (Opuntia wentiana) (MIJARES-URRUTIA et al. 1997). C. lemniscatus produce several clutches per year, each with 1-5 eggs, which hatch after about two months (KOHLER 1997), Teiidae Fig. 389, Adult female of Cnemidophorus lemniscatus (Isla de Utila, Honduras). Cnemidophorus lemniscatus (LINNakUS 1758), Systema Naturae, ed. 10: 209; type locality: “Guinea” [probably in error for Guiana]. SVL to 104 mm. Central America from Guatemala as far as northern South America east of the Andes, including many offshore islands, as well as various Caribbean islands (e.g., Trinidad and Tobago), sea level to at least 1000 m elevation, Only the nominate subspecies occurs in Central America, Further Reading KOx Ler 1996d, Cote & Dessaver 1993, Wright & Vitt 1993, Reever et al, 2002, Moyica et al 2008 © Cnemidophorus lemniscatus Jf ( ~~ t, ro . rane Sure 161162 Xantusiidae Xantusiidae The family Xantusiidae consists of three genera (Xantusia, Lepidophyma, Crico- saura), with only Lepidophyma occuring in Central America (BEzY & CAMARILLO 2002). The distribution ranges of most night lizards are extremely fragmented, and populations are often isolated from one another by hundreds of kilometers. Such disjunct distribution has been taken as an indication of the great geological age of the group (Brzy 1972). Lepidophyma ‘The 17 species of the genus Lepidophyma are distributed from Nuevo Le6n in north- eastern Mexico to as far south as Panama. They are bizarre-looking mid-size lizards, which are secretive ground-dwellers that are seldom observed outside their hiding places (beneath decaying tree stumps and logs, in rock crevices and caves). They have no eyelids, rather having, similar to many gecko species, so-called “spectacles”. ‘The sides of the body are decorated with countless, irregularly arranged tubercular scales, whereas the tail is equipped with whorls of slightly enlarged scales. During the day, these lizards hide under rocks and fallen logs, which provide a moist microclimate. These nocturnal ani- mals are sit-and-wait predators whose prey consists mostly of insects and spi- ders. The remarkable thing about this genus is the livebearing method of repro- duction. There are also populations that consist entirely of females that reproduce parthenogenetically. During the months from April to July, the females of L. flavi- maculatum bear 2-5 (seldom as many as 8) young, which have a SVL of 35-38 mm at birth (ALVAREZ DEL Toro 1960, TeLFORD & CAMPBELL 1970). Fig. 390. Lepidophyma smithii (El Refugio, Ahuachapan, E] Salvador). Fig. 391, Lepidophyma flavimaculatum (near Pueblo Wiso, Jinotega, Nicaragua). mL. tuxtlae YL. chicoasense ¥ L. lipetzi © L. mayae 4 L, roticulatum flavimaculatum smithii LepidophymaLepidophyma _chicoasense Atvanrz & VatenriN 1988, Anales de la Escuela Nacional de Ciencia Biologia México 82: 125; type locality: Cation de Sumidero, 16.3 km N, 1.6 km E Tuxtla Gutiérrez, 600 m, Chiapas, Mexico, SVL to 78 mm. Known only from the type locality. Lepidophyma flavimaculatum A. Dumern. 1851, Cat, Méth. Coll. Rept. Mus. Paris: 137; type locality: El Petén, Guatemala. SVL to 95 mm. Veracruz, Mexico, along the Caribbean side of Central America as far south as the Panama Canal, sea level to 750 m elevation Lepidophyma lipetzi SiH & ALVAREZ DEL Toro 1977, J. Herpetol. 11: 37; type locality: Lago de Mal Paso, headwaters of Rio de la Venta, 30 km N Cintalapa (straight line), Chiapas, Mexico. SVL to 65 mm. Known only from the type locality. Lepidophyma mayae Bezy 1973, Los Angeles County Mus. Contrib, Sei. 239: 1; type locali- ty: near Chinajé, 140 m elevation, Alta Verapaz, Guatemala, SVL to 65° mm. Southern El Petén and the lower slopes of the Sierra de los Cuchumatanes and the Sierra Xucaneb (Departments of Huchuetenango and Alta Verapaz), Guatemala; also Maya Mountains of Belize, 100-300 m elevation. Lepidophyma reticulatum TavLor 1955, Univ. Kansas Sci. Bull. 87: 551; type locality: Agua. Buena, Puntarenas, Costa Rica. SVL to 92 mm. Pacific side of Costa Rica, sea level to 1250 m elevation. Lepidophyma smithii Bocourt 1876, Journ. Zool. Paris 5: 402; type locality: Tehuantepec and western Guatemala. SVL to 81 mm. Guerrero, Mexico, along the Pacific side of Central America as far south as El Salvador, sea level to 1200 m elevation. Lepidophyma tuxtlae WeRLER & SHANNON 1957, Herpetologica 13: 119; type locality: lower slopes of Volcan San Martin, 2500 feet, Veracruz, Mexico. SVL to 97 mm. Isthmus of Tehuantepec (discontinuously from the Sierra de las Tuxtlas, Veracruz, Mexico, to the El Ocote region, Chiapas); sea level to 1500 m elevation, Fig, 393. Lepidophyme flavimaculatum (E] cote region, Chiapas, Mexico). Xantusiidae Key to Lepidophyma 1 a 16 or more divided lamellae under the 4t toe Lepidophyma chicoasense b 15 or fewer divided lamellae under the 4th LO ssersessveees i 2 a Paravertcbral scale row relatively homo: geneous, composed of tubercular scales that are similar in size and nearly contin- uous sss Lepidophyma tuxtlae b Paravertebral scale row heterogeneous, composed of tubercular scales of various sizes interspersed with small granular SCALOS sossentsnees ns 8 3a Parietal eye not visible in adults (Fig. 392a), occasionally present in juveniles with a SVL less than 75 mm . 7 . Lepidophyma smithi b Parietal eye visible (Fig. 392b) .. parietal eyo aL. siithit b.L, flavimaculatum Fig. 392. Dorsal head scalation in two species of Lepidophyma: 163164 Xenosauridae ate a. L. mayae b. L. flavimaculatum Fig. 394. Lateral head scalation in two species of Lepidophyma. Scales between 2nd postorbi- tal supralabial and postocular orange. 4 a Dorsal seales (occiput to base of tail) 179 or fewer; more than 10 longitudinal rows of ventrals on posterior half of body Lepidophyma lipetzi bb Dorsal ecales usually 180 or more; if fewer than 180, 10 or fewer longitudinal rows of ventrals’ on posterior half of body eS 5 a 0-1 Scales between second postorbital supralabial and postoculars (Fig. 394a); lateral tubercular scales arranged in 33- AB OWS ones Lepidophyma mayae b 2 or more scales between second post- orbital supralabial and postocular scales (Big. 394b); lateral tubercular scales arranged in 24-32 rows wns a) 6 a Throat region without obvious dark retic- ular markings os Lepidophyma reticulatum b Throat region without obvious dark retic- ular markings, although some diffuse shading may be present ..... Lepidophyma flavimaculatum Further Reading ‘TELFORD & CAMPBELL 1970, BEdy 1972, 1984, 1989, Bezy & CAMARILLO 2002. Fig. 395. Xenosaurus grandis (near Cordoba, Veracruz, Mexico. Photo: R. E. Ballinger Xenosauridae The family of knob-scaled lizards has a fragmented distribution in Mexico and the northern portion of Central America, as well as in China, which shows it to be a relictual group, known from fossils since the Upper Cretaceous, Xenosaurus In Central America, only one knob-scaled lizard species occurs, Xenosaurus grandis. The ability to autotomize and regenerate the tail is not present in the species of the genus Xenosaurus. The typical habitat of Xenosaurus grandis is open rocky land with hiding places in the form of rock erev- ices, Actually, all the X. grandis observed by BaLincER et al. (1995) in Veracruz, Mexico, were in rock crevices; none were found in the open. Only one knob-scaled lizard lives in each crevice, Social rela- tionships and interactions could not be determined in field studies, which leaves the suspicion that these animals are very solitary, Horizontal rock crevices are pre- ferred over vertical ones (BALLINGER et al 1995). The assumption in earlier literature that knob-scaled lizards are nocturnal (KING & THOMPSON 1968) could not be sub- stantiated by more recent field studies. At night, all of the observed X. grandis sleptFig. 396, Xenosaurus grandis. Photo: R. D. Bartlett with their eyes closed and did not react to light stimulation (BALLINGER et al. 1995). The first activities (e.g. opening the eyes, Jooking out of the rock crevice) were deter- mined to take place about two hours after sunrise, Xenosaurus grandis belongs to the thigmothermic lizards, which make use of the heat in the surrounding sub- strate, but not the heat of the sun’s rays. The body temperature of active knob-scaled lizards corresponds to the temperature of the surrounding rock and lies between 20.0-25.8°C (average 22.7°C); (BALLINGER et al. 1995). In general, these lizards are not shy and seem almost lethargic. The radius of activity is reported to be less than 1 m (ALVAREZ DEL ToRO 1960). If a knob-scaled lizard is grabbed, it will spray a considerable amount of fluid from the cloacal bladder as a defense mechanism, similar to many frogs and toads. Knob- scaled lizards feed primarily on many types of insects, but will occasionally eat small lizards (PRESCH 1981, BALLINGER et al. 1995). Xenosaurus grandis reproduces ovoviviparously, with litters containing 3-7 young (Frirrs 1966), ALVAREZ DEL TORO (1960) reported that births among X. gran- dis occur in Chiapas, Mexico, from March to July, and that newborns have a total length of 45 mm. Xenosauridae Xenosaurus grandis (Gray 1856), Ann. Mag. Nat. Hist. (2) 18: 270; type locality: Cérdoba, Veracruz, Mexico. SVL, to 116 mm. Disjunet populations from central Veracruz and Oaxaca, Mexico, to Guatemala, 750-1800 m elevation, Two subspecies occur in Central America: Xenosaurus g. grandis (Veracruz and Oaxaca) and Xenosaurus g. rackhami Star? 1941b (Chiapas and Guatemala). Fig. 97. Xenosaurus grandis (101 Ocote region, Chiapas, Mexico). Photo: A. Ramirez V. @ Xenosaurus grandis Further Reading Fritts 1966, Presc 1981, BALLINGER et al. 1995.Amphisbaenia Worm Lizards (Amphisbaenia) Although the taxonomic position of the worm lizards has yet to be satisfactorily resolved, this group is generally regarded as an independent suborder related to the lizards (Sauria) and snakes (Serpentes). Evidence exists that the worm lizards had already separated from the snakes and lizards (order Squamata) during the Cretaceous period, even prior to the rise of modern lizard families (GANS 1969, 1978). Earlier authors had considered the worm lizards to be a lizard family (VANZOLINI 1951). A number of anatomical characters distinguish the worm lizards from both the snakes and the lizards. These include the reduction of the right lobe of the lung and the existence of a middle pre-maxil- lary tooth, as well as a characteristic skull and middle ear morphology (Gans 1969). Worm lizards are the true burrowers among the reptiles. Living in self-con- structed, permanent tunnel systems, amphisbaenians have an elongated body and, with the exception of one genus (Bipes), no external limbs. Rather than use already existing tunnels as worm snakes will, or staying in loose soil or sand, they are in the habit of tunneling through com- pact earth. They seldom come to the sur- face and then only at night. Frequently, | 1B Amphisbaena fuliginosa { ¥ Amphisbaena spurrelli 166 they can be found in the underground colo- nies of leaf-cutter ants (GANS 1969). Worm. lizards are equally able to crawl back- wards and forwards in their tunnel: Amphisbaenids are skilled hunters, prey- ing upon insects, spiders, and rodents, as well as worm snakes and other reptiles (Gans 1969). Amphisbaena alba LiNNagus 1758, Systema Naturae, ed. 10: 229; type locality: America. SVL to 670 mm. Tropical South America east of the Andes. Occurrence in Panama ques- tionable. Amphisbaena fuliginosa LINNAEUS 1758, Systema Naturae, ed. 10: 229; type locality: America. SVL to 500 mm, Panama, tropical South America cast of the Andes, as well as western Colombia and Ecuador, Amphisbaena spurrelli (BouLENGER 1915), Proc. Zool, Soc, London 1915: 659; type locali- ty: Andagoya, at junetion of Rios Condoto and San Juan, Colombia. SVL to 300 mm Panama, northern Colombia, and Venezuela. Fig. 398. Amphisbaena fuliginosa (Peru), Photo: B. LehrKey to Amphisbaenia 1 a 65 or more scales at midbody seine é . .. Amphisbaena alba b 55 or fewer scales at midbody .. 2 2 a Irregular pale-dark spotting; more than four precloacal pores... . Amphisbaena fuliginosa b Without spotting; four precloacal pores = . Amphisbaena spurrelli Further Reading VANZOLINI 1951, Gans 1962a, b, 1967, Hooc- mogD 1973, GANS & MarieRs 1977, Gans 1978 prefrontal ocular temporal rostral 1 supralabials AE nasal infralabials prefrontal ocular ps “a Fig. 399. Amphisbaena spurrelli. Modified from BOULENGER (1915). =eeees Amphisbaenia Fig. 400. Amphisbacna alba. Photo: R. W. Van Devender Fig. 401. Amphisbaena alba Photo: R. W. Van Devender Fig. 402. Amphisbaena fuliginosa (Nusagandi, Comarea de San Blas, Panama). 167168 Serpentes Snakes (Serpentes) anterior temporal posterior temporals Fig. 403, Scalation characters on the head of a colubrid snake, infratabials Parietal efrontal aN In Central America, the snakes are well represented by 276 different species. Key to Snake Families 1 a Tail conspicuously flattened laterally (Fig. 685) ...... Elapidae (in part: sea snakes) b Tail more or less round in cross-section 2 2 a Ventral scales not distinctly enlarged, scales around body all approximately of equal 8126 vss teeenstniceoeiciee: b Ventral scales distinctly larger than dorsal scales ee 5 3 a 14 scale rows at midbOdY weds ; asa Leptotyphiopidae b 18 or more scale rows at midbOdY ss 4 4 a 18-20 scale rows at midbody; one preocu- lar; no subocular ..........1.:'Typhlopidae b 21-22 scale rows at midbody; preocular present or not; subocular present or not ‘Anomalepididae 5 a One tube-like loreal pit between nostril and eY€ «0. vee oo iperidae b Without tube-like loreal pit between nostril and €Y€ o.sscscsneseseeeee 6 postoculars ‘supraocular prefrontal Internasal rostral prenasal mental postnasal preocular supralabials ‘4st dorsal row ventrals ( internasal infralabials rage Ce chin shields: eS @ ‘mental "> — = frontal a mental groove Ventrals do not extend across the entire body width (one or more dorsal rows on the ventral side); usually all or most sub- caudals undivided (exception: Loxocemus); often with cloacal spurs on either side of the cloaca .. sence sseeerscone T Ventrals extend across the entire body width (no dorsals on the ventral side); usually all or most subcaudals paired (exception: Pseudoboa); no cloacal spurs PROSONE oo .cessescessssscesessesesesee 9 All or most subeaudals paired . . Loxocemidae b All or most subcaudals undivided ...........8 Fewer than 35 dorsal rows at midbody eons innsnnerestnsiensenee Beopidophiidae More than 35 dorsal rows at midbody A seer . Boidae Upper jaw with 2 immobile, enlarged teeth, otherwise, upper jaw without teeth; no loreal; body pattern with complete black rings ... — . Elapidae (in part: coral snakes) Upper jaw with many teeth, none conspi uously enlarged in front of upper jaw; loreal present or not; coloration and mark: ings variable ColubridaeScolecophidia Blind Snakes (Scolecophidia) pretronial rostral ocular mental Fig. 404. Scalation characters on the head of a blind snake. Three families of comparatively primitive snakes are grouped under the term blind snakes (Seolecophidia). They are charac- terized by shiny, smooth scales of fairly uniform size surrounding the entire body. Fig. 405. Anomalepis mexicanus (Maritza, Guanacaste, Costa Rica). Photo: M. Franzen Anomalepididae The family Anomalepididae is made up of four genera (Anomalepis, Helminthophis, Liotyphlops, and Typhlophis; the last not in Central America), with a total of 15 spe- cies, Little is known about the habits of these small underground-dwelling snakes. In Central America, the genera Anomalepis, Helminthophis, and Liotyphlops are each represented by a single species. Key to Anomalepididae 1 a Dorsal surface of head with two large pre- frontals behind the rostral, broadly in contact with each other and followed posteriorly by a large frontal (Fig. 406s)... seve . . Anomalepis b Scalation on dorsal surface of head diffe- rent than that above isons ennnes aan 2 Prefrontals behind the rostral in contact with each other (Fig. 406b) Helminthophis b Prefrontals not in contact with each other, separated by conspicuously large rostral (Fig. 406¢) ».. Liotyphlops b. Helminthophis a. Anomalepis Fig. 406. Dorsal head scales (prefrontal yellow; parietal brown) ©. Liotyphiops Anomalepis ‘The genus Anomalepis includes four spe- cies (KorRON 1988a), one of which occurs in southern Central America. Anomalepis mexicanus is oviparous, as verified by the dissection of a female, which contained two eggs (KOFRON 1988a) 169170 Anomalepididae Anomalepis mexicanus JAN 1861, Icon. Gén. Livr, 1: pl. 5, fig. 1, pl. 6, fig. 1; type “Mexico” [in error]. TL to 153 mm. Northwestern. Costa Rica and central Panama; also in Peru, 300-500 m elevation; distribution in Colombia and Ecuador not yet demonstrated (KOFRON 1988a). Further Reading TavLOR 1939, DUNN 1941, Ros & SmirH 1966, KOFRON 1988a, Fig. 407. Helminthophis frontalis (San José, Costa Rica). Photo: R.D. Bartlett Helminthophis Little is known about Helminthophis fron- talis, other than that it lives as a secretive ground dweller, It is distributed in south- ern Central America, 4 Anomalepis mexicanus ( ¥ Helminthophis frontalis 1B Liotyphiops albirosiris Fig, 408, Liotyphlops albirostris (Isla Pachaca, Pearl Islands, Panama), Photo: C. W. Myers Helminthophis frontalis (PBTERS 1860), Monats, Akad. Wiss. Berlin 1860: 517; type locality: Costa Rica. TL to 160 mm. Central Costa Rica and southwestern Panama, 90- 1440 m elevation. Further Reading AMARAL 1924a, WALLACH & GUNTHER 1997. Liotyphlops According to the most recent revision of Liotyphlops by Dixon & Korron (1984), this genus is represented in Central America solely by the species L. albiros- tris. The other seven species of this genus are distributed in South America. Liotyphlops albirostris (Peters 1857), Monats. Akad. Wiss. Berlin 1857: 402; type locality: Panama. TL to 223 mm. Southern Central America (southeastern Costa Rica, Panama) and northwestern South America (Ecuador, Colombia, Venezuela), as well as on Curacao (Netherlands Antilles), near sea level to 1640 m elevation. Further Reading Dixon & KorKon 1984.Leptotyphlopidae Leptotyphlopidae ‘The family Leptotyphlopidae contains two genera (Leptotyphlops and Rhinoleptus) with a total of 80 species worldwide. Two species of Leptotyphlops occur in Central America. Leptotyphlops The taxonomy of the Central American [aes ie : Leptotyphlops is poorly understood. A a Sa KOHLER (2001a,b) and SAVAGE (2002) treat- Fig. 409. Leptotyphlops goudotii (Isla de ed Leptotyphlops nasalis and L. ater, re- — Ganaja, Honduras). spectively, as valid species. However, these decisions appear to be only weakly support- ed and I prefer to not recognize these taxa until a comprehensive review of the Central American Leptotyphlops has been published. Leptotyphiops nasalis has been described based on a single specimen from Managua, Nicaragua, which differs from the widely distributed species L. goudotii by a single distinguishing scale character, which might also be due to individual anomaly. Since L. nasalis was first describ- ed over 60 years ago, no further specimens that could be assigned to this taxon have been found. Leptotyphlops goudotii is the most common Leptotyphlops found in es Central America. These not even pencil- Fig. 410. Leptotyphlops goudotii can be found thick snakes have a bright yellow spot on iM termite nests, both the anterior and the posterior end. As the very small head is not well differentia- ted from the body, it is difficult at first glance to distinguish the head from the tail tip. These blind snakes live secretively in rotting logs and in the mounds of termites or ants, which make up the bulk of their food intake (KOHLER 1998f). These little snakes most often eat their prey whole. However, they also have been observed sucking out the soft body contents of insects and leaving behind empty exoskeletons (Scorr 1983c). Leptotyphlops goudotii are oviparous and produce clutches of 8-12 eggs (ALVAREZ DEL TORO 1983), Fig 411. Leptotyphlops macrole Colombia). Photo: M. Lundberg Snakes172 Leptotyphlopidae === Leptotyphlops goudotii (Dum#nt, & BIBRON 1844), rp. Gén. 6: 330; type locality: Valley of Rio Magdalena, Colombia. TL to 185 mm Colima and Veracruz, Mexico, through Central America to Colombia and Venezuela, sea level to 1600 m elevation in dry forest and wet forest. Leptotyphlops macrolepis (Psvexs 1857), Monats. Akad. Wiss. Berlin 1857: 402; type locality: Caracas and Puerto Cabello, Venezuela. TL to 400 mm. Panama and north- ern South America. Key to Leptotyphlops 1 a Without pale longitudinal lines (Fig, 411); supraoculars about as large as prefrontal and frontal ... Leptotyphlops macrolepis b Pale longitudinal lines present (Fig. 410); supraoculars somewhat larger than pre~ frontal and frontal... itn .. Leptotyphiops goudotii Further Reading Hann 1979, Scorr 1983¢, KOHLER 2000b. Cia S aR. braminus b. T. costaricensis_«. T. tenuis, Fig, 412. Lateral head scalation in Typhlops and Ramphotyphiops (rostral brown, prefron- tals yellow). 1B Leptotyphlops goudotii Hi Lepiotyphlops macrolepis Typhlopidae The family Typhlopidae is represented by two genera in Central America, one of which (Ramphotyphlops) has been intro- duced in several countries, The species in this family only have teeth on the upper jaw and not on the lower jaw, unlike the Leptotyphlopidae and Anomalepididae. Similar to these species, however, they are also small, worm-like snakes that live very secretively underground. Their dorsal and ventral seales are similarly smooth and of equal size. Key to Typhlopidae 1a More than 345 dorsal scales between rostral and tail spike; scale suture between nostril and rostral on a diagonal (Fig. 412b.) a Typhlops b Fewer than 245 dorsal scales between rostral and tail spike; scale suture between nostril and rostral almost vertical (Fig. 412a) 0. ... Ramphotyphlops 413. Dorsal head scalation in Typhlops costaricensis (rostral brown, prefrontals yellow). \ [%e I Typhlops tenuis {ll Typhiops stadelmani © Typhlops microstomus ‘4 Typhlops costaricensisFig. 414. Ramphotyphlops braminus. Photo: R. D. Bartlett Ramphotyphlops Ramphotyphlops braminus is a species native to Southeast Asia, but has been introduced throughout the tropics world- wide. In Central America, this species has been reported in Mexico, El Salvador and Guatemala, where it has recently become indigenous (Dixon & HeNpRIcKs 1979, LEE 1996, DuENas et al. 2001). It is assumed that this small snake is transported into foreign countries along with the soil on plants. As it is parthenogenetic, only a single female is required to establish a new population (Nussbaum 1980), Rampho- typhlops braminus produces clutches of 2- 3, possibly up to 6 eggs (Nussraum 1980). Ramphotyphiops braminus (DaUDIN 1803), Hist. Nat. Rept. VII: 279; type locality: Vizagapatam, India, TL to 173 mm Southeast Asia, Africa, Madagascar, Indian Ocean, Hawaii, South Pacific and Australia; introduced in ‘several locations in Central America, Further Reading Dixon & HenpRics 1979, Nussaaumt 1980, Ora et al. 1991 Typhlopidae Typhlops The genus Typhlops is found throughout the tropics and subtropics worldwide. In Central America, there are four species of this genus, one of which (7. stadelmani) has only recently been resurrected (McCranié & WILSON 2001b). These blind snakes live very secretively buried under- ground, where they feed on termites, ants, and other small insects. The Central American species of this genus are ovi- parous. Fig. 415. Typhlops stadelmani (near Quebrada Grande, Copan, Honduras), Fig, 416. Typhlops microstomus (Quintana Roo, Mexico). Photo: H. Bahena B. Snakes 173174 ‘Typhlops Typhlops costaricensis Jiménez & SAvaGE 1962, Rev. Biol. Trop. 10: 199; type localit Monteverde, Sierra de Tilaran, Puntarenas, Costa Rica, 1500 m. TL to 360 mm. Central Honduras to Costa Rica, . Typhlops microstomus Core 1866, Proc. Acad. Nat. Sci. Philadelphia 18: 125; type locality: Yucatan, Mexico. TL to 366 mm. Central Veracruz, Mexico, as well as the Yucatén Peninsula. Typhlops stadelmani Scumipt 1936a, Proc. Biol. Soc. Washington 49: 48; type locality: Subirana Valley, Yoro, Honduras. TL to 310 mm. Northwestern Copan and southwestern Yoro, Honduras, 850 to 1370 m elevation in wet forest, ‘Typhlops tenuis SALVIN 1861a, Proc. Zool. Soc. London 1860: 545; type locality: Coban, Alta Verapaz, Guatemala. TL to 326 mm. Veracruz, Mexico, to Guatemala, sea level to 1500 m elevation in dry forest and wet forest. Macrostomata According to McD1armip et al. (1999), six families of early snakes (eg., Boidae, Bolyeriidae, Loxocemidae, Pythonidae, ‘Tropidophiidae, and Xenopeltidae) are included in the Macrostomata. Boidae The species of the family Boidae are wide- ly distributed in the tropics and subtropics of the Old and New World. Characteristic of the boids are the head, which is clearly differentiated from the neck, vertically slit pupils, remnants of the pelvic girdle (which in some species is recognizable by externally visible cloacal spurs) as well as the presence of functional lungs on both the left and right sides. All boids are non- venomous and kill by constriction, using the muscular coils of their body to suffo- cate prey. The jaws are equipped with strong teeth. All boids are live bearers. Key to Typhlops 1 a 20 scales at midbody . .. Typhlops costaricensis bb 18 soales at MQbOAY sssessssutssem 2 2 a More than 470 dorsal scales between rostral and tail spike; subocular present Typhlops microstomus b Fewer than 450 dorsal scales between rostral and tail spike; subocular absent ... 3 3 a 361-441 dorsal scales between rostral and tail spike; body with brown spots su Typhlops tenuis dorsal scales between rostral and tail spike; body solid color pink (in life) and pale brown (in preservative)... Typhlops stadelmani Further Reading Vita 1978, Dixon & McCrae & WiLson 2001b Henpricks 1979, Fig. 417. Boa constrictor (Bartola, Rio San Juan, Nicaragua) Further Reading KLUGE 1991, ZAHER 1994Boidae Fig. 418. Boa consirictor (Isla de Guanaja, Honduras) Key to Boidae 1 a All scales on dorsal surface of head small and irregular, no enlarged scales on the snout; no labial pits Boa b Snout with several enlarged shields; | al pits present 2 2 a More than 245 ventrals; more than 65 sub- als; usually 3, rarely 2 loreals Corailus b Fewer than 240 ventrals; fewer than 65 subcaudals; usually one large loreal, sel: dom 2 loreals Epicrates Boa Boa constrictor is potentially the largest as well as the best-known snake in Central America. It is a large, heavy- bodied snake that can be found both on the ground as well as in the branches of large trees. Although it is predominantly cre- puseular and nocturnal, it is occasionally encountered outside of its hiding places during the day. Typical daytime hiding places include hollow trees, as well as piles of leaf litter among the buttre large trees. Boas are not too particular about their vertebrate prey and will con- sume any lizard, bird, or mammal they can swallow. If they feel threatened, these well-armed snakes will hiss with their mouths wide open, and they are capable of inflicting bloody and painful injuries with their powerful jaws, Often pieces of the long teeth will break off and remain lodged in the wound. Boa constrictor is live bearer and produces litters of 10-64 young (GREENE 1983), On September 2, 1997, I discovered a female Boa constrictor slightly more than 180 cm in total length in the hollow stump of a dead coconut palm on Utila. The snake was coiled up and had obviously just given birth, as there were 13 newborn snakes, with the remains of umbilical cords, in the the hole. The total length of the young boas was between 49 and 52 cm, their weight around 52 g. roots of back of Snakes 175176 Boidae Boa constrictor LinNAEUS 1758, Systema Naturae, ed. 10: 215; type locality: “India” [in error]. TL to 4450 mm (WaTKINS-COLWELL & LeeNnverS 2003); however, most specimens measure less than two meters). Tamaulipas and Sonora, Mexico, to Peru, Bolivia, and Argentina, as well as in the Lesser Antilles, sea level to 1000 m elevation in dry forest, wet forest, and rain forest as well as man- groves. In Central America, the subspecies Boa ¢. imperator DavpiN’ 1803 (Mexico, Central America excluding Toboga Island, Panama, and northwestern South America) and Boa c. sabogae (BARBOUR 1906) (Toboga Island, Panama) occur. Further Reading STULL 1935, LAZELL 1964, GREENE 1983, WILSON & MEYER 1985 ‘@ Boa constrictor Corallus There are seven different species in the genus Coraillus, all of which are crepuscu- Jar and nocturnal tree dwellers in rain forest areas. The two Central American species are extremely variable in terms of coloration and pattern; this is particularly the case with Corallus ruschenbergerit. Corallus species will consume frogs, lizards, small mammals, and birds, which they kill by constriction. Corallus are usually very aggressive, and a bite from their particularly long front teeth can pro- duce a deep wound. These snakes are live- bearers, producing litters of 7 to 30 young (Ross & Marzec 1990). Corallus annulatus (Cove 1876), J. Acad. Nat. Sci. Philadelphia (2) 8: 129; type locality: Costa Rica. TL to 1335 mm. Disjunet from eastern Guatemala to southwestern Ecuador, sea level to 400 m elevation in rain forest. Corallus ruschendergerii (Core 1876), J. ‘Acad. Nat. Sci. Philadelphia (2) 8: 129; type locality: Panama, TL to 870 mm. South- western Costa Rica to Panama, as well as across northern South America east of the ‘andes, sea level to 1000 m elevation in rain forest. Key to Corallus 1 a Nasal shields separated from one another by rostral; seales posterior to nasal shields not enlarged fewer than 90 subcaudals Corallus annulatus in contact with one another; scales posterior to nasal shields usually distinctly enlarged; more than 90 sub- caudals .......... Corallus ruschenbergerii © & a. C. annulatus b. C ruschenbergerii Fig. 419. Snout scalation in Corailus (nasal shields orange). Further Reading HENDERSON 1993a-c, StarrorD & HENDERSON 1996, HENDERSON 1997, E. Smrrn & AcrvEpo 1997, HENDERSON 2002. 1B Corallus annulatus Corallus ruschenbergeniBoidae Fig. 420, C ruschenber (Cérdoba, Colombia), Photo: M. Lundberg. Fig. 421, Coral Limén, Costa Ri near Tortuguero, Photo: M. Burg Fig. 423. Corallus annulatus (Limén Provine Costa Rica) Photo: R.W, Van Devender178 Boidae Epicrates Only one species of the genus Epicrates is found in southern Central America, E. cen- chria, whose common name, “Rainbow Boa”, comes from its iridescent coloration. It is a slim, arboreal, crepuscular, and noc- turnal species found in the rain forest. It is a livebearer (15-28 per litter; Ross & Marzec 1990) that will eat frogs, lizards, and small mammals. Epicrates cenchria (LiNNaBUs 1758), Systema Naturae, ed. 10: 215; type locality: Surinam, TL to 1500 mm. Costa Rica and Panama, as well as widely distributed through tropical South America; sea level to 500 m elevation in rain forest. The subspecies Epicrates ¢ maurus GRAY 1849a occurs in Central Ameri Further Reading KLUGE 1989. Fig. 424. Epicrates cenchria (Guanacaste, Costa Rica), Photo: R. W. Van Devender Fig. 425. Loxocemus bicolor from Costa Rica. Insert: L. bicolor from Mexico (Playa Azul, Oaxaca), Tr / Epicrates cenchria @~*\y N Loxocemus bicolorLoxocemidae Loxocemus The Mexican burrowing python (Loxocemus bicolor) is the only species in the genus. It lives in dry forest and savan- na habitat and can be seen both during the day as well as at night outside of its hiding place (personal observation). It digs underground passages with its raised and pointed snout and hunts whiptails (Aspidoscelis) and small mammals. This snake is also well known for successfully tracking down and raiding iguana nests (Mora 1987). Loxocemus bicolor reprodu- ces oviparously, with the largest documen- ted clutch containing four eggs (Ross & MarzeEc 1990). Loxocemus bicolor Core 18610, Proc. Acad, Nat. Sci. Philadelphia 18: 77; type locality: unknown, TL to 1530 mm. Nayarit, Mexico, along the Pacific coast to northwestern Costa Rica; isolated occurrences also on the Caribbean side, sea level to 600 m elevation in dry forest and savanna, Further Reading Ne.son & MEYER 1967 Tropidophiidae The species of the family Tropidophiidae are small snakes that are distributed in southern Mexico, Central America to northwestern Pacific South America, and on the West Indies. Contrary to an earlier arrangement in which these snakes were included as a subfamily (Tropidophiinae) in the Boidae, the tropidophiids are now recognized as a distinct family (GREENE 1997, McDiarmi et al 1999). Key to Tropidophiidae 1 a Dorsals keeled; more than 27 dorsal rows at midbody Trachyboa b Dorsal smooth; fewer than 27 dorsal rows at midbody «0.0.0... . Ungaliophis Tropidophiidae Trachyboa Little is known about the two species of the genus Trachyboa. In Ecuador, together with R. Seip, I found a specimen of T. boulengeri that was lying well camoufla- ged and motionless on the ground during the day. The animal behaved quite calmly and made no attempt to flee. Rather, it seemed to stiffen into whatever shape it was manipulated. The species is live bear- ing and produces 5-7 young per litter (Ross & Marzec 1990). Trachyboa_boulengeri_ Peracca 1910, Ann. Mus. Zool, Univ. Napoli 3 (12): 1; type locality: unknown, TL to 430. mm. Choco region of Panama through Pacfic Colombia to western poe sea level to 750 m elevation in rain. forest. ¥ Ungaliophis continentalis 4 Ungaliophis panamensis = Trachyboa 179‘Tropidiphiidae Ungaliophis The dwarf boas of the genus Ungaliophis are represented by two species, which are distributed from Mexico across Central America and into northeastern Colombia. Little is known about these secretive snakes. In Selva Negra, Nicaragua, about two hours after dusk, I discovered an U. continentalis on a tree overgrown with tillandsias. The animal was flicking its tongue as it hung about two meters above the ground. Other observers have reported Ungaliophis as a nocturnal tree-dweller that prefers to live in dense epiphytic growth (CoRN 1974). These are attractive snakes that are completely harmless and make no attempt to bite when handled. A specimen of U. continentalis that was kept at the Senckenberg Museum for several 428. Ungaliophis continentalis (Selva years ate new-born mice. Ungaliophis con- Negra, Matagalpa, Nicaragua), tinentalis bears 5-6 young per litter (Ross & Marzec 1990). Ungaliophis continentalis MOLLER 1882, Verh, Naturforsch. Ges. Basel 7: 142; type locality: Retalhuléu, Guatemala. TL to 482 mm. Chiapas, Mexico, to northern Nicaragua, 1000-2130 m elevation in wet forest and cloud forest Ungaliophis panamensis Scumit 193: Smithson. Mise, Coll. 89 (1): 12; type locality: Cerro Brujo, Panama. TL to 470 mm Southern Nicaragua to western Colombia, sea level to 2100 m elevation in rain forest and cloud forest Key to Ungaliophis 1 a Rostral and prefrontal broadly in contact, therefore, internasals separate from one another; blotches on body sides oval; 25 dorsal rows at midbody, 15 dorsal rows one headlength anterior of cloaca Ungaliophis continentalis b Rostral and prefrontal not in contact, internasals in contact with one anothe blotches on body sides triangular; 19-23 dorsal rows at midbody, 17 dorsal rows one headlength anterior of cloaca . un. Ungatiophis panamensis Further Reading Fig. 427. Ungaliophis panamensis (Tortuguero, BocrRT 1968a, CoRN 1974, Via & Wiison Limén, Costa Photo: M. Burger 1990 180Colubridae Approximately 80% of the known snake species belong to the family Colubridae. This family is distributed worldwide and, on almost all continents, it is the dominant family in terms of both the number of spe- cies and their frequency. The one exception is Australia, where the cobras, kraits and elapids (family Elapidae) are most com- monly represented. There is some contro- versy regarding the intrafamilial structure of Colubridae. In Central America, 64 genera from this family are known. The many species (divided among 270 genera worldwide) of the family Colubridae dis- play tremendous morphological and ecolog- ical variety. Thus, in Central America, there are slenderly built climbing species, (Oxybelis spp., Leptophis spp., Imantodes spp), ground-dwellers with stocky (Xenodon rabdocephalus) or very slim body shape (Leptodrymus pulcherrimus) and species that live in streams or swamps (Tretanorhinus nigroluteus, Hydromor phus concolor, Nerodia rhombifera). Some species reach over two meters in total length (Clelia clelia, Leptophis ahaetulla, Masticophis mentovarius), while others are fully grown at less than 30 em (e.g. Geophis hoffmanni, Tantilla schistosa, Tantillita lintoni) Some Colubrids have enlarged grooved fangs in the back of the upper jaw and pro- duce a secretion that is moderately venom- ous (Conophis lineatus, Oxybelis spp.) Although the colubrid snakes of Central America do not pose any mortal danger to humans, their bite can lead to serious localized swelling and considerable pain. Colubridae Fig, 429, Dipsas brevifacies (southeast of Coba, Quintana Roo, Mexico). Photo: J. C. Lee Fig. 480. Oxybelis aeneus (Isla de Roatan, Honduras) Snakes Fig. 431. Nerodia rhombifera on branches Photo: P. Kornacker 181182 Colubridae Key to Colubridae 1 a Dorsal surface of head covered with small, irregular seales (Fig. 432) ....... Nothopsis b Dorsal surface of head covered with large symmetrical shields (Fig. 433) ...o.00. 2 a Rostral shield protruding and pointed (Fig. 438)... me scence b Rostral shield normal (Fig. 433) 4 3 a Fewer than 160 ventrals; cloacal scute divided (Fig. 437b); no loreal Ficimia b More than 165 ventrals; cloacal scute undivided; loreal present ....... Phimophis Dorsal scales in even number of rows at midbody (compare Fig. 436) «os... 5 b Dorsal scales in uneven number of rows at midbody 6 10-12 dorsal rows at midbody; fewer than 170 ventrals; cloaeal scute usually divided (Big, 487) oeesesee sesso Chironius b 14-18 dorsal rows at midbody, more than 190 ventrals; cloacal scute undivided (Fig. 4870). .». Spilotes 6 a Without longitudinal mental groove (Fig. BB) coer 7 Dipsas b Longitudinal mental groove present.........7 7 a Body extremely slender; head large and distinet from thin neck (Fig. 434a); with large protruding eyes; 195-262 ventrals; 106-178 subcaudals s,s... Imantodes b Combination of characters different from that above snsaoeccec eats By Number of dorsal rows constant from mid- body to cloaca, no reduction oo... D b Number of dorsal rows one headlength anterior to cloaca at least two rows fewer than at midbody arvana ST Cloacal scute undivided (Fig. 4372) «1... 10 b Cloacal scute divided (Fig. 437b) ... 17 No anterior temporal col b One or two anterior temporals soe AZ 13 dorsal rows at midbody Sibon (in part) b 15-17 dorsal rows at midbody...... Geophis Dorsal scales strongly keeled Ninia b Dorsal scales smooth or only slightly keeled ce 18 Unicolor black (adults) or red with pale neck band (juveniles) ...... Clelia (in part) b Coloration different from above .... 14 Preocular present; more than 210 ventrals Siphlophis Fig. 432. Nothopsis rugosus; note the small irregular scales on top of head. Photo: R.W. Van Devender Fig. 433. Dryadophis melanolomus; note the large, symmetrical shields on top of head. e0-- a. Imantodes Riser b. Oxyrhopus open Se Fig. 435. Ficimia public (from DuMERIL et al, 1870-1909) Fig. 434. Head shape in two colubrids Fig. 436. How to count dorsal scale rows; the example shows 21 dorsal scale rows in a Rhadinaea.lia 18a b 19a Usually without preocular; fewer than 210 ventrals ... eos 15 13-15 dorsal rows at midbody Sibon (in part 17-19 dorsal rows at midbody ... 16 Sixth supralabial not conspicuously en- larged, not in contact with postocular, an- terior and posterior temporal; fewer than 40 subeaudals ... .. Atractus Sixth supralabial enlarged and in contact with postocular, anterior and posterior temporal; more than 40 subcaudals ..... osssetmmn Tropidedipsas Dorsal scales keeled (Figs. 441¢,d) Storeria Dorsal scales smooth (Figs. 441a,b) ....... 18 Prenasal fused with bordering internasal (Fig. 438) . 19 Prenasal not fused with bordering interna- sal ... ae 2 20 17 dorsal rows at midbody; fewer than 55 subcaudals : . Stenorrhina 15 dorsal rows at midbody; more than 90 subeaudals Symphimus (in part) Body with complete black rings 0.20 Body without complete black rings, at most dark bands that are not closed on the ven tral side 17 dorsal rows at midbody; fewer than 150 ventrals sun Pliocereus 15 dorsal rows at midbody; more than 160 ventrals ee 22 One posterior temporal; black rings at regular intervals; in life, a bright orange- red dorsal blotch between each of the black rings (Fig, 440); in preservative, without these blotehes, only with black and white rings ... Seolecophis ‘Two posterior temporals; black rings not at regular intervals, either a pair of black rings or rings set somewhat irregularly or partially along the vertebral line; without orange-red blotch dorsally between the black rings in life .. Erythrolamprus 15 dorsal rows at midbody = 24 17-21 dorsal rows at midbody ......0:.s10: 29 Preocular and loreal present, therefore, at least three scales between nostril and eye 25 fither preocular or loreal missing, there- fore, only two seales between nostril and eye 26 Colubridae Fig. 437 Subcaudals can be paired (as shown) or unpaired, Fig. 438. Stenorrhina degenhardtii; note that the prenasal is fused to the adjacent internasal (orange). Fig. 439. Ninia sebae; the loreal (orange) reaches to the eye; there is no preocular. Fig. 440. Seolecophis atrocinctus (Nicaragua). a, Stenorrhina b. Dryadophis . Senticolis d. Thamnophis Fig 441. Dorsal scales a, smooth, without apical scale pits b, smooth, with two apical seale pits c. keeled, with two apical scale pits d. keeled, without apical scale pits 183