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BULLETIN OF THE Museum of Comparative Zoology SPECIAL PUBLICATIONS SERIES BOARD OF EDITORS Editor: Jonathan Losos Managing Editor: Catherine Weisel Associate Editors: Andrew Biewener, Scott Edwards, Brian Farrell, Gonzalo Giribet, James Hanken, Hopi Hoekstra, George Lauder, James McCarthy, Naomi Pierce, Stephanie Pierce, Mansi Srivastava and Robert Woollacott Publications Issued or Distributed by the Museum of Comparative Zoology Harvard University Bulletin 1863- Breviora 1952- Memoirs 1865-1938 Johnsonia, Department of Mollusks, 1941-1974 Occasional Papers on Mollusks, 1945- General queries, questions about author guidelines, or permissions for MCZ Publications should be directed to the editorial assistant: MCZ Publications Museum of Comparative Zoology Harvard University 26 Oxford Street Cambridge, MA 02138 [email protected] EXCHANGES AND REPRINTS All of our publications are offered for free on our website: http://www.mcz.harvard.edu/Publications/index.html To purchase individual reprints or to join our exchange program, please contact April Mullins at the Ernst Mayr Library: [email protected]. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2018. The Lizards, Crocodiles, and Turtles of Honduras Systematics, Distribution, and Conservation JAMES R. MCCRANIE Bulletin of the Museum of Comparative Zoology Special Publications Series, No. 2 All photos and images are by the author unless otherwise credited. The Lizards, Crocodiles, and Turtles of Honduras Systematics, Distribution, and Conservation JAMES R. MCCRANIE Harvard University Museum of Comparative Zoology 2018 Distributed by Harvard University Press Cambridge, Massachusetts, and London, England Copyright © 2018 by the President and Fellows of Harvard College Printed in the United States of America ISBN 978-0-674-98416-5 PREFACE suspected represented new cryptic species. When I started working seriously on the Those suspicions came from carefully ex¬ taxonomy and geographic distribution of the amining and taking morphological data for amphibians and reptiles of Honduras in some specimens in my personal collection 1976, pristine forest was easily reachable and in U.S. museums. Those plans of either by car, a four-wheel-drive vehicle, discovering new species were stopped by walking, or by riding horses or mules. Those the current governmental agency in 2014 days are gone forever, as very little forest in when they began to deny new collecting good condition remains in the country permits. today. Deforestation began to become I need to stress that my taxonomic and serious during the early 1980s. That defor¬ geographical distribution work requires estation has accelerated to the point where fieldwork and the need to collect select at least 90% of pristine forest that was in the species at select localities. The days of country in 1976 is now lost. The forest “general collecting” are long gone, and the devastation began to increase significantly in few select specimens I have collected over the last few years (around 2005) under the the last 10 years (up to 2013) can in no way last years of control of Administracion damage the populations of the few species Forestal del Estado, Corporacion Hon- involved, especially when compared with durena de Desarrollo Forestal, Tegucigalpa, the accelerated habitat destruction that is the Honduran group responsible for pro¬ going on in Honduras. My collecting only tection of its forest and fauna until 2008. requires select species to continue to gain That deforestation has drastically increased knowledge of species taxonomy and distri¬ under the watch of the Instituto Nacional de bution in the country. During the most 10 Conservacion y Desarrollo Forestal, Areas recent years, most individuals of most Protegidas y Vida Silvestre (ICF), Teguci¬ species have been released on spot. My galpa, that took over in March 2008, even in only desire is to collect specimens from the World Heritage Biosphere Reserve Rio species complexes that are likely unde¬ Platano (Reserva del Hombre y la Biosfera scribed, before they become extinct when del Rio Platano). Flying from La Ceiba to their few remaining forested habitats are Puerto Lempira will take you above the gone. Looking back at my earlier work on heart of that Biosphere Reserve and will amphibians, at least five of the new species I reveal the increasing devastation having discovered have apparently gone extinct. If taken place, especially since about 2010, in coworkers and I had not been allowed to do that reserve, with that devastation currently that fieldwork, those species would have close to its center. disappeared from the planet without any In 2010, I began working with colleagues human knowledge that they formerly exist¬ gathering molecular samples of reptiles, ed. I fear there are other undiscovered which has already helped discover six species awaiting discovery that will now previously undescribed cryptic reptile spe¬ suffer that fate. That extinction rate can only cies. I also had ideas of where to target be expected to increase with the current several other taxa that certainly represent uncontested and illegal forest destruction complexes of undescribed cryptic species. I taking place in Honduras. made a list of localities where I was sure undescribed species occurred among sever¬ James R. McCranie al species complexes. I also had field trips Miami, Florida, USA planned to collect tissues from specimens 8 October 2015, updated 20 December lor molecular study of various populations I 2016 THE LIZARDS, CROCODILES, AND TURTLES OF HONDURAS. SYSTEMATICS, DISTRIBUTION, AND CONSERVATION JAMES R. McCRANIE1'2 CONTENTS Preface. v Superfamily Scincoidea.360 Abstract. 2 Family Scincidae.360 Introduction. 2 Infraorder Teiformata.370 Materials and Methods. 3 Superfamily Gymnophthalmoidea.371 An Update on the History of Reptilian Study of Family Gymnophthalmidae.371 Species Occurring in Honduras. 6 Family Teiidae.378 Class Reptilia. 9 Order Crocodylia.422 Species Accounts. 9 Superfamily Alligatoroidea and Family Order Squamata. 9 Alligatoridae.424 Infraorder Cordylomorpha. 18 Superfamily Crocodyloidea and Family Superfamily Xantusioidea. 19 Crocodylidae.430 Family Xantusiidae. 20 Order Testudinata.437 Infraorder Gekkomorpha. 29 Infraorder Cryptodira.438 Superfamily Chelonioidea.442 Superfamily Eublepharoidea. 29 Family Cheloniidae.444 Family Eublepharidae. 29 Family Dermochelyidae.465 Superfamily Gekkonoidea. 34 Superfamily Chelydroidea.469 Family Gekkonidae. 35 Family Chelydridae.469 Family Phyllodactylidae. 59 Superfamily Kinosternoidea.479 Family Sphaerodactylidae. 78 Family Kinosternidae.481 Infraorder Neoanguimorpha. 134 Superfamily Testudinoidea.501 Superfamily Anguioidea. 134 Family Emydidae.503 Family Anguidae. 135 Family Geoemydidae.528 Family Diploglossidae. 149 Species of Probable Occurrence in Honduras ... 549 Infraorder Neoiguania. 162 Distribution of the Lizards, Crocodiles, and Superfamily Iguanoidea. 162 Turtles in Honduras.551 Family Coiytophanidae. 168 Distribution Within Departments.551 Family Dactyloidae.218 Distribution Within Ecological (Forest) Family Iguanidae.225 Formations and by Elevation.557 Family Leiocephalidae.288 Distribution Within Physiographic Regions.... 563 Family Phrynosomatidae.291 Distribution Within Ecophysiographic Areas .. 564 Family Polychrotidae.331 Broad Patterns of Geographic Distribution.... 565 Infraorder Scincomorpha.334 Honduras as a Distributional Endpoint.574 Superfamily Lygosomoidea.335 Conservation Status of the Lizards, Crocodiles, Family Mabuyidae.338 and Turtles in Honduras.575 Family Sphenomorphidae.348 Vulnerability Gauges.575 CITES Categories and Species Occurring in Honduras.580 1 Smithsonian Research Associate, 10770 SW IUCN Red List Categories.581 164th Street, Miami, Florida 33157 (jmccrani@ Species Occurring in Protected Areas.584 bellsouth.net). Acknowledgments.593 Dedication: To the memory of my suegro Literature Cited.596 Emiliano Green (Warunta to his many friends), Index to Scientific Names.647 who was not only my father-in-law, but also a valued Index to Authors.656 Iriend. I miss you, man, especially remembering those long boat rides and walks we made together. Bull. Mus. Comp. Zook, SPS(2): 1-666, 2018 1 2 Bulletin of the Museum of Comparative Zoology, Special Publication Series, No. 2 Abstract. I discuss the taxonomy, distribution, and work in broadleaf humid forests away from conservation of the lizards, crocodiles, and turtles roads, whether at moderate or intermediate occurring in the Central American country of Hon¬ elevations, the author quickly began to duras. This is the fourth and final work in this series discover species of amphibians and reptiles documenting the amphibians and reptiles of that country. I propose three new species: one of new to science. Also, that fieldwork at those Laemanctus and two of the Sceloporus malachiticus elevations was expanded to regions in species complex of the S. formosus group. I also Honduras close to the international border suggest several other taxonomic changes, including elevating Laemanctus waltersi, Leiocephalus varins, with other countries, as well as into the vast Sceloporus schmidti, and Kinostemon albogulare to (at that time) lowland rainforests of the valid species. I also place Ctenosaura praeocularis in Mosquitia in northeastern Honduras. That the synonymy of C. quinquecarinata and Laemanctus extended fieldwork also resulted in the serratus mccoyi in the synonymy of L. serratus. All of those decisions are documented with specimens documentation of various amphibian and examined (with the exception of L. s. mccoyi) and reptilian species previously unknown from data and were reviewed by experts on those groups. Honduras. Several geographically wide-ranging species appear to An idea of the scope of that additional represent more than one species (i.e., Phyllodactylus tuberculosus, Sphaerodactylus continental^, Basiliscus fieldwork can be gained by the following vittatus). Also discussed is the uncontested, illegal discussion. Meyer and Wilson (1973) listed forest destruction occurring throughout the country, 2,351 specimens of lizards and close to 100 including in areas designated as national parks and more of crocodiles and nonmarine turtles in other types of reserves. The reader is urged to refer to the materials and methods section for an explanation museum collections around the world from of the procedures used in this work. In total, 107 Honduras. However, those authors made no species of lizards, 2 species of crocodiles, and 17 effort to examine those specimens they species of turtles are discussed. Full species accounts of 86 nominal forms are included herein, since a listed. A serious and misleading example of recent review of the anoles had included full species not examining those specimens is that accounts for 39 of those species (one anole species Meyer and Wilson (1973) listed the holo- recently added not included in a species account). type of Sceloporus schmidti, but that specimen was already lost about 1 year after its description in 1927. Currently, close to INTRODUCTION 14,000 Honduran specimens of those three Meyer and Wilson (1973) provided an groups reside in museum collections, almost annotated checklist of the lizards, croco¬ all of which I have examined. Those diles, and nonmarine turtles of Honduras. additional specimens, including marine Those authors included a total of 59 known turtles and specimens from the Swan species for the country. That total repre¬ Islands and Cayos Miskitos, not treated by sented 48 lizard species, 2 crocodile species, Meyer and Wilson (1973) have resulted in and 9 turtle species. The Meyer and Wilson the knowledge that 126 species of those (1973) effort represented “a revised version three groups (107 lizard, 2 crocodile, and 17 of a doctoral thesis” completed by the first turtle species) currently are documented to author in 1969. Thus, the majority of the occur in Honduras (with several additional work for that thesis was finished some 50 lizard species descriptions in press or in years ago. Needless to say, much progress preparation), as opposed to the 59 species has occurred in those 50 years regarding our recognized by Meyer and Wilson (1973). understanding of the systematics and tax¬ Thus, those 50 years of additional study of onomy of those reptile groups. Those 50 these three reptile groups have been years have also seen significant fieldwork productive for increasing our knowledge taking place in Honduras, especially at about them, but the rate of forest devasta¬ moderate and intermediate elevations, in¬ tion over those same 50 years have crucially cluding in isolated cloud forest areas. In that destroyed any hope that those types of