Caiman Yacare) of SOUTH AMERICA
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PHYLOGEOGRAPHY AND CONSERVATION GENETICS OF THE YACARE CAIMAN (Caiman yacare) OF SOUTH AMERICA By ROBERT E. GODSHALK A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2006 Copyright 2006 by Robert Godshalk This document is dedicated to Dr. Federico Medem – a true renaissance scholar with a keen interest in knowledge, the supreme ‘caimanologo’, friend. He spearheaded detailed studies of Neotropical crocodilians, an inspired legacy that continues today in many countries. He encouraged me when I first began the study of these fascinating creatures. With his support, I undertook the first surveys in Venezuela of the endangered Orinoco crocodile, a cause dear to his heart. Our enthusiastic discussions on his patio at the Instituto Roberto Franco remain fond memories. His wonderful spirit and love of life and all living things was truly inspirational. ACKNOWLEDGMENTS First I want to thank the current and former members of my committee. Dr. F. Wayne King, committee chair, made it possible for me to attend the University of Florida. His broad knowledge of caimans and position in the world of crocodilian conservation assisted me in many ways. Franklin Percival and Ray Carthy posed thought provoking questions on wildlife ecology and conservation. Charlie Baer generously became a committee member at a late date and, with his phylogeographic experience, guided me through the complex genetic ananlysis. Clyde Kiker, my outside committee member, continues to stimulate my interest in natural resource economics. My gratitude goes to former members Richard Bodmer, currently in England, and John Eisenberg (deceased) for their time and interest in my project. The nature of fieldwork in a foreign country placed me in constant need of assistance. Warm thanks go to the many individuals that helped me during my fieldwork and sample collection. Inclusion of all is impossible; I apologize to anyone inadvertently overlooked. Gracias to Tito and Rosario Ortiz for hospitality in Trinidad, Bolivia. My appreciation goes to the many ranchers who spent time with me, including Eduardo Añez, Rodolfo Brükner, Ron Larsen Dr. Romulo Lozada, Osvaldo Monasterios, Andres Parra, Armando Pedacassé, Cristina Quaino de Paz, Rubens Rivero, Ernesto Salas and Hugo Serrate. The indigenous people were very open to new ideas and keen on developing their resources in a sustainable fashion. Valentin Muiba Guaji and Wigberto Rivero took iv particular interest in the CITES project. The various non-governmental organizations and individuals involved with Indian issues were also enthusiastic and helped at various times, thanks in part to Sylvia Chalukián, Cristian Gallegos, Jürgen Riester, Allyn Stearman, and Wendy Townsend. My warmest thanks go to so many government employees in various offices that extended their time and data after only a brief introduction: Ing. Freddy Arteaga, Teresa Centruión, Oscar Méndez, Ivan Pinaya, Luis Alberto Vaca, Lilian Villalba, and Jorge Montaño. Special thanks go to the Wildlife Department employees in Beni who assisted during my trips: Gerardo Coimbra, Enrique Cuellar, and Sergio Salazar. The various biologists I met, both Bolivian and expatriate, offered opinions, advice and help: James Aparicio, Juan Pablo Arce, Claudio Barra, Stephen Beck, Paolo Bertella (deceased), Werner Honnegger, René Ibañez, Tim Killeen, Carmen Miranda, Luis Pacheco, Damian Ruiz, Andrew Tabor, and Marcelo Zalles. Thanks go to Dr. Mario Baudoin for finding time in his busy schedule to talk with me. Arturo Moscoso gave vigorous support and financial assistance from the Netherlands embassy. And last, but certainly not least, I thank Dante Videz R., who was my assistant for nearly all of the fieldwork. He had also accompanied Dr. King on surveys in 1986-87, and proved invaluable during our projects. My sample pool was substantially enhanced with the help of Crocodile Specialist Group members: Alejandro Larriera sent yacare samples from Argentina, Alvaro Velasco in Venezuela, Perran Ross arranged use of the Honduras specimens, and Kent Vliet provided samples for both Paleosuchus species from the St. Augustine Alligator Farm in Florida. v My entrance into the new labyrinth of molecular science was eased by the mentoring by Dan Brazeau, Ginger Clark, Bill Farmerie and Charlie Baer. Highest thanks and praise go to Ginger Clark, head of the ICBR Genetic Analysis Laboratory, who initially allowed me to have bench space, mentored me continuously, and later employed me. Special thanks go to the ICBR people, especially David Moraga and Sharon Norton of the Education Core for all the informative workshops on essential techniques. Kevin Holland and Ernesto Almira of the ICBR-GSSL assisted me with the bulk of the sequence data generation on the Amersham MegaBACE 1000. Savita Shankar of the ICBR Sequencing Core performed her sequencing magic on stubborn templates and for haplotype verification. Tara Paton of the Sick Children’s Hospital in Toronto generated most of the raw microsatellite data. Naomi Phillips gave me much needed assistance, especially with her thorough analysis, advanced computer skills and moral support. Ed Braun helped me get through the myriad of molecular clock models and David Reed assisted with the phylogeographic analysis. Financial support for this project was supplied initially by a Tropical Conservation and Development (TCD) research grant, Programs for Studies on Tropical Conservation (PSTC) research grant, and by my father, Ernest L. Godshalk. Funding for the extensive population surveys and sample collection came from the CITES central office in Switzerland. Additional funding towards laboratory processing costs were supplied by my father and my wife, Vicki McGrath. Without Vicki’s organizational support this project might still be “in progress.” Tireless effort was spent by many individuals in editing drafts. I specifically want to thank F. Wayne King, Charlie Baer, Franklin Percival, Ray Carthy, Clyde Kiker, Paul vi Moler, John Thorbjarnarson, Kent Vliet and Ginger Clark for comments on the manuscript. A special note of thanks goes to my parents, Ernest and Trudy, for their constant moral support and encouragement during my unconventional life. My warmest gratitude and love go to my wonderful wife, Vicki McGrath, who reviewed manuscripts, organized details for deadlines, endured endless piles of literature cluttering her life and was very instrumental in the final product. My gratitude goes to anyone else unnamed that contributed in some part to the final product. The journey was a long time in the making and I had contact with many people in several countries; thanks go to all. vii TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................................................................................. iv LIST OF TABLES............................................................................................................. xi LIST OF FIGURES ......................................................................................................... xiv ABSTRACT..................................................................................................................... xix CHAPTER 1 INTRODUCTION ........................................................................................................1 Background...................................................................................................................2 Evolving Conservation .................................................................................................4 Current Study................................................................................................................8 2 YACARE CAIMAN DISTRIBUTION – RIVER SYSTEMS, HABITATS AND DETAILS ON THE BOLIVIAN LOWLANDS ........................................................11 Introduction.................................................................................................................11 Amazon River Basin...................................................................................................15 Hydrology............................................................................................................15 Environment ........................................................................................................23 Temperature .................................................................................................23 Precipitation .................................................................................................24 Soils..............................................................................................................26 Biota ....................................................................................................................26 Flora .............................................................................................................26 Fauna ............................................................................................................28 Human Dimension...............................................................................................30 Paraguay-Paraná River Basin .....................................................................................32 Hydrology............................................................................................................32 Environment ........................................................................................................36