Effects of Habitat Fragmentation on Non-Volant Small Mammals of the Interior Atlantic Forest of Eastern Paraguay
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EFFECTS OF HABITAT FRAGMENTATION ON NON-VOLANT SMALL MAMMALS OF THE INTERIOR ATLANTIC FOREST OF EASTERN PARAGUAY by NOÉ ULISES DE LA SANCHA SAENZ, B.S. A DISSERTATION IN ZOOLOGY Submitted to the Graduate Faculty of Texas Tech University in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Approved Jorge Salazar-Bravo Chairperson of the Committee Richard E. Strauss Guillermo D’Elía Tina Delahunty Nancy McIntyre Ralph Ferguson, Dean of the Graduate School December, 2010 ©2010, Noé Ulises de la Sancha Saenz Texas Tech University, Noé U. de la Sancha, December 2010 ACKNOWLEDGMENTS I would like to thank first and foremost my family – my parents, Pedro de la Sancha and Cruz Saenz, and my siblings, Anais de la Sancha and Eduardo ―Lalo‖ de la Sancha – for all of their love and support throughout my life; this would not have been possible without all of them. I want to thank Shayna S. Harris for all of her love and limitless support. I also thank Lee and Tarie Harris for their continuous support and hospitality. I would like to thank all the members of my committee, including Dr. Jorge Salazar-Bravo, Dr. Richard Strauss, Dr. Nancy McIntyre, Dr. Guillermo D’Elía, and Dr. Tina Delahunty. Special thanks to Dr. Jorge Salazar-Bravo for chairing my committee; Dr. Richard Strauss for all of his guidance, advice, and unselfish assistance and support; Dr. Nancy McIntyre who was always supportive throughout the thick and thin; and Dr. Guillermo D’Elía for his constant support, guidance, and friendship. This work would not have been possible without the help of many people in Paraguay. First and foremost I would like to thank Pastor ―Paco‖ Pérez and María Luisa ―Malu‖ Ortiz for their friendship and unselfish support, had it not been for them, the field work for this work would not have been possible. I want to thank all the volunteers (many of them great friends) including Lourdes Valdez, Julio Torres, Carlos Molina, Mario Maldonado, Natha Mujica, Alberto Carosini, Angélica Troche, Víctor Martínez Vázquez, Mirta Ruiz, Mario Méndez, Gretta Schaerer Lezcano, Sebastián Galeano Luraschi, Rodrigo Santander, Alejandro Bonzi, Johana López, Marcela Giménez, José Martínez Villalba and Andrea Gabriaguez. I would like to sincerely thank the Netto-Sisa ii Texas Tech University, Noé U. de la Sancha, December 2010 family for opening the doors of their home and lives to me (especially Flavia, Claudia, and Chiqui). I need to thank everyone at the Fundación Moisés Bertoni, with special thanks to Freddy Ramírez, Sixto Fernández, Myriam Velasquez, Rene Palacios, Alcides Ramírez, Roberto Ariel Ramírez, Emiliano Ramón, Fernández, Guido Fernández, Osvaldo ―Don Tito‖ Fernández Pintos, and all the Guardaparques Mbaracayú; all the community members of Arrollo Bandera, especially Felipe Bejarogi; Guyra Paraguay, including Alberto Yanosky, Oscar Rodas, Jose Cartes, Guardaparques of Reserva San Rafael, and Leticia López; Victor Vera, Abilio Ramírez, and Derlis Ojeda at Reserva Morombí; Nelson Pérez and officials at Itaipu Binacional; and the Guardaparques at Reserva Limoy. And finally I would like to thank all of the local workers whom worked on the grid systems. Quiero agradezerle a todos los que ayudaron en este proyecto; estoy eternamente agradecido con el apoyo y la ayuda que me brindaron. I have to give a very special thanks to Dr. Bruce Patterson, Dr. Ron Pine, Dr. John Zak, Dr. Clyde Jones, Dr. Llewellyn D. Densmore, Dr. Richard Stevens, Dr. Celia Lopez-González, Dr. Steven Presley, and Dr. Kevin Mulligan for their help and support throughout these years. I would like to thank countless friends during my tenure at Texas Tech for all the good times and experiences including the entire ―LA LATIN MAFIA‖ et al., Hugo Mantilla, Carlitos Dick, Nicté Ordóñez, Juan Pablo Carrera, Bryan Reece, David Rodríguez, Faisal Anwarali, Molly McDonough, Adam Ferguson, John Harting, Cibele Caio, Liz Siles, Sergio Solari, Miriam Venegas, Chris Higgins, Chris Bloch, Matt Campbell, Becky Roper, Vicki Swier, Ken Griffith, Raquel Marchán, Cody Thompson, and Tyla Holsombach. I want to thank the ladies who really run the Biology Department, iii Texas Tech University, Noé U. de la Sancha, December 2010 and who have always gone out of their way to help me, including Pat Moore, Judy Easterwood, Carol Espinoza, Lanita Ladd, and Jane Fowler. I want to thank Miguel Licona and Jean Long for believing in me, Irene Blakely for helping me believe in myself and pushing me to keep going forward, and Dr. Richard ―Dick‖ Neumann for introducing me to Mammalogy. Financial support was partially provided by a Fulbright Fellowship (Institute of International Education - US Department of State), the Mary Rice Foundation (special thanks to Mrs. Mary Rice), The Marshall Field Collection Fund of Field Museum of Natural History (The Field Museum of Natural History), the American Philosophical Society through the Lewis and Clark Exploration Fund, a Latin American Award (American Society of Mammalogists), research assistantships from the Department of Biological Sciences, Texas Tech University (TTU), a Texas Tech Association of Biologists Mini-grant (TTUAB), the J. Knox Jones, Jr., Memorial Endowed Scholarship (TTU), the Michelle C. Knapp Memorial Scholarship (TTU), a AT&T McNair Fellowship, and a Hispanic Scholarship Fund Award. Thanks to Dale and Louie Hanson for their valuable donation to my project. I thank Robert J. Baker and Heath Gardner (Natural Sciences Research Laboratories, TTU) for all their help at the NSRL. I thank Dr. Nancy McIntyre and Dr. Richard A. Nisbett (Dept. of Kinesiology, Rice University) who helped with traps. I thank all the curators who opened the doors to their collections for their assistance in learning about Paraguayan fauna, including Dr. Bruce Patterson (the Field Museum Natural History), Dr. Phil Myers and Steve Hinshaw (University of Michigan Museum of Zoology), Drs. Al Gardner and Don Wilson (Smithsonian), Drs. Joseph Cook and Jon iv Texas Tech University, Noé U. de la Sancha, December 2010 Dunnum (Museum of Southwestern Biology), Dr. Jim Patton and Chris Conroy (Museum of Vertebrate Zoology), Dr. Kurt Schwenk (Connecticut State Museum of Natural History), Eilene Westwig and Dr. Robert Voss (American Museum of Natural History), and Isabel Gamarra de Fox (Museo Nacional de Historia Natural del Paraguay). Without a doubt, Paraguay will forever hold a piece of my being. Espero esto ayude un poco a avanzar nuestro conocimiento de la naturaleza de Paraguay. Rohayhu Paraguay!! v Texas Tech University, Noé U. de la Sancha, December 2010 TABLE OF CONTENTS ACKNOWLEDGMENTS ................................................................................................ iii LIST OF TABLES ............................................................................................................ ix LIST OF FIGURES .......................................................................................................... xi CHAPTER I. INTRODUCTION: PARAGUAY, ITS MAMMALS, AND ITS ATLANTIC FOREST ...................................................................................... 1 General Information about Paraguay ...................................................... 1 Deforestation in the Interior Atlantic Forest of Paraguay ....................... 2 Mammal research in Paraguay ................................................................ 3 Literature Cited ....................................................................................... 8 II. A PREDICTIVE MODEL OF NON-VOLANT SMALL MAMMAL ASSEMBLAGES IN EASTERN PARAGUAY ATLANTIC FOREST REMNANTS.................................................................................................. 26 Introduction ........................................................................................... 26 Materials and Methods .......................................................................... 30 Basic framework: Non-volant small mammals ......................... 30 Brazilian dataset ....................................................................... 31 Species identification ................................................................ 31 Trapping effort .......................................................................... 32 Species predictor models .......................................................... 32 Paraguayan remnants ............................................................... 32 Paraguayan data/models .......................................................... 32 Endemics and non-forest species .............................................. 34 Principal Component Analysis (PCA) ...................................... 35 Results ................................................................................................... 36 Brazilian models ....................................................................... 36 Paraguay remnants ................................................................... 37 Paraguayan models .................................................................. 37 Species-area relationships ........................................................ 38 Effects of trapping effort ........................................................... 38 Area reduction .......................................................................... 39 Forest, non-forest, and endemic species ................................... 40 Principal Component Analysis ................................................. 40 Discussion ............................................................................................