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Integrative Taxonomy Reveals the Chortís Block of Central America As an Underestimated Hotspot of Amphibian Diversity

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Integrative Taxonomy Reveals the Chortís Block of Central America As an Underestimated Hotspot of Amphibian Diversity INTEGRATIVE TAXONOMY REVEALS THE CHORTÍS BLOCK OF CENTRAL AMERICA AS AN UNDERESTIMATED HOTSPOT OF AMPHIBIAN DIVERSITY By JOSIAH HAROLD TOWNSEND 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 2011 1 © 2011 Josiah Harold Townsend 2 Para Ileana And in loving memory of my grandfather, Vernon Lynn Boyd 3 ACKNOWLEDGMENTS I want to begin by expressing my gratitude for the support shown by my advisory committee during the course of preparing this dissertation. My Chair, James Austin, supported me both academically and personally throughout this process; his leadership and steadfast support was instrumental in my success. I want to thank my committee members, Rob Fletcher, Mike Miyamoto, Rick Stepp, and Larry David Wilson, for their guidance, advice, and support through the process of researching and writing this dissertation. Writing a dissertation must be both a singularly selfish act and one that relies on the extended patience and support of loved ones, labmates, and colleagues. This certainly was the case for me. To the one person who is literally all three of those things, my wife Ileana Luque-Montes, I cannot adequately express the amazement with which I have watched you begin your own graduate studies over the past four months; maintaining a rigorous field schedule and supervising undergraduates, while I was of minimal help and fully focused on writing the following document. I simply would not have completed this without your remarkable dedication and support, and my gratitude will be best expressed by giving you the same degree of support as you pursue your own goals. I am blessed with the devoted support and seemingly endless patience and understanding of a wonderful and inspirational family, my parents Steve and Terri Boyd Townsend, my sister Katielynn, and my grandparents Vernon Lynn and Norma Jean Boyd and Willa Parker Townsend, who are largely responsible for any past and future successes I may have. In many ways, I am who I am today thanks to the mentorship of Larry David Wilson. As a mentor, colleague, and friend, Larry introduced me to Honduras in 1999 4 and guided my developing research interests there throughout my community college and undergraduate years, becoming my colleague and supporter as I progressed through my graduate studies. I owe no small debt of gratitude to my friends and colleagues in the Austin Lab, and, in particular, for actively assisting with my labwork and providing countless impromptu instructional sessions, I want to thank Jason Butler, John Hargrove, Nathan Johnson, Emily Saarinen, Matt Shirley, and Aria (Johnson) St. Louis. I also benefited from the dedicated work of a series of undergraduate researchers and volunteers: Teresa Burlingame, Dania Gutierrez, Jaclyn Irwin, Rachel Shapiro, Vicki Villanova, and Lauryn Walter. My work for the past six years has benefited greatly from the support of the Section of Protected Areas and Wildlife, Instituto Nacional de Conservación y Desarrollo Forestal, Áreas Protegidas y Vida Silvestre, and particularly Iris Acosta, Carla Cárcamo, Saíd Lainez O., Andrés Alegria, Ramón Alvarez L., Wilson Zúniga D., Sonia Martínez Moreno, and Wendy Aronne (Instituto Nacional de Conservación y Desarrollo Forestal, Áreas Protegidas y Vida Silvestre [ICF]). Fieldwork was carried out under a series of research permits issued by ICF, most recently Resolución DE-MP-086-2010 and Dictamen DVS-ICF-045-2010. In Honduras, my work would not have been possible without the hard work and support variously given by Ileana Luque-Montes (UNAH), Melissa Medina-Flores (UNAH, UNA), Luis A. Herrera (UNAH), Allan J. Fuentes and Eduardo J. Zavala (PROLANSATE), Alcalde Adolfo Pagoada-Saybe (Municipalidad de Arizona), Alcadesa Teresa Espinosa Aguilar (Municipalidad de Marale), Oliver Komar, Jose Mora, Jorge Iván Restrepo, and Fredy Membreño (Centro Zamorano de 5 Biodiversidad), Efrain Aguilar (San José de Texíguat), Alfonso Contreras (Mezapita), Alionso Portillo (Jilamito Nuevo), J. Dubón (La Liberación), Mario Orellana Leiva (El Playon), Leonel Erazo Chávez (El Cedral), Rafael Ulloa (Municipalidad de Gualaco), Eduardo Rico (ICF-Gualaco), Carlos Perdomo (Aldea Global), Paul House (UNAH, Herbario TEFH), Robert Dale (Los Naranjos), and Alicia Ward (Santa Bárbara). I would like to acknowledge the following people for their work in the field in support of this project: Carlos Andino, Ben K. Atkinson, James D. Austin, Christopher Begley, Mark Bonta, Jason M. Butler, Brian Campesano, César A. Cerrato, Gabriela Diaz, Anne Donnelly, Matthew Donnelly, Yensi Flores, Sergio C. Gonzalez, Levi Gray, Vladlen Henriquez, Luis A. Herrera, Paul House, Robert Hyman, Lorraine Ketzler, Ileana Luque, Christina Martin, David Medina, Melissa Medina-Flores, Mayron Mckewy-Mejía, Aaron Mendoza, Wendy Naira, Ciro Navarro, Lenin Obando, Sandy Pereira, Onán A. Reyes, John Slapcinsky, Mario Solís, Fito Steiner, Nathaniel Stewart, Alexander Stubbs, Katielynn Townsend, Steve Townsend, Scott L. Travers, Rony Valle, Hermes Vega, Alicia Ward, and Christopher Wolf. I am very grateful for the support of Amy Driscoll, Dan Mulcahy, Andrea Ormos, and the SI Barcoding Project (Smithsonian Institution Laboratory of Analytical Biology), and Roy McDiarmid (USNM) for supporting my ―Barcoding the Herpetofauna of Eastern Nuclear Central America‖ initiative. Deposition of voucher specimens and timely acquisition of catalog numbers was facilitated by: Jim McGuire, Ted Papenfuss, Sean Rovito, Carol Spencer, and David Wake (MVZ), Jose Rosado (MCZ), and Steve Gotte, Jeremy Jacobs, John Poindexter, and Robert Wilson (USNM). Jason Butler, Ileana 6 Luque, Javier Sunyer, and Scott Travers kindly contributed photographs for use in this dissertation. Various portions of this dissertation was funded by Critical Ecosystem Partnership Fund (CEPF), a Summer Research Grant from the Working Forests in the Tropics IGERT Program (National Science Foundation DGE–0221599) at the University of Florida, a grant to Kirsten E. Nicholson (Central Michigan University; National Science Foundation DEB-0949359), and the Explorer’s Club (via Robert Hyman; Flag #93). During my dissertation studies, I was supported by a 2007–09 NSF GK-12 Fellowship from the UF SPICE Program, and as much as any experience during my graduate education, my participation in this program was truly formative and help to shape the conceptual framework within which I propose to carry out research. I am especially grateful to Doug Levey and Suzan Smith, my partner-teacher Nate Stewart, and fellow fellows Jackson Frechette, Rachel Naumann, and Tom Tidyman, for helping to make SPICE a truly great experience. Following SPICE, my work with the UFTeach Program as an instructor for their core Research Methods class was key during the final two years of my dissertation, and I thank Alan Dorsey, Dimple Flesner, Griff Jones, Linda Jones, Katrina Short, Gloria Weber, and our RM students for making these last two spring semesters so enjoyable. Finally, I want to take this opportunity to dedicate my work in the past and future to the memory of three friends that I lost while completing this degree. I met Thad Owens as a student in my Herpetology class at UF and we quickly became friends, as he did with just about everyone who knew him. Thad was one of the most honest, unreserved, and enthusiastic people I will ever have the pleasure of knowing, and rarely a day 7 passes since he was lost in May 2009 that I do not think about him or find inspiration in his memory. Wade Wassenberg was a good friend since high school, and was one of the most loyal friends anyone could hope to have. After losing track of him for years, I learned that Wade had joined the US Army and served in the elite 1st Battalion of the 75th Ranger Regiment, surviving four tours of duty in Afghanistan and two more in Iraq from 2002 to 2008. After retiring to rejoin life with his beautiful young family, he was suddenly diagnosed with brain cancer and passed soon thereafter at the VA Hospital in Gainesville, walking distance from my office. I will always regret that while my friend suffered and passed literally minutes away from my home, I was unaware and in the field in Honduras. Finally, Don Mario Guiffaro of Olancho, Honduras, was an influential and remarkable man whose legendary life story included being a feared pistolero, a frontier gold miner, a jaguar hunter, and in his later years, and outstanding advocate and conservationist in the Patuca region. In 2007, Don Mario’s forceful opposition to illegal logging and drug trafficking in the Mosquitia was ultimately answered by violence, and he was gunned down in front of his son and friends. The memories of, and examples set, by these three friends, to each of whom I would not have hesitated to entrust with my life, and in some cases did just that, will always be my compass and my motivation as I continue through life’s journey. 8 TABLE OF CONTENTS page ACKNOWLEDGMENTS .................................................................................................. 4 LIST OF TABLES .......................................................................................................... 14 LIST OF FIGURES ........................................................................................................ 15 ABSTRACT ................................................................................................................... 18 CHAPTER 1 INTRODUCTION
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