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Ecology, Habitat Preference, and Conservation of Neotropical Non-Volant Mammal Communities in Costa Rica’S Caribbean Lowlands

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Ecology, Habitat Preference, and Conservation of Neotropical Non-Volant Mammal Communities in Costa Rica’S Caribbean Lowlands Ecology, habitat preference, and conservation of Neotropical non-volant mammal communities in Costa Rica’s Caribbean lowlands By Andrea Romero Submitted to the graduate degree program in the Department of Ecology and Evolutionary Biology and the Graduate Faculty of the University of Kansas in partial fulfillment of the requirements for the degree of Doctor of Philosophy. ________________________________ Chairperson: Robert M. Timm ________________________________ Norman A. Slade ________________________________ Helen M. Alexander ________________________________ Mark E. Mort ________________________________ Xingong Li Date Defended: 19 April 2013 The Dissertation Committee for Andrea Romero certifies that this is the approved version of the following dissertation: Ecology, habitat preference, and conservation of Neotropical non-volant mammal communities in Costa Rica’s Caribbean lowlands ________________________________ Chairperson: Robert M. Timm Date approved: 19 April 2013 ii ABSTRACT Biodiversity in the tropics is continually being threatened by anthropogenic disturbances such as habitat degradation and fragmentation. Unfortunately, conservation decisions and management of wildlife are difficult tasks to undertake without adequate scientific knowledge. In the tropics, little is known about the conservation value of secondary forests for mammals, the manner in which fragmentation can affect low-density populations, how mammalian assemblages have changed over time, and even some of the most basic ecology of organisms. Herein, I use multi-species and single-species approaches at different spatial scales to address these topics using the non-volant mammals in the Caribbean lowland rainforests of Costa Rica. I first tested the null hypothesis that primary and secondary forests have equivalent assemblages of non-volant mammals. I found no significant differences in the community composition of non- volant mammals in these two forest types. This community-based study led me to address the population biology of the collared peccary, one of the most common species in these faunal surveys. Using both my data and historical records, I established that this species has been greatly affected by anthropogenic disturbances in the area, and increased after the extirpation of white-lipped peccaries. My work at both the community and population level emphasized that the rodent communities in the Caribbean lowlands are at low densities, as represented by trap success. Fragmentation of tropical forests may therefore have particular consequences for mammalian communities. I thus performed a landscape level study with 15 fragments in Costa Rica’s Caribbean lowlands. This work revealed that forest fragments showed differences in species diversity and relative abundance. Larger fragments (>9 km2) had higher relative abundance for all species. I also focused on the population biology of two species, the spiny pocket mouse, Heteromys desmarestianus and the arboreal vesper mouse, Nyctomys sumichrasti. My work on the former species included molecular genetic studies which revealed that the iii Caribbean lowlands have cryptic diversity that has not been previously explored. In overview, non-volant mammalian communities of the Caribbean lowlands represent a rich assemblage of organisms that are crucial to the health of tropical ecosystems. While anthropogenic disturbances are affecting these populations, a good understanding of the conservation value of secondary forests, how fragmentation can affect populations, and the ecology of organisms is crucial for making data-driven management and conservation decisions. iv ACKNOWLEDGMENTS This body of work is a result of the help, encouragement, and support provided by many people throughout my graduate career. First and foremost, I’d like to extend a most sincere thank you to my advisor, Bob Timm. Bob had the patience to teach me how to trap rodents where it was difficult to do so, and encouraged me to pursue my research interests, well before I even started graduate school. I am most appreciative of his support throughout my graduate career and his help in serving as a sounding board for my project ideas, writing countless letters of recommendation, and reading many drafts of this dissertation, and doing so always with a smile. I’ve learned so much about mammalogy, teaching, mentoring, and writing from Bob, and look forward to working with him in the future. The spark of my interest in mammalogy can be credited to Deedra McClearn, who taught me how to catch my first mouse in South Africa. Deedra recruited a naïve college student to spend many amazing months in South Africa, learn many field techniques, and simply put, think that mammals were just awesome. When I asked Deedra for advice on where to look for jobs so that I could get more field experience before graduate school, she offered me one. I was lucky to work with Deedra in Costa Rica, and have her as my mentor as I began to look at graduate schools and think about research ideas. It was a joy to see her each summer when I worked in Costa Rica, and get her constant support, often after long days in the field with 0 mouse captures. I have had the pleasure to embark in so many different adventures with her throughout the world, and look forward to many more in the future. I extend my gratitude to my committee members: Helen M. Alexander, Mark E. Mort, Norman A. Slade, and Xingong Li for their critical review of my work, ideas, interpretations, and statistics. I appreciate their honest feedback on my work, encouragement, and help with sometimes difficult datasets or project setbacks. Kenneth G. Gerow and Ford Ballantyne IV v provided advice on complex statistical matters for several projects. J. Andrew DeWoody, Anna Fahey, Nicholas Marra, Kendra Abts, and John Patton provided support for molecular studies at Purdue University, and graciously allowed me to use their facilities and taught me various molecular techniques. Mark Mort and Oscar Toro helped me deal with molecular data results and taught me how to use various phylogenetic programs. Javier Guevara Sequeira and SINAC– MINAET provided permits for my work in Costa Rica. Felix Corrales, Paul S. Foster, Alex Gilman, and Richard LaVal collected and saved various important specimens used in this dissertation. Bernal Rodríguez (UCR and MNCR), Sue McLaren (CMNH), Bruce Patterson (FMNH), Ronald Pine (KU), Burton Lim and Mark Engstrom (ROM), and Neal Woodman (USNM) provided access to data from collections under their care. Isaias Alvarado-Díaz, James H. Beach, Amos Bien, Deborah A. Clark, Paul S. Foster, David Janos, Daniel H. Janzen, Gary S. Hartshorn, Mirjam Knörnschild, Richard K. LaVal, F. Gary Stiles, Don E. Wilson, Joe Wunderle, and Bruce E. Young provided critical historical data and recollections of peccaries in Costa Rica. This research was funded by several sources, including the Madison and Lila Self Graduate Fellowship, OTS Research Grant, Tinker Grant, Idea Wild, Explorer’s Club, KU’s Travel Grants, Ida H. Hyde Grant, Endowment Grant, and the KU–UCR Grant. Terry McGlynn provided me with great opportunities to mentor students in the field, and funded some of my time at La Selva via the Tropical Ecology Mentorship Program NSF-OISE 0854259. I would like to thank the Organization for Tropical Studies (OTS) and the staff of La Selva Biological Station for supporting of my research. Isabelle Boittin, Marisol Luna, Ronald Vargas, Orlando Vargas, Ragde Sánchez, Karla Baquero, Sara Hu, Erick Berlin, and various private landowners helped with the logistics, provided access to land, and aided with data collection. A special thank you is owed to Guillermo Fandos Guzmán and Virginia Weigand- vi Noble for assistance in the field over various summers. Although trap success was always low and the field conditions harsh, they continued to work with me in the field over many summers, and always with a smile and contagious enthusiasm. Marcia Snyder, Malia Fincher, Steven Whitfield, Jenny Stynoski, Alex Gilman, and Felix Corrales provided hours of “playtime” and tomfoolery after the most difficult field days. Finally, a special thank you goes out to my family, who has supported all of my decisions (even the crazy ones), has always encouraged me to travel the world, and take an interest in the natural world, even when they were unsure about what exactly I did. My father, mother, and sister are my biggest cheerleaders and I am grateful for all of their support throughout the years. My husband, Brian O’Neill, always pushes me to work hard, but reminds me to have fun too. His patience and supportiveness knows no bounds. This body of work is a tribute to all of these people who made it possible. vii TABLE OF CONTENTS Title page…………………………………………………………………………………………..i Acceptance Page………………………………………………………………………..…………ii Abstract………………………………………………………………...…………………………iii Acknowledgments………………………………………………………………………..………..v Table of Contents…………………………………………………………………………….….viii List of Figures………………………………………………………………………….…………ix List of Tables…………………………………………………………………………………...…x Introduction………………………………………………………………………………………1 Chapter 1. Non-volant mammalian populations in primary and secondary Neotropical rainforests as revealed by transect surveys………………………………………………………………..…5 Chapter 2. Group dynamics, behavior, and current and historical abundance of peccaries in Costa Rica’s Caribbean lowlands………………………………………………………………..44 Chapter 3. Forest fragments and rodent abundance and diversity in Costa Rica’s Caribbean lowlands……………………………………………………………………………………….....89
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