Ecology and Behavior of the Bioko Island Drill (Mandrillus leucophaeus poensis) A Thesis Submitted to the Faculty of Drexel University by Jacob Robert Owens in partial fulfillment of the requirements for the degree of Doctor of Philosophy December 2013 i © Copyright 2013 Jacob Robert Owens. All Rights Reserved ii Dedications To my wife, Jen. iii Acknowledgments The research presented herein was made possible by the financial support provided by Primate Conservation Inc., ExxonMobil Foundation, Mobil Equatorial Guinea, Inc., Margo Marsh Biodiversity Fund, and the Los Angeles Zoo. I would also like to express my gratitude to Dr. Teck-Kah Lim and the Drexel University Office of Graduate Studies for the Dissertation Fellowship and the invaluable time it provided me during the writing process. I thank the Government of Equatorial Guinea, the Ministry of Fisheries and the Environment, Ministry of Information, Press, and Radio, and the Ministry of Culture and Tourism for the opportunity to work and live in one of the most beautiful and unique places in the world. I am grateful to the faculty and staff of the National University of Equatorial Guinea who helped me navigate the geographic and bureaucratic landscape of Bioko Island. I would especially like to thank Jose Manuel Esara Echube, Claudio Posa Bohome, Maximilliano Fero Meñe, Eusebio Ondo Nguema, and Mariano Obama Bibang. The journey to my Ph.D. has been considerably more taxing than I expected, and I would not have been able to complete it without the assistance of an expansive list of people. I would like to thank all of you who have helped me through this process, many of whom I lack the space to do so specifically here. However, I would like to express my gratitude to: My advisor, Gail Hearn, for convincing me to switch to primates, and providing me with what has been the most exciting and rewarding venture of my life. Words cannot express how grateful I am for the opportunity you provided me to study one of the most interesting and least known primates, in one of the most fascinating locations. Thank you for letting me find my own way, for giving me the independence to make mistakes, and providing me the support to learn from them. I am so much more capable as a result. Shaya Honarvar, who has filled a large number of roles over these years, each as important to me as the next. As a committee member, she worked tirelessly to help develop and refine this project, and edit countless reports, proposals, presentations, posters, and chapters. In the field she provided the logistical support, data, volunteers, and samples from Playa Moaba. As a friend, she gave me the support, encouragement, iv and reality checks I needed to continue. If it was not for our walks, I would not be writing this acknowledgement today. Michael O’Connor, for his uncanny ability to make me continually question what I think I know, and for making me strive to answer that question. His guidance has helped to build my self confidence in statistical analysis and hypothesis testing, and ultimately in my ability to succeed in this field. Susan Kilham, for her positivity, and for always helping me to see the bigger picture. Your insight has helped me to appreciate the interrelatedness of the world, and think more broadly about the theory and impact of ecology and evolution. Katy Gonder, for her consistently upbeat personality, excitement, and interest. Her knowledge of the ecology and evolution of primates helped me to shape this study into a coherent, manageable, and worthwhile endeavor. Christos Astaras, for the camaraderie, and for sharing with me your knowledge of drills, data and photographs, and methodological tips and tricks. Richard Bergl, Nelson Ting, and Josh Linder, for their technical, theoretical, and field assistance, and for introducing Drew and me to the primatology community with only a moderate amount of hazing. All those involved with the BBPP and Drexel Study Abroad, including Sally Vickland, Elizabeth Congdon, Heidi Ruffler, and Daniela Ascarelli. David Montgomery, Mark Andrews, Andrew Fertig, Karim Shnan, Matt Cuzzocreo, Livy Lewis, Nabil Nasseri for the logistical support, friendship, and much needed good times while in Bioko. Mary and Pete Johnson, for always treating me like family and for the best sausage gravy and biscuits in the eastern hemisphere. All of the Equatoguinean field workers who continually impress me with their skill and knowledge in the field. I would especially like to thank Cirilo Riaco, Miguel Angel Silochi, Fermin Muatiché, and Florentino Motove for collecting fecal samples, and for laughing at my excitement about said fecal samples. Filemon Rioso Etingue, the single most capable person I have ever had the opportunity to work with in the field. Thank you for teaching me how to identify Bioko’s primates and how to work and live in tropical Africa, and also for the friendship. v Esteban Muatiche, for opening your home to me, your friendship, work ethic, and beautifully high falsetto. I will always cherish our daily breakfasts and nightcaps together. My field assistants Brent Barry, Krissy Copeland, Tess Dornfeld, Julianna Gehant, Elissa Gordon, and Colleen Weathers, and lab assistants Rumaan Malhotra, Gian Bonetti, Atika Mehmood, Mark Nessel, Tyler Short, and Christian Brown, who volunteered a portion of their lives to drill feces and made this dissertation possible. Justin Jay, for countless hours in the blind watching drills from a few meters away. Thank you for the conversations, laughs, support, and thank you for your effort to help conserve the drills. Stephen (Steve Dubbs) Woloszynek, for your help with stats, conversations about religion and politics, and the inane video forwards at 4 am. Faculty and staff of the Drexel University Bio/BEES departments, especially Susan Cole, Ken Lacovara, Dan Duran, and Walt Bien. My fellow travelers of the Drexel University graduate experience. I was initially drawn to Drexel University because of the camaraderie and honesty I saw between the graduate students in this department. Over these years, I have come to realize that I had stumbled into something unique and extremely special in the world of academia. The friendship and support we have shared has enriched not only my graduate work, but also my outside life. Thank you all so much. The Hearn Lab members, for everything. Drew, Pat, Deme, and Miki, you have made life and field work so much more enjoyable. Thank you for all of the help and for the amazing and often ridiculous memories, working with you guys has been unforgettable. My friends from CHS and Stockton who continually stick by me, support me, and never take me seriously. Daniel Duran, Ilene Bean Eberly, Margaret Lewis, Ekaterina Sedia, Linda Smith, Roger Wood, and the other faculty of Richard Stockton College who convinced me I wasn’t finished with a bachelor’s degree, and have continued to support me ever since. Coach Todd Curll, Matt Markey, Bill Pino, Matt Waterhouse, Kim Kryscnski, Jason Mackie, and the other vaulters at RSC for giving me sporadic breaks from my vi terrestrial existence. Lily, Tuba, Noodle, and Oscar, for their love, the excuse to take a break and go outside, and for helping me maintain my sanity during this process. I would like to thank my family for their enduring support and belief in me. There is no way I can express how truly grateful I am for all of your patience, love, reprieve, and food. The Jones and Edwards family, thank you for the continual support and for taking me in as family. Levi and Sarah, you are the best siblings I could have ever asked for, thank you for your patience, friendship, and the amazing nieces and nephews who have brought me so much joy. Mom and dad, you made this possible through your encouragement, love, support, and by always trusting me to make the right decisions in my life. Thank you for investing in my future, and for giving me the opportunity to see the world. Finally, my wife and best friend, Jen, you are my foundation. You are the most selfless person I know, and I am so incredibly lucky to have been with you all of these years. Thank you for always telling me to pursue my aspirations, and working to making sure I achieve them. I am eternally grateful. vii Table of Contents LIST OF TABLES .........................................................................................................x LIST OF FIGURES .......................................................................................................xii ABSTRACT ....................................................................................................................xiv CHAPTER 1: INTRODUCTION TO THE DISSERTATION .................................1 Introduction to the Drill ....................................................................................1 Physical Description and Phylogeny ..............................................................1 Range ...............................................................................................................4 Conservation Status .........................................................................................5 Previous Studies ..............................................................................................9 Dissertation Objectives ......................................................................................10 Description of the Study Sites ...........................................................................11 Primate Community .........................................................................................13