Socioecology, stress, and reproduction among female Diana monkeys (Cercopithecus diana) in Côte d’Ivoire’s Taï National Park DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Erin Elizabeth Kane Graduate Program in Anthropology The Ohio State University 2017 Dissertation Committee: Professor Debra Guatelli-Steinberg Professor Dawn Kitchen Professor W. Scott McGraw, Advisor Professor Barbara Piperata Copyrighted by Erin Elizabeth Kane 2017 Abstract Socioecological models have been used to explore the relationship between female sociality and feeding ecology for nearly 40 years. Models typically distinguish between species eating ubiquitously distributed resources (e.g., leaves) in which females do not exhibit strong social bonds, engage in minimal feeding competition, and minimal territorial defense. These are contrasted with species relying on patchily distributed resources (e.g., ripe fruit) in which females do exhibit social bonds, engage in feeding competition, and defend group territories. Although these models have been critiqued and extended, these basic predictions have been widely used to develop hypotheses about the relationship between ecology and sociality among primates. Much of this research has focused on open-habitat and/or terrestrial primates, while relationships between ecology and sociality remain unexamined in many forest-dwelling and/or arboreal taxa. This is detrimental to our ability to develop unifying models of primate sociality and ecology, and problematic considering the arboreal, forest-dwelling niche filled by the earliest primates and primates in their evolutionary environment. This dissertation examines the relationship between ecological variables and diet, social behavior, reproduction, and stress in arboreal, forest-dwelling guenons. Previous research has demonstrated that Diana monkeys (Cercopithecus diana) exemplify many predictions of socioecological; models: they are ripe fruit specialists, females are philopatric and socially bonded, and feeding competition both within and between groups ii is relatively high compared to closely related taxa. Here, I use socioecological data collected between July 2013-August 2015 on habituated adult female Diana monkeys living in the Taï Forest, Côte d’Ivoire, and fecal samples collected over the same period to test the hypothesis that fluctuations in food availability have significant consequences for Diana monkeys in terms of their diet, social relationships while foraging, the timing of reproduction, and metabolic stress. Diana monkeys preferentially fed at very large, relatively rare trees, supplementing this diet with invertebrates and fruits from less-preferred trees that tend to be smaller but ubiquitously distributed. Diana monkeys timed birth and lactation for periods with of relatively high fruit availability; however, there was no relationships between fluctuations in food availability, diets, and fecal glucocorticoid concentrations. While fecal glucocorticoid concentrations did fluctuate interannually, concentrations never reached pathological levels. Instead, I argue that these fluctuations allow Diana monkeys to cope with the normal strains of the life of wild primates. Diana monkeys are thus able to maintain their fruit-rich diet with minimal competition and few serious stressors. This has implications for socioecological models of primate behavior that derive primate sociality from resource distribution, with particular emphasis on competition over access to resources. Diana monkeys complicate these models because, although they eat high-value, discontinuously distributed resources, feeding competition does not drive their social behavior and there is little evidence that they experience negative consequences during periods of low food availability. This affirms the need to test not just the causes of primate behavioral ecology, but also its consequences. iii This dissertation is dedicated to my undergraduate advisors, Tab Rasmussen and Bob Sussman. You helped set me on this path. Thank you. iv Acknowledgments Thank you to my advisor, Scott McGraw, and all my committee members – Dawn Kitchen, Barbara Piperata, and Debbie Guatelli-Steinberg. You have all been patient, interested, helpful, and constructive, and I appreciate the time you have spent reading, rereading, writing, rewriting, and discussing more than I can say. Scott, thank you for sharing the monkeys of the Taï Forest with me, and supporting me throughout fieldwork and writing. Thank you also to Mark Hubbe, Mark Moritz, Kris Gremillion, and Julie Field for immeasurably enhancing my graduate education. Thank you to Dr. Larsen for department leadership, and to Wayne Miller, Jean Whipple, and Elizabeth Freeman for keeping the anthropology department functioning. Thank you to all funders of this research, including Sigma Xi, the OSU Office of International Affairs, the Graduate School, the Department of Anthropology, and the American Society of Primatologists. For permission to work in the Taï Forest, I thank the Ministere d’Enseignment Superireur et Recherche Scientifique, the Ministere d’Agriculture et Resources Animales, and the Centre Suisse de Recherches Scientifiques. Without the help of Dr. Anderson Bitty, conducting this research would have been much more of a challenge! I am hugely indebted to the Reproductive Endocrinology lab at the Smithsonian Conservation Biology Institute, notably Dr. Janine Brown, Nikki Haley, Dr. Natalia v Prado-Ovieda, Juthapathra Dechanupong, Morgan Maly, Deanna Moore, and Arianna Bond. Thank you for welcoming me, training me, and sharing resources, ideas, and fun in Front Royal! Thank you to my field friends in Taï for their camaraderie – Claudia Stephan, Fredy Quintero, Martina Magria, Sonia Touistiti, and Adam van Casteren made time in the field enriching, enjoyable, and manageable. Thank you to all the friends who have supported me in Columbus, especially my Artisan family (especially Sarah Dorn, Aisha Bradshaw, Jessica Kehn) and Triceratops. I am so grateful for the close social bonds I’ve formed with the other primatology graduate students at Ohio State. Dara Adams, Noah Dunham, Tim Sefczek, Alex Wilkins, Ashley Edes, Jessica Walz, Michelle Rodrigues, and Cathy Cooke – thank you! I am also grateful for the warm and collegial environment of the anthropology graduate program, with particular gratitude to Lisa Beiswenger, Barbara Betz, Erin Blankenship-Sefczek, Abby Buffington, Addy Cary, Ana Casado, Melissa Clark, Aaron Comstock, Jesse Goliath, Sarah Holt, Daryin Hummel, Katy Marklein, Leigh Oldershaw, Brian Padgett, Genevieve Ritchie-Ewing, Marissa Stewart, Gordon Ulmer, and Andrew Weiland. Thank you to my family – grandparents, great grandparents, aunts, uncles, cousins – who love and support me in my wild dream to go chase monkeys, despite the many missed family occasions. Thank goodness for my little brother, Brendan, who has always been there to make me feel short, capable, and loved. My parents, Jessica and Derek, have not once balked at what I do with my life (well, OK. Maybe the second and third times I called to say I had malaria, they may have wished I made slightly different choices). Their unconditional support and love have kept me going. vi Finally, this wouldn’t have been possible without the skilled, wonderful employees of the Taï Monkey Project and their families. Nasuo, aséo, merci beaucoup à vous! Frederic Gnépa, Ferdinand Belé, Richard Pého, Gerard Gah, Bertin Dioré, and Paterson Kalo kept me alive, sane, and sciencing in Taï National Park. They welcomed me into their communities, villages, families with incredible kindness and generosity. I know that when I go back to Côte d’Ivoire and arrive in Ponan, Daobly, Tai, Gouliyako, or Paulioula, I’ve made it back home. vii Vita June 2006 .......................................................Central High School 2009................................................................B.A. Anthropology and Environmental Studies, Washington University in St. Louis 2012................................................................M.A. Anthropology, The Ohio State University 2010 to present ..............................................Distinguished University Fellow, Department of Anthropology, The Ohio State University Publications Dunham NT, Kane EE, McGraw WS. 2017 Humeral correlates of forelimb elevation in four West African cercopithecid monkeys. American Journal of Physical Anthropology 162: 337-349. Dunham NT, Kane EE, Rodriguez-Saona L. 2016. Quantifying soluble carbohydrates in tropical leaves using a portable mid-infrared sensor: implications for primate feeding ecology. American Journal of Primatology. 78: 701-706. Kane EE, Gnepa F. 2016. An infanticide attempt after male takeover in Diana monkeys (Cercopithecus diana diana) in Taï, Côte d’Ivoire. African Primates. 11: 37-40. viii Watsa M, Erkenswick G, Halloran D, Kane E, Porier A, Klonoski K, Cassalet S, Maciag E, Mangalea M, Dinsmore M, McCready H, Boughan B, Parker C, Hickmott A, Nole Bazán I, Zuñig A.2015. A field protocol for the capture and release of callitrichids. Neotropical Primates 22: 59-68. Dunham NT, Kane EE, McGraw WS. 2015. Scapular morphology and forelimb use during foraging in four sympatric cercopithecids. Folia Primatologica 86: 474- 489. McGraw WS, van Casteren A, Kane EE, Geissler E, Burrows B, Daegling DJ. 2015. Ingestive and oral processing behaviors of two colobine monkeys in Tai Forest,