Feeding ecology of black and white colobus monkeys from south coastal Kenya: the influence of spatial availability, nutritional composition, and mechanical properties of food items DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Noah Thomas Dunham Graduate Program in Anthropology The Ohio State University 2017 Dissertation Committee: W. Scott McGraw, Advisor Debbie Guatelli-Steinberg Dawn Kitchen Barbara Piperata Copyrighted by Noah Thomas Dunham 2017 ABSTRACT Identifying the degree to which primates alter their behavior and diets to different ecological conditions has significant implications for examining functional morphology, modeling socioecology and feeding competition, and developing primate conservation strategies. This study seeks to determine if Angola black and white colobus monkeys (Colobus angolensis palliatus) employ consistent dietary section criteria by investigating the behavior and diet of three groups inhabiting ecologically distinct areas of Kenya’s Diani Forest. The primary goals of this research are to examine feeding ecology, dietary flexibility, and food selection in relation to 1) seasonal and spatial availability, 2) mechanical toughness, and 3) nutritional composition of food items. Behavioral data were collected on three habituated groups (Ujamaa, Ufalme, and Nyumbani) over 267 days from July 2014 – December 2015. Behavioral data were recorded using a combination of instantaneous scan sampling and focal follows. Food availability was estimated by combining tree species composition profiles of home ranges with phenology data. Mechanical toughness was recorded with a portable test instrument. Nutritional composition of food items was calculated using a combination of traditional wet chemistry assays and near-infrared reflectance spectroscopy (NIRS) predictive models. Home ranges of the three groups differed significantly with regard to tree species composition and food availability. Diets differed considerably with regard to plant ii species and species-specific plant parts: only three species ranked in the top 20 food species for all three groups and mean monthly dietary overlap was just 10.4% among all groups. Dietary idiosyncrasies were not readily explained by differences in spatial and temporal availability of the most abundant tree species within the groups’ home ranges (i.e., all groups selected rare tree species and plant parts from their ranges). Leaf toughness was not a strong predictor of food selection; however, toughness significantly impacted foraging efficiency. Leaf toughness negatively correlated with ingestion rate (i.e., g/min) and positively correlated with masticatory investment (i.e., chews/g). NIRS models of nutritional components had strong predictive power despite the highly heterogeneous sample set. Conventional fiber limitation and protein to fiber ratio maximization models explained leaf selection in two of the three groups and one of the three groups, respectively. Despite significant differences in consumption of species- specific plant parts and quantity of kilocalories consumed per day, individuals of different groups balanced their intake of non-protein energy (NPE) and available protein energy (AP) to a consistent ratio of approximately 2:1. This study emphasizes that aspects of behavior and diet can vary considerably among groups living in different areas within the same forest. While availability, mechanical toughness, and nutritional composition of plant parts influenced food selection to varying degrees, maintaining a consistent NPE to AP intake (i.e., intake target) was the only consistent pattern among all three groups. Intake targets can be achieved by consistently consuming foods whose nutritional composition is close to or equal to that of the target or by consuming foods with disparate, yet complementary iii nutritional compositions. Unlike traditional models of food selection (e.g. protein maximization), the Geometric Framework provides a theoretical approach that can be universally applied to all investigations of primate feeding ecology. iv Acknowledgments This dissertation could not have been completed without the support and encouragement of many people. I would first like to thank my advisor, Scott McGraw. You have been higly influential in every aspect of my graduate studies and dissertation research. I am particularly grateful for your support during the ups and downs of fieldwork, for challenging me to be a better scholar and writer, and for your infectioius enthusiasm for primatology and scientific research. I would also like to thank my other dissertation committee members, Debbie Guatelli-Steinberg, Dawn Kitchen, and Barbara Piperata. I have greatly enjoyed our discussions over the years. The diversity of your theoretical approaches and research specaialities has strengthened my research and made me consider my work in relation to other subfields of biological anthropology. Finally, thanks to Clark Larsen and Dave Daegling for their guidance and support during my graduate career, particularly as I prepare for the future. I would also like to thank many others that have been highly influential and supportive throughout the course of my research prior to the dissertation. I am highly appreciative of my undergraduate faculty at Illinois Wesleyan University, Rebecca Gearhart Mafazy, Chuck Springwood, and Martin Nickels, for introducing me to the field of anthropology and for motivating me to pursue graudate studies. Thanks to Matthew v Preston who helped me through my initial fieldwork days as an undergraduate student in Tanzania, and for helpful discussions thereafter. My dissertation research was possible thanks to fincancial support from American Society of Mammalogists, Columbus Zoo and Aquarium, The Graduate School at The Ohio State University, Nacey Maggioncalda Foundation, National Geographic Society, National Science Foundation, The Ohio State University Department of Anthropology, Phi Kappa Phi Academic Honor Society, Pittsburgh Zoo and PPG Aquarium, and Priamte Society of Great Britain. Thanks to the Kenya Wildlife Society and the National Commission for Science, Technology, and Innovation for permission to conduct the research. During my time in the field, I am greatly appreciative of the support provided by those at Colobus Conervation. Special thanks to Pam Cunneyworth, Andera Donaldson, Keith Thompson, Kelly-Marie Martin, and Luciana Parazzi for making my time in Kenya both productive and enjoyable. I am grateful to have made many friends during my stay. Thanks especially to Joy Shirrazi for all of the memorable moments and to Henry Njoroge Kibue for being an excellent friend, roommate, and confidant. I send enormous thanks to my research assistants, John Ndege and Paul Opere. To John, your expertise in botany and plant identification, as well as your passion for scientific research, greatly enhanced the quality of the dissertation. To Paul, your energetic and optimistic disposition, relentless determination, and unparalleld kindness made every day in the field delightful. I am happy and humbled to consider you one of my closet friends. I also thank Saidi Mtali, Ferdinand Bele, Hassan Kidapa, and vi especially Bakari Mnyenze for their expertise and assistance in data collection over the years. I am greatly indebted to my friend and colleague, Alexander Lambert, who designed and assembled the mechanical toughness tester used for my dissertation research. What started as a casual conversation over beers at Little Bar slowly turned into a functioning prototype, and eventually a field-ready instrument. Alex, I cannot thank you enough for your dedication to making this project a success. Special thanks go to Luis Rodriguez-Saona for allowing me to conduct nutritional analyses in his laboratory, for several helpful methodological discussions, and for teaching me how to operate different infrared spectrometers. Your knowledge and expertise, particularly with regard to chemometrics and infrared predictive equations, elevated the quality of my dissertation research immensely. I would also like to thank Emilee Landers, Marcal Plans-Pujolras, Huseyin Ayvaz, and especially Mei-Ling Shotts and Peren Aykas for their assistance and patience in the lab. Thanks to Kevin Jewell and STAR Lab for a portion of the nutritional analyses. Thanks to Jessica Rothman for helpful discussions about nutritional analyses and nutritional balancing theory and methods. Thank you to my fellow graduate students and friends that helped shape my time at Ohio State. I am especially apprectiave of my cohort members that helped me enjoy and endure the intital years of coursework. Thanks to Dara Adams, Ashley Edes, Tim Sefczek, Jessica Walz, and Alex Wilkins for helpful discussions about primatology. vii Special thanks go to Erin Kane for being an excellent friend and for helping me through all aspects of my graduate career. I would like to thank my parents and siblings for supporting me all of these years and for inspiring me to pursue my passions. Thank you to Jacob Dunham for assistance and advice on statistical analyses. Lastly, I want to thank my wife, Veronica, for all of your loving support. I thank you for visiting me in Kenya, remaining patient as I spent months abroad, and for always beliveing in me. viii Vita 2007 ............................................................ Pontiac Township High School 2011 ...........................................................