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EFFECTS OF FOOD NUTRITIONAL AND MECHANICAL PROPERTIES ON FORAGING OF JUVENILE IN WILD TANA RIVER MANGABEYS, CERCOCEBUS GALERITUS, KENYA by STANISLAUS KIVAI A dissertation submitted to the School of Graduate Studies Rutgers, The State University of New Jersey In partial fulfillment of the requirements For the degree of Doctor of Philosophy Graduate Program in Anthropology Written under the direction of Ryne A. Palombit And approved by _____________________________________ _____________________________________ _____________________________________ _____________________________________ New Brunswick, New Jersey January, 2018 ABSTRACT OF THE DISSERTATION “Effects of food mechanical and nutrition properties on foraging of juvenile in wild Tana River mangabeys, Cercocebus galeritus, Kenya” By STANISLAUS KIVAI Dissertation Director: Ryne A. Palombit Food is an essential for individual survival and reproduction, but intrinsically presents a variety of mechanical obstacles that must be met and nutritional requirements that must be overcome by the consumer. Mechanically, foods may be hard, tough, or stiff enough to inflict dental, damage on teeth, cranium, or facial form while chemically, it may be packed with toxins, nutrient deficiencies or in excess, which may harm the consumers' fitness. Therefore, feeding efficiency and challenges associated with foods provide a strong selective mechanism potentially influencing the evolution of foraging behavior and diet. The problems of accessing food and ensuring sufficient intake of energy and protein are likely to have a greater impact on juveniles than adults, as juveniles are smaller, less experienced and generally more inefficient foragers. Detailed studies testing the effects of these formative factors on juvenile feeding have yet to be conducted, however, and there is a poor understanding of how adult-juvenile foraging differs in nonhuman primates. The main debate on this issue centers on the development of juvenile ii foraging efficiency, particularly on the effects of body size, nutritional needs, and experience in driving the suggested adult-juvenile differences in feeding. In order to clarify understanding of adult-juvenile foraging differences, I collected data and compared the diet choice, metabolizable energy intake, the mechanical and nutritional properties of foods, and the interactive effects of these properties on food selectivity in the Tana River mangabeys, Cercocebus galeritus. This research addressed the following questions about the implications of mechanical and nutritional properties of foods on foraging. Compared to adult lactating females, how do juveniles differ in: (1) diet, particularly in food choice and energy intake?; (2) the influence of mechanical and nutritional properties on foraging decisions?; and, (3) food electivity as a result of interactive effects of the mechanical and nutritional properties of foods? I collected data on feeding behavior using focal animal sampling from two wild groups (Kitere & Mchelelo groups) in lower Tana River forest fragments, Kenya. I measured the fracture toughness and elastic modulus of the primary and fallbacks foods of the mangabeys using a portable FLS-1 food tester machine. Finally, I performed laboratory analyses to determine the nutritional properties of the foods. Overall, I found that dietary breath (BA) was wider in lactating females (BA = 0.13) than in juveniles (BA = 0.11) and was higher in Mchelelo group (0.22 & 0.17) than in the more anthropogenically impacted Kitere group (0.15 & 0.11) for both age classes, respectively. Juveniles’ mean metabolizable energy intake per metabolic body mass per minute was higher than in lactating females. Juveniles also ingested more available protein per metabolic body mass per minute than the lactating females. Lactating females ingested foods with iii significantly higher fracture toughness than juveniles, but elastic modulus values of the food did not differ between the two age classes. Moreover, fruit availability and the mechanical and nutritional properties of foods interacted to influence the diet selection across seasons of different fruit availability, but there were no age class differences. These findings supported my predictions that juveniles will ingest more protein and metabolizable energy per unit of metabolic body mass compared to lactating females potentially due to increased nutritional needs for growth. Additionally, differences in the mechanical properties of the foods ingested by juveniles and lactating females support the Need-To-Learn hypothesis that juveniles that require time to learn and acquire competence in feeding skills. iv ACKNOWLEDGEMENT Foremost, I would like to express my sincere gratitude to Rutgers, The State University of New Jersey and the National Museums of Kenya, who were very instrumental facilitating my PhD training. I am also thankful to Institutional Animal Care and Use Committee Rutgers University, Kenya National Council of Science Technology & Innovation, and Kenya Wildlife Service who approved my field study and issued necessary research permits. My research could not be possible without their endorsement. My entire research work could not have happened without the financial support by different funding agencies that supported my field and laboratory work. Thus, I am highly indebted to The Leakey Foundation, Paleontological Scientific Trust (PAST), Conservation International’s Primate Action Fund, Primate Conservation Inc., International Primatological Society, Nacey Maggioncalda foundation, Rutgers Center for Human Evolutionary Studies, Rutgers Department of Anthropology, Harvey Waterman Students Research Fund, and Primate Nutrition Ecology Laboratory at Hunter College New York. Moreover, it would be hard to go unrecognized the effort of different people in these organizations who helped tirelessly in the grants application process as well as encouraging me even when the funds were not forthcoming, and for showing interest in my work and linking me to various funding organization to solicit financial support for my work. In particular, I am thankful to Prof. Jack Harris, Antony Ryland, Noel Rowe, Paddy Moore, Anne Maggioncalda, Ella Outlaw, and Janette Wallis. v Besides my dissertation committee, the professional guidance and technical support in undertaking my dissertation would not be complete without help or guidance from Peter Lucas, Mark Wegner, Scott McGraw, Julie Wieczkowski, Peter Morin, Chris Free, Nancy Moinde, Padimin Iyer, Emily Lynch, Lisa Danish, and Jack Harris. I salute them all. Special thanks also goes to Dr. Robert Scott, who through the course I took with him on hominin diet inspired my study. Jungle life by the waters of Tana River could have been gruesome, where the Nile crocodiles crave for human and nonhuman primate blood while the African elephants and savanna buffaloes fearlessly fight intruders in their territory. However, my field research assistants, Kenya Wildlife Rangers and staff, and my Pokomo friends were instrumental for maintaining my safety and comfort both at Mchelelo and Kitere field sites. They included; Said Rhova, Komora Said, Dabasa Mohamed, Hussein Galana, Buya Mohamed, Mwalimu Bahatisha, Dadho Bahatisha, Coprol Kitheka, Warden Simon, Abio Gafo, Omar Kakawa, Aisha Abae, and Abed Said the camp manager. My research was also endorsed and supported by the Ndera Community Conservancy who allowed me to carry out my research within their area in Kitere and showed a lot of interest in my work. Father Ndegwa at St. John the Baptist Catholic Church in Wenje was a crucial pillar to my spiritual nourishment and made my life feel complete despite the jungle challenges away from my family and amid the terror threats of Somali militia. My dream to share my knowledge with the young generation and make some impact in conserving the Tana River biodiversity would have been far from realization if it were not for the unwavering support and invitation by local schools vi to educate the young people. I am truly thankful to head teachers in Maroni, Baomo, Makere, and Hara primary schools for invitation to their school to educate the pupil’s on conservation. Undertaking my research and coordinating the community outreach program would have worn me out without the tremendous support I received from my two Kenyan students, Charles Maingi Kivasu (MSc. Student University of Nairobi) and Janeserah Mukundi (Undergraduate student of Chuka University). Training them and their participation in helping me in my field research made a great progress towards achieving my goals, besides building the local research capacity in primatology. The fieldwork would not have been possible without successful completion of my coursework. This was made possible through a healthy life at graduate school at Rutgers. The graduate students and faculty fraternity in Department of Anthropology always made me feel home away from home and were always helpful whenever their assistance was needed. Writing the dissertation was the most difficult part in my training, learning the statistics and editing my work was indeed difficult, but special thanks goes to Shauhin Alavi, Alex Pritchard, Rebbeca Ferrell, Fred Foester, Timothy Bransford, and Chris Free whom their amazing encouragement and support was instrumental in overcoming these challenges. I too wish to express my heartfelt appreciation to all the departments’ support staff particularly Penny Burness, Marilyn Reyes, Ginny Caputo, Maydelle Romero, Shellie Harden, Jovani Reaves, and Bibi
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