Inter-individual Relationships of Female Rhesus M acaques (M a c a c a m u l a t t a ) BY K r ist in E lise P hillips D o c t o r of P h il o so ph y IN A nthropology AT University C ollege London 1998 ProQuest Number: U642096 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest. ProQuest U642096 Published by ProQuest LLC(2015). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. Microform Edition © ProQuest LLC. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 A b s t r a c t Rhesus macaques {Macaca mulatta) live in large, female philopatric social groups containing several lineages of related females and multiple breeding males who have generally immigrated from other groups. In this study, I examined the social relationships between 27 female rhesus macaques and other adults living in a single, free ranging group on Cayo Santiago, Puerto Rico. I first examined the time budgets of rhesus macaque females in order to investigate how variation in time for activities affect each female's strategies in maintaining relationships with important allies and to see whether females of different classes — particularly rank, age, and kinship — differed in their time allocation patterns. In this study, females spent an average of only 12.5% of their time engaged in subsistence activity {eat, drink, and forage), while sleep and rest together constituted 77.3% of the female activity budget. Thus rhesus macaque females did not seem to face time constraints in meeting their subsistence needs. Nonetheless, there were some class effects on female time budgets, particularly rank. High ranking females spent more time eating foods concentrated at feeding hoppers and drinking from water spigots than did either middle or low ranking females. Both of these results were obtained whether females were alone or with other individuals. In fact, high ranking females spent more overall time in social contact with other animals than did lower ranking females. I next identified the set of preferred partners {affiliates and associates: sensu Smuts 1985) with whom the focal female had close social relationships, using proximity scans collected at 10 minute intervals during focal samples in concert with more detailed data on grooming interactions among dyads. Focal females had between 1 and 5 preferred female partners and between 1 and 6 preferred male partners, and neither female rank, nor age, nor availability of close kin had a significant effect on the number of preferred partnerships a female participated in. Female preferred partners tended to be close kin or members of the female’s cohort (bom within a year of her). Whether kin or nonkin, female preferred partners tended to be dominant individuals. Male preferred partners also tended to be high ranking. Moreover, preferred males typically were those with longer periods of tenure in the group and, in contrast to female partners, were nearly all nonkin. These same general patterns hold even considering only important preferred partnerships, or those that the focal female was responsible for maintaining. Finally, I tested several hypotheses regarding the function of relationships for female rhesus macaques: what benefits do these females receive from participating in these relationships? One major hypothesis suggests that females appear to form relationships in order to improve their access to clumped resources such as monkey chow in the feeding hoppers or water from drinking troughs. Predictions based on this hypothesis, however, were not substantiated. A second major hypothesis suggests that females gain a benefit from partnerships in terms of protection from general aggression. Several findings argue in favor of this hypothesis: [1] females received less aggression from preferred partners then from other individuals (when the amount of time spent with each partner type is taken into account), and [ 2] females were aided more frequently by both female and male preferred partners than they were by other animals. Two other hypotheses — that preferred partners benefit females in the care and protection of their infants and that preferred male partners benefit females as mates — have mixed results. T h is t h e sis is d e d ic a t e d t o th e l o v e a n d k in d n e s s of A lice a n d E d w a r d N e e l y AND R o b e r t a a n d Ca r l B oegershausen 111 A cknowledgements The generous help and care of many individuals has contributed to the completion of this thesis. First, it is important to thank Robin Dunbar. He has been very kind in dealing with a self-funding student who typically veered far from the path of least resistance in completing a thesis. He has also been a supportive and inspirational mentor — many, many thanks. Special gratitude must be extended to Fred Brett for his kindness as my supervisor in London during the latter part of the thesis and for wading through a multitude of administrative questions for me. Thank you! The Biological Anthropology Group at University College London, including Leslie Aiello, Catherine Homewood and Simon Strickland, has been wonderfully kind and supportive over the years. Thank you for thought-provoking seminars and meetings over the teakettle. In addition, I would like to thank many of the postgraduates at UCL, particularly Heidi Marriott, Guy Colishaw, Bettina Greiser and Craig Roberts. The Committee of Vice Chancellors and Principals of the Universities of the United Kingdom granted me an Overseas Research Studentship Award that made working towards a doctorate in the U.K. possible. As this doctorate has been a rewarding experience, I greatly appreciate both the financial assistance provided and faith in me by this organisation. A grant from the Sigma Xi partially funded earlier research on female macaques among the Arashiyama West snow monkeys. I would like to thank John Berard and Matt Kessler for permission to collect data on the rhesus macaques of Cayo Santiago (CPRC) and for access to such a wonderfully rich research site. A thesis on detailed social relationships would not have been possible without the years of work that they and countless other individuals have devoted to Cayo. Thank you. John Berard and the staff of CPRC also provided necessary logistical support. Finally, the staff that maintained the island and censused the monkeys were wonderfully kind to me during my stay in Punta Santiago. Thank you to Edgar Davila, Guelo Figueroa, Hector Rolon, Hector Vasquez, Tito Vasquez and William Walker for your cheery smiles, mangoes and help — again, none of this would have been possible without your hard work. The Anthropology Department of the University of California, Davis has been extremely supportive throughout the write-up stage of this thesis. A very special thanks to Peter Rodman, Henry McHenry, Sandy Harcourt, Kelly Stewart, Monique Borgerhoff Mulder, Sarah Hrdy and Lynne Isbell for inviting me to seminars and classes, allowing me to work as a teaching assistant and making me a part of the large primatological community at Davis. I hope to some day repay each for their kindness. Discussions with and the support of many students in Davis were crucial to finishing this work, and Mary Towner, Barney Luttbeg, Kimber Haddix, Lore Ruttan and Anthony Di Fiore contributed immeasurably with ideas and help in our weekly dissertation workshop. Finally, a special thank you must be said to Mary Monroe Rodman for her very timely help and kindness over the last few years. Ximena Bans, Ivan Cornejo, Peter English, Robin Foster, Else Magord, Liz Marberry, Sean O'Donnell, Peter Rodman, Michelle Rogers, Lore Ruttan, Brian Smith, Jens-Christian Svennig and especially Anthony Di Fiore lent support during the two years spent in Ecuador. Many other visitors provided welcome social outlet to living in Yasuni National Park: thank you. The Universidad Catolica provided wonderful living facilities in Yasuni National Park under the guidance of Laura Arcos Teran. Maxus IV Ecuador, especially those stationed at the Northern Production Facility, provided logistical support, friendship and a much-needed good meal from time to time. Finally, thank you to Luis Albuja for supporting my application for an intercultural visa and the chance to live in Ecuador. My network of friends has been very supportive of me in immeasurable ways. Thank you to Lisa Collins, Pete & Chris Coppolillo, Leslie Digby, Fran Dolins, Eduardo Femandez-Duque, Sara Hansen, Craig Kirkpatrick, Lynne Knight, Heidi Marriott, Drew & Karen Rendall, Laurel Saito, Mary Towner, Claudia Valeggia, Kate Walters, Sylvia Wikler, Liz Williamson, Use Wolfram and especially Michael Weiss for the visits, letters, tapes, books, discussions and friendship. Lynne Fukuda, Steig Johnson, And Kazem, Mairi Knight and Jennifer Lippold were my oxygen on Cayo Santiago — from making me laugh over monkey antics to joining me on quick trips to San Juan. A special thank you must be said to Alyn Brereton. He was an incredibly supportive friend throughout the time collecting data for this thesis. My family has also been supportive in many, many ways: thank you to Elise Phillips, Augie Phillips, Patricia Ortega, John Phillips, Edward Neely Jr., Alice Neely, Matthew Di Fiore, Amy Di Fiore and Josephine Gemingiani. Those to whom this thesis is dedicated - my grandparents and great aunt & uncle - provided me with financial assistance when it was most needed.