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UNIVERSITY of CALGARY Determinants of Group Size and Composition for Colobus Vellerosus at Boabeng-Fiema, Ghana

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UNIVERSITY of CALGARY Determinants of Group Size and Composition for Colobus Vellerosus at Boabeng-Fiema, Ghana UNIVERSITY OF CALGARY Determinants of Group Size and Composition for Colobus vellerosus at Boabeng-Fiema, Ghana: Ecological versus Social Factors by Julie A. Teichroeb A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY DEPARTMENT OF ANTHROPOLOGY CALGARY, ALBERTA MARCH, 2009 © Julie A. Teichroeb 2009 UNIVERSITY OF CALGARY FACULTY OF GRADUATE STUDIES The undersigned certify that they have read, and recommend to the Faculty of Graduate Studies for acceptance, a thesis titled “Determinants of Group Size and Composition for Colobus vellerosus at Boabeng-Fiema, Ghana: Ecological versus Social Factors” submitted by Julie A. Teichroeb in partial fulfillment of the requirements for the degree of Doctor of Philosophy. ___________________________________ Supervisor, Dr. Pascale Sicotte, Department of Anthropology ___________________________________ Dr. Linda Fedigan, Department of Anthropology ___________________________________ Dr. Robert Longair, Department of Biological Sciences ___________________________________ Dr. Warren Wilson, Department of Archaeology ___________________________________ External Examiner, Dr. Rasanayagam Rudran, Scientist Emeritus, Smithsonian Institution _____________ Date ii ABSTRACT Intense within-group food competition is not thought to influence social organization for folivorous primates. Thus, they are expected by the socioecological model to live in large groups, exploiting benefits such as predation avoidance. However, many folivores form small groups, well below the threshold to avoid within-group scramble competition for food (the “folivore paradox”, Janson & Goldsmith, 1995). Social factors, rather than ecological factors, are thought to determine social organization for folivores, though this has rarely been systematically tested. In this study, the relative contribution of ecological versus social factors on the group size and composition of ursine colobus monkeys (Colobus vellerosus ) at the Boabeng-Fiema Monkey Sanctuary, Ghana was explored. Four groups were followed for 22-months (from 2002-2005) and behavioural observations were done using focal-animal sampling and ranging scans. Phenology, a large-tree survey, and a quadrat survey were used to determine home-range quality. It was found that within-group scramble competition for food constrained group size, despite a highly folivorous diet. Social factors also constrained overall group size and appeared to have more influence on group composition. High-ranking males used infanticide and other aggressive behaviours to influence female mate choice. Female dispersal, forced female emigration, and female resistance to female immigration indicated female competition for group membership. Therefore, females actively sought to maintain small group sizes. Within groups, females mated promiscuously, despite differing male agonistic display outputs, which appeared to indicate fighting ability. Promiscuous mating was likely a female counter-strategy to the high rate of infanticide. However, females may have attempted to immigrate into groups where the male(s) won in previous encounters with the male(s) in the females’ original group. This suggests a female preference for residing with strong males and may be linked to protection against infanticide. Theoretical examination of the folivore paradox has suggested that infanticide risk should affect females at smaller group sizes than increased food competition, which has a detrimental effect at larger group sizes. Colobus vellerosus seems to conform to this idea, in that social factors appear to constrain group size at an earlier point than ecological factors while also showing more effect on group composition. iii ACKNOWLEDGEMENTS This dissertation could not have been completed without the help of many people. My supervisor, Dr. Pascale Sicotte, was wonderfully supportive throughout the entire process. I would also like to thank the rest of my supervisory committee, Dr. Linda Fedigan and Dr. Robert Longair for their support and helpful advice over the years. I am grateful to my external examiners Dr. Rudy Rudran and Dr. Warren Wilson for attending my defence and Dr. Josie Smart for being the neutral chair. Thanks go out to Dr. James Paterson and Dr. Drew Rendall who formed part of my candidacy exam committee. The staff in the Department of Anthropology at the University of Calgary, Susan Moisik, Monika Davidson, May Ives, Jill Ogle, and Julie Boyd helped me innumerable times, as did the heads of the department, Dr. Doyle Hatt and Dr. Mary Pavelka. Permission to conduct this research was given by the Ghana Wildlife Division and the management committee of the Boabeng-Fiema Monkey Sanctuary (BFMS). The chiefs and elders of Boabeng and Fiema aided me several times over my years of research there. Thank you to the Herbarium at the University of Legon, Accra for plant identification. For being my family in Ghana and for helping me so much over the years, my love goes out to Alfred, Bea, Eva, Vida, Jennifer, Seth, and Steven Annor-Donyina. In addition, my research would not have been possible without the hard work and more importantly friendship of my field assistants in Ghana: Tony Dassah, Robert Koranteng, Kwame Michael Duodo, Kwaku Eric Amponsah, Afia Boahen, Rachel Boratto, Moses Ampofo, Constance Serwaa, Charles Kodom, and Kwabena Samuel Pepra. Staff of the Ghana Wildlife Division at Boabeng-Fiema were also always very kind, my gratitude goes to Mercy, Isaac, Jonas, and the late Osei. My testosterone analysis was done at the Wisconsin National Primate Research Centre in the laboratory of Dr. Toni Ziegler with the help of Dan Wittwer. Sarah Carnegie also aided in the interpretation of my testosterone results. The Hormone laboratory at the Wisconsin National Primate Research Center, University of Wisconsin- Madison was made possible in part by Grant Number P51 RR000167 from the National Center for Research Resources (NCRR), a component of the National Institutes of Health (NIH). This research was conducted in part at a facility constructed with support from Research Facilities Improvement Program grant numbers RR15459-01 and RR020141- iv 01. The chapter on female dispersal patterns in this dissertation (Chapter Four) benefited from additional research assistance by Rachel Boratto, Johanna Hedlund, Danica Stark, and Lucy Anderson. Tania Saj, Andrew MacIntosh, Sarah Marteinson and Lauren Brent provided contributions to the database on dispersals and immigrations at BFMS. A big thank you to all the people who I spent time with in the field: Nikki Porter, Sarah Wong, Tania Saj, Lauren Brent, Sarah Marteinson, Eva Wikberg, Rachel Boratto, Dan Barthmeier, Joshua Middleman, Paige Shaw, Anne Dixon, Peter Hinchcoiffe, Bright Kankam, Yehudi Fleising, Evelyn Tan, and Scott Crawford. We laughed, cried, got parasites and malaria, but most of all cared for one another and had a great time. A special note of thanks to Tracy Wyman who was always just yelling-distance away from helping me with stats, computers, GIS, etc. during my write up process. Thank you to Greg Bridgett and Dr. Tak Fung for help with statistics. For reviewing papers and giving helpful advice I’d like to thank Dr. Tania Saj, Dr. Kathy Jack, Dr. Steig Johnson, Dr. Colin Chapman, Dr. Joanna Lambert, Dr. Lynne Isbell, Eva Wikberg, Sarah Carnegie, Dr. Hugh Notman, Alison Behie, Amanda Melin, Fernando Campos, Mackenzie Bergstrom, Kira Delmore, Sara Martin, and several anonymous reviewers. Funding for this project was provided by the Natural Sciences and Engineering Research Council of Canada (NSERC), the Department of Anthropology, Research Services, and the Faculty of Graduate Studies at the University of Calgary, the Province of Alberta, and the American Society of Primatologists. To my family and friends, Alan and Michael Teichroeb, Mary Ann and Jake Teichroeb, Mary and Robert Schalla, Kara and Scott Wilson, Christine Duffey, Brent Watson, Ritchie Dong, and Wilson Wong, thanks for all your support. Thank you to Greg Bridgett, for your help, love, and support over the years. This work seemed so much easier because I had you there to peel me off the walls, calm me down, and set me on course. Finally, all my gratitude goes to my parents, Herman and Susan Teichroeb, for their unconditional love and support and for telling me I could accomplish anything. v TABLE OF CONTENTS Approval Page…………………………………………………………………………….ii Abstract…………………………………………………………………………………...iii Acknowledgements……………………………………………………………………….iv Table of Contents…………………………………………………………………………vi List of Tables……………………………………………………………………………...x List of Figures…………………………………………………………………………….xi 1.0 Chapter One: The Research Problem and an Overview of the Dissertation…….1 1.1 Introduction to the Research Problem…………………………………………2 1.1.1 The “Folivore Paradox”…………………………………………...4 1.2 The Research Objective……………………………………………………….6 1.2.1 Research Questions and Hypotheses………………………………..8 1.3 The Study Site…………………………………………………………………9 1.3.1 History of the Boabeng-Fiema Monkey Sanctuary..........................11 1.4 The Study Species............................................................................................12 1.5 Research Significance………………………………………………………..15 1.6 Limitations of the Study……………………………………………………..16 1.7 Overview of the Dissertation………………………………………………...17 2.0 Chapter Two – Test of the Ecological-Constraints Model on Ursine Colobus Monkeys ( Colobus vellerosus ) in Ghana………………………………………20 2.1 Abstract………………………………………………………………………21
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