The Feeding, Ranging, and Positional Behaviors of Cercocebus torquatus, the Red- Capped Mangabey, in Sette Cama Gabon: A Phylogenetic Perspective. Dissertation Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Catherine Agnes Cooke, M.A. Graduate Program in Anthropology The Ohio State University 2012 Dissertation Committee: W. Scott McGraw, Advisor Debbie Guatelli-Steinberg Dawn Kitchen Jeff McKee Copyright by Catherine Agnes Cooke 2012 Abstract The feeding, ranging, and positional behaviors of Cercocebus torquatus (red- capped mangabey) were studied in Sette Cama, Gabon from 2008-2009. It has been argued by several authors that hard-object feeding is the key adaptation of the Cercocebus-Mandrillus clade and that durophagy influences many aspects of Cercocebus social and behavioral ecology. The goal of this study was to evaluate the impact of obdurate foods on C. torquatus adaptation and evolution by combining multiple lines of evidence. Additionally, this information was used to test the hypothesis that C. torquatus is the most primitive member of its genus and that C. torquatus is a sister taxon to Mandrillus. C. torquatus in Sette Cama fed predominantly on fruits and seeds, but seed consumption was higher than that reported for other C. torquatus populations. The C. torquatus diet in Sette Cama was dominated by foods with intermediate to high hardness values and their foods were comparable in hardness values to those reported in other studies. The consumption of obdurate foods remained constant throughout the wet and dry seasons, and C. torquatus showed no preference for any food group. This suggests that durophagy is habitual among this population and does not serve as a fallback strategy. ii C. torquatus in Sette Cama occupies a smaller home range than other Cercocebus species of similar group sizes. The ranging behaviors of C. torquatus were influenced by the seasonal distribution and availability of fruits, particularly the location of Sacoglottis gabonensis seeds. The lack of competing primate species in their habitat along with the intensive use of both terrestrial and arboreal resources by C. torquatus may explain their small home range. C. torquatus also frequently divided into subgroups. C. torquatus spent the majority of time on the ground followed by the understory (one to five meters) and forest canopy at heights of five to twenty meters. Contrary to predictions based on phylogenetic relationships and morphology, C. torquatus is possibly one of the least terrestrial of all Cercocebus species studied. They used quadrupedal locomotion most frequently during travel and feeding followed by climbing and leaping. The most common postural behaviors were sitting and quadrupedal standing. The feeding, ranging, and positional behavior data from this study supports a close relationship between C. torquatus, C. atys, and C. lunulatus within the Cercocebus- Mandrillus clade. C. torquatus and C. atys are similar in their large group sizes and reliance on hard-object foods such as Sacoglottis gabonensis seeds throughout the year. Previous cranio-facial studies specifically positioned C. torquatus as the sister taxon to Mandrillus. Nevertheless, the behavioral evidence suggests that C. atys occupies a more similar niche to mandrills than does C. torquatus. Since C. torquatus and Mandrillus iii species are sympatric throughout much of their distribution, several behavioral differences would be expected between these groups. This study does not refute the hypothesis that C. torquatus and Mandrillus are sister taxa. Their current sympatry and niche partitioning suggest they could have evolved from a common terrestrial ancestor by sympatric speciation. The recognition of a 1.5 to 2 million year old fossil species closely resembling C. torquatus and the behavioral ecology of C. torquatus suggests that the polarity of the features shared by C. torquatus and Mandrillus are ancestral rather than derived for this clade. iv Dedication To my mother, Agnes Cooke, for never giving up, Nathaniel Cooke Moussopo, my little mangabey, and in memory of Thomas Howard Cooke, Sr. v Acknowledgements I would like to thank Dr. W. Scott McGraw for your guidance and advising on this dissertation and throughout my graduate study career. Thank you for never giving up hope that I would finish. I also thank the other members of my candidacy and dissertation committee for their patience and support. I thank the Ministere d’Eaux et Fôrets of Gabon, CENAREST, and the village chief of Sette Cama for permission to study the mangabeys and live at the Brigade d’Eaux et Forets. I am indebted to the WWF- Gabon, in particular Bas Huijbregts and Bas Verhage, for their logistical support and assistance in this project. Thank you to all the members of “Equipe torquatus” including Joseph Ibinda Igouwe, Zico Ibamba Igalla, and Richard Moussopo Ibessa. Joseph, I would not have survived my two years in Gabon without your knowledge of the forest, companionship, humor, and good cooking. Richard, thank you for your helpful comments, map making skills, and your support and belief in my ability to finish this dissertation. I am indebted to the various family members and friends who helped fund this endeavour, gave me books for the field, or moral support when I needed it! In particular I thank Agnes Cooke, Mary Cooke-Hall, Tom Cooke, Mary Adduci, Bonnie Martuarano, Ariane Payen, Lucy Keith, Elisabeth Hellmer, Michelle Rodrigues, and Indus Films. Thanks to Mary Cooke-Hall, Patrick Hall, Tom Cooke, Mary Beavers, and Elisabeth Hellmer for housing my cats at one time or another while I was off chasing mangabeys. This research was funded by grants from The Ohio State University (Department of Anthropology, Office of International Affairs, Alumni Grant, OSU-Sigma Xi), Sigma Xi, Primate Conservation, Inc., and the International Primatological Society. vi Vita 1999………………………………………………B.A., Anthropology, University of Missouri-Columbia 2003………………………………………………M.A., Anthropology, The Ohio State University Publications McGraw WS, Cooke C, and Shultz S. 2006. Primate remains from African Crowned eagle (Stephanoaetus coronatus) nests in Ivory Coast’s Tai Forest: Implications for primate predation and early hominid taphonomy in South Africa. Amer J Phys Anthropol 131:151-165. Newell E, Guatelli-Steinberg D, Field M, Cooke C, and Feeney R. 2006. Life history, enamel formation, and linear enamel hypoplasia in the Ceboidea. Amer J Phys Anthropol 131:252-260. Fields of Study Major Field: Anthropology vii Table of Contents Abstract ................................................................................................................................. ii Dedication ............................................................................................................................. v Acknowledgements............................................................................................................... vi Vita vii List of Tables ......................................................................................................................... xi List of Figures ....................................................................................................................... xv Chapter One: Introduction .....................................................................................................1 1.1 Introduction ..................................................................................................................... 1 1.2 Species Profile: Cercocebus torquatus torquatus Kerr (1792) ........................................ 1 1.3 Background on the Cercocebus-Mandrillus clade ............................................................ 7 1.4 Cercocebus-Mandrillus Morphology and Hard-object Feeding Adaptations................. 21 1.5 Cercocebus-Mandrillus Biogeography and Evolution .................................................... 26 1.6 Cercocebus-Mandrillus: The impact of obdurate feeding? ........................................... 29 1.7 Research Questions ....................................................................................................... 30 1.8 Organization of dissertation .......................................................................................... 31 Chapter Two: General Methods and Basic Data .................................................................... 42 2.1 Introduction ................................................................................................................... 42 2.2 Background .................................................................................................................... 42 2.3 Research Questions ....................................................................................................... 48 2.4 Methods ......................................................................................................................... 48 2.5 Data Analyses ................................................................................................................. 61 2.6 Results ............................................................................................................................ 64 2.7 Discussion ...................................................................................................................... 76 2.8 Conclusions ...................................................................................................................