AT the DUKE LEMUR CENTER by Kathleen Marie Mcguire B
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i THE SOCIAL BEHAVIOR AND DYNAMICS OF OLD RING-TAILED LEMURS (LEMUR CATTA) AT THE DUKE LEMUR CENTER By Kathleen Marie McGuire B.S., Georgia Institute of Technology, 2014 A thesis submitted to the faculty of the University of Colorado in partial fulfillment of the requirement for the degree of Master of Arts. Department of Anthropology 2017 ii This thesis entitled: The Social Behavior and Dynamics of Old Ring-tailed Lemurs (Lemur catta) at the Duke Lemur Center written by Kathleen Marie McGuire has been approved for the Department of Anthropology _____________________________________________________ Dr. Michelle L. Sauther (Committee Chair) _____________________________________________________ Dr. Herbert H. Covert _____________________________________________________ Dr. Joanna E. Lambert Date The final copy of this thesis has been examined by the signatories, and we find that both the content and the form meet acceptable presentation standards of scholarly work in the above mentioned discipline. IACUC protocol # A168-14-07 iii Abstract McGuire, Kathleen Marie (M.A. Anthropology) The Social Behavior and Dynamics of Old Ring-tailed Lemurs (Lemur catta) at the Duke Lemur Center Thesis directed by Professor Michelle L. Sauther There has been little emphasis within primatology on the social and behavioral strategies old primates might use to meet the challenges of senescence while maintaining social engagement, such as assuming a group role like navigator. Understanding how old primates maintain sociality can reveal how behavioral flexibility might have facilitated an increase in longevity within the order. Using focal sampling of old (N = 9, 10+ years) and adult (N = 6, <10 years) Lemur catta at the Duke Lemur Center, activity budgets, social interactions, and group traveling information were recorded and compared from May to August of 2016. I found that both male and female old lemurs maintained sociality in the group, with older females being more social than adults. I failed to support the second hypothesis that older individuals would have a behavioral profile distinct from adults. Finally, I found preliminary support that older females help care for a daughter’s offspring in the form of carrying. These results indicate that social manifestations of age and senescence depend on a myriad of factors such as environment, life history, and individual personalities. This research also reveals the importance of decoupling ideas of chronological age, being an old individual, with senescence because somatic decline depends on these other factors besides age. Understanding these complex interactions is essential as we strive to define senescence and explore how age has shaped evolutionary trajectories among primates. iv Dedication This thesis is dedicated to “Mom” Syble Sweat, a primate grandmother who taught me the importance of family and the pursuit of knowledge. v Acknowledgements First, I would like to thank my advisor Dr. Michelle Sauther for her patience and essential support during all stages of this research and thesis. I would also like to thank Dr. Herbert Covert and Dr. Joanna Lambert for their mentorship as I developed and executed this work. I am also incredibly grateful to the Duke Lemur Center staff for permitting me to conduct my research. In particular, I thank Dr. Sara Zher, Andrea Katz, Erin Shaw, and Dr. Erin Ehmke for their unwavering enthusiasm and for providing essential information about the histories of these lemurs. I am incredibly grateful to Dr. James Millette for taking the time out to discuss aspects of my data collection, management, and analysis. I would also like to thank my family Michael P., Michael C., and Oliver McGuire for their love and support. I appreciate the many hours of conversation with Brittany Miles and Elizabeth McMillian about statistics, old age, and writing that helped to foster this research and thesis. Getting the opportunity to pursue my passion for primate social behavior would not have been possible without Dr. Linda Green and Dr. Joe Mendelson and their research project at Zoo Atlanta. Finally, thank you to Charles Hughes who endures my endless discussion of primates and their social lives, while relentlessly encouraging me to pursue this work. Funding for this study was provided by the University of Colorado, Department of Anthropology and the University of Colorado Boulder Graduate School. This work would have not been possible without the support of these institutions. vi Table of Contents Chapter 1: Introduction ............................................................................................................... 1 Senescence .................................................................................................................................. 1 Why Studying “Old” Primates Matters ...................................................................................... 4 Old Social Involvement in Nonhuman Primates ........................................................................ 7 Individual Factors and Old Primate Social Behaviors .............................................................. 10 The Social Role of Older Primates ........................................................................................... 11 Lemur catta as a Suitable Model for Primate Aging ................................................................ 12 Studying Aging in Captivity ..................................................................................................... 13 Lemur catta Mortality at the Duke Lemur Center .................................................................... 14 Infants and Juveniles (Birth to 2 years) ................................................................................ 14 Adults (2-10 years) ............................................................................................................... 17 Old Adults (10+ years) ......................................................................................................... 18 Study Aims and Hypotheses ..................................................................................................... 19 Chapter 2: Methods .................................................................................................................... 21 Study Site .................................................................................................................................. 21 Study Animals ........................................................................................................................... 23 Behavioral Methods .................................................................................................................. 23 Data and Statistical Methods .................................................................................................... 26 Chapter 3: Results ...................................................................................................................... 28 Degree of Sociality ................................................................................................................... 28 Proximity ............................................................................................................................... 28 Physical Contact ................................................................................................................... 29 Activity Budgets ....................................................................................................................... 31 Locomotion (Travel and Movement) ..................................................................................... 31 Feeding and Foraging .......................................................................................................... 34 Resting .................................................................................................................................. 34 Frequency of Behaviors ............................................................................................................ 40 Agonistic Point Events .......................................................................................................... 40 Affiliative Point Events ......................................................................................................... 41 Grooming .............................................................................................................................. 42 Stink Fights ........................................................................................................................... 43 Vocalizations ......................................................................................................................... 44 vii Individual Factors ..................................................................................................................... 46 Dominance Rank ................................................................................................................... 46 Social Roles .............................................................................................................................. 48 Case Studies .............................................................................................................................. 48 Aracus ................................................................................................................................... 48 Sprite ....................................................................................................................................