Abstract of the Dissertation…………………..……………………………… X
Total Page：16
File Type：pdf, Size：1020Kb
UNIVERSITY OF CALIFORNIA, SAN DIEGO Why Do Females Live in Groups? Dominance, Grooming, and Experimental Testing of the Effects of Food Distribution on the Behavior of Commensal Bonnet Macaque (Macaca radiata) Females A dissertation submitted in partial satisfaction of the requirements for The Degree of Doctor of Philosophy in Anthropology by Sandra Rose Chacko Committee in Charge: Professor James Moore, Chair Professor Christine Harris Professor Donald Macleod Professor Margaret Schoeninger Professor Shirley Strum 2014 © Sandra Rose Chacko, 2014 All rights reserved The Dissertation of Sandra Rose Chacko is approved, and it is acceptable in quality and form for publication on microfilm and electronically: Chair University of California, San Diego 2014 iii DEDICATION This dissertation is dedicated to my parents John and Blanche McLaughlin for their unwavering love and support and to my nephew Jacob Murphy Douglas, future primatologist. iv TABLE OF CONTENTS Signature Page……………………………………………………………… iii Dedication…………………………………………………………………… iv Table of Contents…………………………………………………………… v List of Graphs………………………………………………………………. vi List of Tables……………………………………………………………….. vii Acknowledgements…………………………………………………………. vii Vita………………………………………………………………………….. ix Abstract of the Dissertation…………………..……………………………… x Introduction…………………………………………………………………. 1 Chapter1: Differences in dominance interactions among commensal female bonnet macaques (Macaca radiata) under natural foraging and experimental feeding conditions in Mysore, India…………..…………………………… 14 References………………………………………………………….. 45 Chapter 2: Grooming reciprocity among commensal female bonnet macaques (Macaca radiata) in Mysore, India……………..………………. 50 References………………………………………………………… .. 81 Chapter 3: Experimental Tests of the Effects of Food Distribution on the Behavior of Commensal Bonnet Macaque (Macaca radiata) Females……………………………………………………………………… 85 References…………………………………………………………… 113 v LIST OF GRAPHS Graph 1.1a: Normalized David Scores weighted by number of encounters (Dij) by rank for both the Non-Experimental and Experimental Observations….. 41 Graph 1.1b: Normalized David Scores (Pij) weighted by rank for both the Non-Experimental and Experimental Observations………………………… 42 Graph 1.2: Female Dominance on an Interval Scale………………………… 43 Graph 1.3: Distribution of Modified Landau Indices (MLI) by frequency for Macaque grades 1-3 as categorized by [Thierry, 2000]……………………… 44 Graph 2.1a: Grooming Given By Rank……………………………………… 71 Graph 2.1b: Grooming Received by Rank…………………………………… 72 Graph 2.2a: Grooming Reciprocity within Individual Bouts……………….. 73 Graph 2.2b: Grooming Reciprocity within Reciprocal Bouts……………… 74 Graph 2.3: Grooming Reciprocity across Bouts for Reciprocating Dyads……………………………………………………….. 75 Graph 2.4: Percentage of Grooming of a Subordinate as a Function of Rank Distance…………………………………………………. 76 Graph 2.5a: Reciprocal and Non-reciprocal Grooming Given by Rank……………………………………………………………… 77 Graph 2.5b: Reciprocal and Non-reciprocal Grooming Received by Rank………………………………………………………… 78 Graph 2.6: Grooming of Adjacent and Non-adjacent Partners……………… 79 Graph 2.7: Grooming of High and Low Adjacent Partners………………….. 80 Graph 3.1: Kcal Consumption by Rank and Food Distribution……………… 110 Graph 3.2: Aggressive Acts by Food Distribution……………………………. 111 Graph 3.3: Time on platform vs. Aggression Received………………………. 112 vi LIST OF TABLES Table 1.1a: Dominance matrix: Non-Experimental Data…………………… 34 Table 1.1b: Dominance matrix: Experimental Data…………………………. 35 Table 1.2: Differences in Normalized David Scores (Dij) and Subsequent Assigned Rank for Individual Females During Non-Experimental Observations and During Feeding Experiments……… 36 Table 1.3a: Macaque Dominance Hierarchy Strengths……………………… 37 Table1.3b: Mean Dominance Hierarchies for Macaque Species……………. 40 vii ACKNOWLEDGEMENTS I would like to thank the University of California-San Diego Department of Anthropology for funding this project. I would also like to thank my thesis committee, especially Dr. Jim Moore and Dr. Shirley Strum for their comments and support. In India I would like to thank Dr. Mewa Singh for his invaluable guidance and support, without which this project could not have been completed. Thank you also to the University of Mysore for permission to study and work on their grounds. Also I owe a big thank you to the bonnet macaques. This research complied with protocols approved by the University of California, San Diego Institutional Animal Care Committee. This research also adhered to the legal requirements for animal welfare put forth by the Governments of India and the State of Karnataka. Chapters 1, 2, and 3 are being prepared for submission for publication of the material. The dissertation author was the primary investigator and author of this material. viii VITA EDUCATION 2000 BS magna cum laude Ecology & Evolutionary Biology University of Arizona 2000 BA magna cum laude Anthropology University of Arizona 2004 MA Anthropology University of California, San Diego 2006 PhD candidacy Anthropology University of California, San Diego 2014 PhD Anthropology University of California, San Diego APPOINTMENTS 2009-Present Adjunct Faculty Pima Community College Department of Sociology & Anthropology 2006-2007 Teaching Assistant University of California, San Diego Eleanor Roosevelt College 2006 Academic Reader University of California, San Diego Department of Religious Studies 2006 Teaching Assistant University of California, San Diego Department of Religious Studies 2004-2006 Senior Teaching University of California, San Diego Assistant Eleanor Roosevelt College 2003-2004 Teaching Assistant University of California, San Diego Eleanor Roosevelt College 2001 Teaching Assistant University of Arizona Department of Ecology & Evolutionary Biology PUBLICATIONS 2010. Chacko, S.R. Dominance Hierarchy and Social Grooming among Commensal Female Bonnet Macaques (Macaca radiata) in Mysore, India. Primate Research 26: 136. 2009. Chacko, S. R. Experimental Testing of Three Models of Female Behavior in a Group of Commensal Bonnet Macaques (Macaca radiata) in Mysore, India. American Journal of Primatology 71: 68. 2008. McLaughlin-Chacko, S. Experimental Testing of Three Models of the Evolution of Female Behaviour in a Group of Commensal Bonnet Macaques (Macaca radiata) in Mysore, India. Primate Eye ix ABSTRACT OF THE DISSERTATION Why Do Females Live in Groups? Dominance, Grooming, and Experimental Testing of the Effects of Food Distribution on the Behavior of Commensal Bonnet Macaque (Macaca radiata) Females by Sandra Rose Chacko Doctor of Philosophy in Anthropology University of California, San Diego 2014 Professor James Moore, Chair According to the socioecological model of female sociality, females living in environments with clumped resources should exhibit strong, linear dominance hierarchies and females in the upper part of these hierarchies should receive less aggression, gain better access to food and receive more grooming than their lower ranked counterparts. x This dissertation investigates the predictions of this model using females in a group of bonnet macaques (Macaca radiata). It also investigates the effects of three food distributions (0.25 m², 1 m², and 4 m²) on female aggression. This group had a moderately linear dominance hierarchy (0.63 Modified Landau Index). Although aggressive behavior increased during the feeding experiments, the overall strength of the dominance hierarchy was not significantly affected by an increase in food clumping. Higher ranked females did not receive more grooming than lower ranked females. Females engaged in both reciprocal (12.6%) and non-reciprocal bouts (87.4 %) of grooming and reciprocal grooming was time matched. Although most grooming was not immediately reciprocated, females did not appear to be trading grooming for another commodity, except, potentially, for coalitionary support. Grooming was more reciprocal across bouts than within bouts showing increased grooming reciprocity between partners over time. This long-term reciprocity suggests that bonnet macaque females reciprocate grooming over long spans of time effectively increasing the proportion of reciprocal grooming and decreasing the amount traded in a biological market. Females showed different types of competition as the experimental food distribution was decreased. High ranking females consumed more than low ranking females in each distribution. The greatest rank asymmetry in feeding was found in the highly clumped condition due to the ability of dominant individuals to monopolize the feeding platform. However, the number of aggressive acts among females was significantly lower in both clumped distributions than in the dispersed condition. Usurpation, as measured by agonistic acts, was higher in the dispersed condition, but xi competition through monopolization was greatest in the highly clumped condition as expected by the socioecological model. Competition in the highly clumped condition could not be measured by quantity of agonistic behavior illustrating the importance of distinguishing between monopolizability and usurpability when analyzing levels of competition. xii INTRODUCTION Sociality, or group living, is a major characteristic of the primate order. However this sociality comes with costs, particularly in the context of feeding [Alexander, 1974; Pulliam & Caraco, 1984]. These costs may be particularly high for females due to the increased energetic requirements that accompany gestation and lactation. Thus