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Behaviour, the Social Cercopithecus Biology BEHAVIOUR, BIOLOGY AND THE SOCIAL CONDITION OF CERCOPITHECUS AETHIOPS . THE VERVET MONKEY. S.G. TOLLMAN Submitted in partial fulfillment of the requirements f or the degree of Doctor of Philosophy in the Department of Psychology, University of Natal, Durban. September 1984. ACKNOWLEDGEMENTS. I should like to express my thanks and appreciation to all who assisted me. I am particularly indebted to the following: Mr C.O. Murray, senior lecturer in psychology at the University of Natal in Durban. His concern and generosity after taking over supervision of the project, and his invaluable statistical advice, is much appreciated. Former staff members and supervisors, Mr J. Lucas and Dr P. Krige. The inspiration of John Lucas as supervisor and mentor led to the initiation of the project, and his continued advice from England is gratefully recorded. Dr P. Krige efficiently supervised the launching of the research. Professor Pam Sharratt, head of the Department of Psychology at the University of Natal, Durban, for her critical comments on reading the draft of the thesis. Ted Tollman, my husband, of the Department of Architecture at the University of Natal in Durban, for assisting with the layout of this thesis, and without whom this project could not have been undertaken. Peter Tollman, B.Sc. (Electronic Engineering), for his guidance with the analysis of the body temperature data, and computerised graphical presentations. Professor K.F. Poole and Mr J. Watts of the Department of Electronic Engineering at the University of Natal, Durban, for monitoring the characteristics of the temperature sensitive transmitters used in the project. Professor T. Downing and Mr T. Klomfass of the Natal Institute of Immunology, for providing assistance, ad~iCe, ::-,\~nd facilities to conduct all operative procedures. _ -\~~ -'0. \ - \\ ·l Mr J. Mueskins, followed by Mr A.B. Stead; ~hfef Horticulturalists, University of Natal, Durban, and staff, for permitting. construction of the dome in the Natal University gardens) and ongoing cooperation. My colleagues Professor R. Miller, Dr P. Henzi, Dr A. Bentley, David Basckin, and all the staff, for their continued support and encouragement. Mr B.H. Clark, architect, for drawing the diagrams. Mr K. Devadas, of the Department of Architecture, for his photographic work. Professor D.A. Scogings, former head of the Department of Survey, University of Natal, Durban, for providing special aerial photographs. The technicians, Mr P. Goodburn, Mr F. Sutton and Miss P. Daniel, who built and have maintained the cage. Marcia Lockett for her helpful comments and efficiency in typing this project. This project was supported by a grant from the University Research Fund. Opinions expressed and conclusions reached are those of the author. 11 ABSTRACT Biotelemetry has been coupled with an ethological approach to investigate a postulate that the physiological, behavioural, and social functioning of individuals coact in order to maintain homeostasis in an everchanging environment. Attention was focussed upon body temperature, behaviour, and the social situation, as they occurred together in each of five 'undisturbed' adult vervet monkeys. One male and one female were housed alone in single cages, and the other three, all females, were part of a natural troop that live together in captivity in a 6.5 metre radius geodesic dome. Intensive studies, including in-depth and multiple repeated measures at each level of functioning permitted comparisons between and within subjects, so that the connections between body temperature, and individual and social behaviour, could be scrutinised. Techniques for observation, data processing, and factor analysis have been considered, and procedures to facilitate the organisation and interpretation of information are suggested. The results pointed to individual variations superimposed upon a rhythmic underpinning of all the functions monitored. A synthesis of the data of body temperature with individual and social behaviour supports the contention that the individual responds to the vagaries of the environment as an integrated system within which the different levels of functioning are linked. It was found that the oscillation in body temperature was greater in the vervets that lived alone than in the vervets that lived in a troop. These results were supported behaviourally since the isolated subjects could only complement autonomic thermoregulatory responses with individually based behavioural strategies, whereas their troop-living conspecifics could utilise both individual and socially directed behavioural mechanisms. Within the troop an inverse relationship between body temperature variation and social status was revealed that is, the lower the status of the subject, the more the body temperature fluctuated around the mean. Behaviourally, it was found that the lower the status of the subject, the more difficult it became to gain access to resources, and to manipulate interpersonal space. In addition, harassment by troop III conspecifics increased and, consequently, the efficiency with which behavioural patterns could be executed, was decreased. An analysis of the data also led to the proposal that social grooming has evolved as a thermoregulatory mechanism; to the identification of three different facets of individual behaviour and of social behaviour; and to the idea that the rhythmic changes in the troop's spatial conformation reflected cyclical patterns in behavioural and social activity. lV CONTENTS Page CHAPTER ONE: INTRODUCTION 1 CHAPTER TWO : THE BACKGROUND TO THE STUDY 3 2.1 INTRODUCTION 3 2.2 STUDY OF STRATEGIES FOR ADAPTATION 3 2.3 STUDY OF FEEDING BEHAVIOUR : A BIOLOGICAL NECESSITY 5 2.4 INFORMAL STUDY OF ECOLOGY AND BEHAVIOUR 6 2.5 PRELIMINARY STUDY OF PSYCHOLOGY, PHYSIOLOGY AND THE SOCIAL GROUP 7 2.6 CONCLUDING COMMENT 8 CHAPTER THREE : THE SUBJECTS AND THE RESEARCH ENVIRONMENT 9 3.1 INTRODUCTION 9 3.2 CERCOPITHECUS AETHIOPS PYGERYTHRUS 9 3.2.1 Classification 9 3.2.2 Habitat and the adaptability of the vervet monkey 12 3.2.3 The appropriateness of Cercopithecus aethiops for the study of Homo sapiens 14 3.3 THE RESEARCH SUBJECTS 16 3.4 THE RESEARCH AREA: DURBAN, NATAL 17 3.4.1 The study area 20 3.5 CONCLUDING COMMENT 25 CHAPTER FOUR : THE RESEARCH OBJECTIVES 27 4.1 INTRODUCTION 27 4.2 OBJECTIVES 27 CHAPTER FIVE: THERMOREGULATION 29 5.1 INTRODUCTION 29 5.2 LITERATURE REVIEW 30 5.2.1 Body temperature 30 5.2.2 Circadian rhythms 35 5.2.3 Thermoregulation 42 v Page CHAPTER FIVE (CONTINUED) 5.3 PROCEDURE 51 5.3.1 Apparatus 51 5.3.2 Subjects 53 5.3.3 The recording schedule 54 5.3.4 Data collection 56 5.3.5 The implant operation 56 5.3.5.1 Capturing the subjects 56 5.3.5.2 The anaesthetic 57 5.3.5.3 Surgery 57 5.3.5.4 Recovery 59 5.3.6 The Minimitters 59 5.4 RESULTS 62 5.4.1 Body temperature means and standard deviations 64 5.4.2 Body temperature graphical representation of data 69 5.4.3 Body temperature analysis of variance 69 5.4.4 Body temperature Multiple regression on behavioural factors 73 5.5 DISCUSSION 74 5.6 CONCLUDING COMMENT 76 CHAPTER SIX : BEHAVIOUR 77 6.1 INTRODUCTION 77 6.2 LITERATURE REVIEW 78 6.2.1 The activity cycle 78 6.2.2 The categorisation of behaviour 82 6.2.3 Individual differences 85 6.3 PROCEDURE 88 6.3.1 Data collection 88 6.3.2 Data processing 89 6.4 RESULTS 100 6.4.1 Factor analytic treatment of the data 103 6.4.2 Multivariate analysis of variance of behavioural factors 111 6.4.2.1 Multivariate analysis of variance of functional categories 112 6.4.2.2 Multivariate analysis of vanance of derived factors 116 6.4.3 Individual behaviours 120 Vl Page CHAPTER SIX (CONTINUED) 125 6.5 DISCUSSION 126 6.5.1 The factors 6.5.2 The activity cycle 131 6.5.3 Individual differences 133 6.6 CONCLUDING COMMENT 134 CHAPTER SEVEN : THE SOCIAL GROUP 136 7.1 INTRODUCTION 136 7.2 LITERATURE REVIEW 137 7.2.1 Social organisation 137 7.2.2 Social structure 144 7.2.3 Spatial organisation 151 7.2.4 Aggression, dominance and hierarchy 158 7.2.5 Social roles 170 7.2.6 The adult female 173 7.2.7 Group integrative mechanisms 176 7.2.8 Grooming 179 7.3 PROCEDURE 182 7.3.1 Data collection 182 7.3.2 Data processing 182 7.4 RESULTS 184 7.4.1 The dominance hierarchy 184 7.4.2 Social interactions 184 7.5 DISCUSSION 190 7.6 CONCLUDING COMMENT 196 CHAPTER EIGHT : BEHAVIOUR, THERMOREGULATION AND THE SOCIAL GROUP 199 8.1 INTRODUCTION 199 8.2 LITERATURE REVIEW 200 8.2.1 Behaviour, thermoregulation and the social group 200 8.2.2 The theories of Satinoff, Luria and Salzen 200 8.2.3 Functions of the group 204 8.3 PROCEDURE 207 8.4 RESULTS 207 8.5 DISCUSSION 207 8.6 CONCLUDING COMMENT Vll Page CHAPTER NINE: AN EVALUATION OF THE RESEARCH 223 9.1 INTRODUCTION 223 9.2 METHODOLOGY 224 9.2.1 Behavioural and social data 225 9.2.2 Thermoregulatory data 230 9.3 CONSTRAINTS AND DIFFICULTIES 231 9.4 SUGGESTIONS FOR FUTURE RESEARCH 234 9.5 CONCLUDING COMMENT 235 CHAPTER TEN : SUMMARY AND CONCLUSIONS 236 GLOSSARY 241 APPENDICES : APPENDIX 1 : APPARATUS AND MEDICATION 243 1.1 Biotelemetry 243 1.2 Recording instruments 243 1.3 Surgery 243 APPENDIX 2 : BODY TEMPERATURES: INDIVIDUAL STRAIGHT LINE PLOTS ILLUSTRATING WEEKLY DAY LEVEL MEANS OVER A FIVE WEEK PERIOD. Plots are complementary to the three-dimensional plots of Graph 5.1. Subjects living alone 244 Subjects living in a troop 245 APPENDIX 3 : ETHOGRAM : NUMERICAL CODING FOR ALL .. BEHAVIOURS RECORDED DURING THIS RESEARCH PROJECT 246 APPENDIX 4: FUNCTIONAL CATEGORIES: BEHAVIOURS INCLUDED IN EACH CATEGORY 247 APPENDIX 5 : STATISTICAL ANALYSES: DETAILS OF PRINCIPAL AXES
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