DOCTORAL THESIS the Behavioural Ecology of Personality in Wild Barbary Macaques Tkaczynski, Patrick
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DOCTORAL THESIS The Behavioural Ecology of Personality in Wild Barbary Macaques Tkaczynski, Patrick Award date: 2017 General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Download date: 07. Oct. 2021 The Behavioural Ecology of Personality in Wild Barbary Macaques By Patrick John Tkaczynski BSc (Hons), MRes A thesis submitted in partial fulfilment of the requirements for the degree of PhD Department of Life Sciences University of Roehampton 2016 1 2 “A personality is the product of a clash between two opposing forces: the urge to create a life of one's own and the insistence by the world around us that we conform.” Hermann Hesse Soul of the Age: Selected Letters, 1891-1962 3 Abstract Personality, that is intra-individual consistency and inter-individual variation in behaviour, is widespread throughout the animal kingdom. This challenges traditional evolutionary assumptions that selection should favour behavioural flexibility, and that variation in behavioural strategies reflects stochastic variation around a single optimal behavioural strategy. Adaptive models to explain personality within the framework of evolutionary and behavioural ecology exist, and are typically empirically explored by identifying proximate associations to, and the functional consequences of, personality expression. To date, such studies have typically quantified a narrow range of personality traits within a species, and focused on captive populations or species with relatively limited behavioural or social repertoires. In this thesis, personality is studied in wild Barbary macaques (Macaca sylvanus). Quantification of personality structure in the species was conducted using a multi-method approach, and subsequently, it was examined whether physiological stress response (a proximate association) was related to personality expression, and whether personality expression affected social (functional) outcomes for individuals. Seven personality constructs were identified in Barbary macaques. Three personality constructs were related to physiological stress responses (Excitability, Tactility and Exploration), with the relationship between stress and personality expression dependent on sex, and in some cases rank or age. Two personality constructs (Excitability and Exploration) were associated with measures of social integration. Subjects generally socially assorted themselves according to personality, tending to be in proximity to individuals with a similar personality to themselves. This study contributes methodologically by 4 demonstrating the plausibility of multi-method approaches to measuring personality in wild primates, and empirically, by generating evidence supporting adaptive models for the evolution of personality, namely that intra-individual consistency in behaviour may be mediated by physiology and that inter- individual variation in behaviour has functional benefits in the formation of social relationships and social structures. 5 Table of Contents Abstract .................................................................................................. 4 List of Figures ....................................................................................... 12 List of tables ......................................................................................... 15 Acknowledgements ............................................................................... 21 1. Introduction ...................................................................................... 24 1.1 The Concept of Personality ................................................................ 25 1.2 Adaptive Models for the Origin and Maintenance of Personality .............. 31 1.3 Quantifying Personality in Animals ..................................................... 35 1.4 Mechanistic Associations of Personality ............................................... 38 1.5 Functional Outcomes of Personality .................................................... 42 1.6 Present Project and Aims .................................................................. 46 2. General Methods ............................................................................... 49 2.1 Study Species ................................................................................. 49 2.2 Study Site and Population ................................................................. 51 2.3 Study Subjects ................................................................................ 54 2.4 Data Collection ................................................................................ 58 2.4.1 Behavioural Sampling ................................................................. 58 2.4.2 Female Reproductive State .......................................................... 63 2.4.3 Field Data Collection Time Periods ................................................ 64 2.5 Dominance Hierarchy Calculations ................................................... 65 2.6 Personality Assessment .................................................................... 66 6 2.7 Stress Physiology Data ..................................................................... 71 2.7.1 Faecal Sample Collection ............................................................. 72 2.7.2 fGC Extraction ........................................................................... 72 2.7.3 Enzyme Immunoassay ................................................................ 73 2.8 Social Network Analysis .................................................................... 75 2.9 Statistics ........................................................................................ 78 2.9.1 Repeatability Analysis ................................................................. 78 2.9.2 Modelling Approaches ................................................................. 78 3. Quantifying Personality in Barbary Macaques: A Multi-Method Approach ............................................................................................... 81 3.1 Introduction .................................................................................... 82 3.1.1 Aims ......................................................................................... 87 3.2 Methods ......................................................................................... 88 3.2.1 Quantifying Personality Constructs ............................................... 88 Behavioural Coding .................................................................................. 88 Trait Rating ............................................................................................. 91 Experimental Assays ................................................................................ 92 Personality Expression and Experimental Design .......................................... 97 3.2.2 Determining Intra-Individual Consistency and Inter-Individual Variation in the Expression of Personality Constructs .............................. 98 3.2.3 Relation of Behavioural Variables to Constructs Generated Outside of a Natural Setting ................................................................................... 99 7 3.2.4 Testing Convergent and Discriminant Validity of Personality Constructs ........................................................................................................ 99 3.2.5 Determining Barbary Macaque Personality Structure ....................... 99 3.3 Results ......................................................................................... 100 3.3.1 Personality Constructs .............................................................. 100 Behavioural Coding ................................................................................ 100 Trait Rating ........................................................................................... 105 Experimental assays ............................................................................... 110 3.3.2 Inter-Individual Variation and Intra-Individual Consistency in the Expression of Personality Constructs ................................................... 123 Intra-Individual Consistency .................................................................... 123 Inter-Individual Variation ........................................................................ 124 3.3.3 Relation of Behavioural Variables to Trait Rating- and Experimental Assay-Derived Personality Constructs.................................................. 133 3.3.4 Convergent and Discriminant Validity of Personality Constructs ...... 138 3.3.5 Barbary Macaque Personality Structure ....................................... 138 3.4 Discussion .................................................................................... 143 3.4.1 Personality Constructs of Wild Barbary Macaques ........................