Within-Group Consequences Of The Risk And Occurrence Of Between-Group Conflict In Crested Macaques (Macaca nigra ) James O Waterman A thesis submitted in partial fulfilment of the requirements of Liverpool John Moores University for the degree of Doctor of Philosophy March 2021 1 Contents Contents ........................................................................................................... 1 List of Tables ................................................................................................... 4 List of Figures .................................................................................................. 6 Abbreviations and Acronyms .......................................................................... 7 Abstract ............................................................................................................ 8 Declaration ...................................................................................................... 9 Acknowledgements ....................................................................................... 10 Chapter 1 ....................................................................................................... 13 General Introduction ........................................................................................................13 1.1 Intergroup competition ......................................................................................... 13 1.2 Crested macaques ................................................................................................. 17 1.3 Crested macaque intergroup conflict .................................................................... 18 1.4 Intergroup encounter risk perception .................................................................... 19 1.5 Thesis aims and structure ...................................................................................... 20 Chapter 2 ....................................................................................................... 22 General Methods .............................................................................................................22 2.1 Ethical statement ................................................................................................... 22 2.2 Study site .............................................................................................................. 22 2.3 Study subjects ....................................................................................................... 24 2.4 Data collection ..................................................................................................... 24 2.5 Predators ............................................................................................................... 25 2.6 Tourist encounters ................................................................................................ 25 2.7 Behavioural sampling ........................................................................................... 26 2.8 Intergroup encounters ........................................................................................... 30 2.9 Range use .............................................................................................................. 31 2.10 Data processing and analysis .............................................................................. 31 Chapter 3 ....................................................................................................... 38 Intergroup encounter risk in wild crested macaques ( Macaca nigra ): winner/loser effects on spatial cohesion and behavioural synchrony ..................................................38 Abstract ....................................................................................................................... 38 3.1 Introduction ........................................................................................................... 39 3.2 Methods ................................................................................................................ 47 3.3 Results ................................................................................................................... 53 2 3.4 Discussion ............................................................................................................. 61 Chapter 4 ....................................................................................................... 66 Potential intergroup conflict influences current intragroup behaviour in wild crested macaques ( Macaca nigra ) ...............................................................................................66 Abstract ....................................................................................................................... 66 4.1 Introduction ........................................................................................................... 67 4.2 Methods ................................................................................................................ 73 4.3 Results ................................................................................................................... 78 4.4 Discussion ............................................................................................................. 87 Chapter 5 ....................................................................................................... 94 Impacts of intergroup conflict on intragroup social behaviour in wild crested macaques (Macaca nigra ) ................................................................................................................94 Abstract ....................................................................................................................... 94 5.1 Introduction ........................................................................................................... 95 5.2 Methods .............................................................................................................. 102 5.3 Results ................................................................................................................. 110 5.4 Discussion ........................................................................................................... 124 Chapter 6 ..................................................................................................... 131 General Discussion ........................................................................................................131 6.1 Discussion ........................................................................................................... 131 6.2 Limitations and future directions ........................................................................ 137 6.3 General conclusion ............................................................................................. 140 References ................................................................................................... 141 Appendices .................................................................................................. 164 Appendix 1: Predator encounters .............................................................................. 164 Appendix 2: Tourist encounters ................................................................................ 165 Appendix 3: IGEs ..................................................................................................... 167 Appendix 4: Home range & overlap ......................................................................... 169 Appendix 5: Relative IGE risk ................................................................................. 171 3 List of Tables List of Tables Table 2.1 Observation duration, composition and size of crested macaque study groups in TNR, Sulawesi, Indonesia. ............................................................................................ 24 Table 2.2 Definitions of focal sampling behaviours, following Thierry et al. (2000) and Duboscq et al. (2013). ....................................................................................................... 27 Table 2.3 Definitions of scan sampling activities, following Thierry et al. (2000), Duboscq et al. (2013), and Allan & Hill (2018). ............................................................... 29 Table 2.4 Definitions of female sexual swelling states. ................................................... 34 Table 3.1 Hypotheses to test the influence of perceived IGE risk on anxiety, group cohesion, and behavioural synchrony among crested macaques. ..................................... 46 Table 3.2 GLMM (M1) investigating the influence of perceived IGE risk on self- scratching in crested macaques. ........................................................................................ 57 Table 3.3 GLMM (M2): Influence of perceived IGE risk and recent win/loss record on spatial cohesion in crested macaques. ............................................................................... 58 Table 3.4 GLMM (M3): Influence of perceived IGE risk and recent win/loss record on general behavioural synchrony in crested macaques. ....................................................... 59 Table 3.5 GLMM (M4): Influence of perceived IGE risk and recent win/loss record on active behavioural synchrony in crested macaques. .......................................................... 60 Table 3.6 Summary of hypotheses, predictions, and support provided by this study. ..... 61 Table 4.1 Hypotheses to test