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Cooperation, Conflict and Warfare in Wild Banded Mongooses

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Cooperation, conflict and warfare in wild banded mongooses Submitted by Elizabeth Fiona Rosemary Preston to the University of Exeter as a thesis for the degree of Doctor of Philosophy in Biological Sciences September, 2019 This thesis is available for Library use on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. I certify that all material in this thesis which is not my own work has been identified and that any material that has previously been submitted and approved for the award of a degree by this or any other University has been acknowledged. (Signature) ……………………………………………………………………………… 1 2 SUMMARY Intergroup conflict can be a strong force in the lives of social species. Conflict can be dramatic, leading to serious injury or death, loss of territory or dominance status, and can impact behaviour, reproductive success and fitness. The impact of intergroup conflict on within-group behaviour is a growing area of research, and evidence for increased affiliation between group members after exposure to intergroup conflict has been found in several species. However, these studies focus on short timescales, the minutes and hours post-conflict, and it is unclear what effect intergroup conflict has on within-group behaviour in the longer term. In this thesis I use the banded mongoose (Mungos mungo) as a model system to investigate the effects of intergroup conflict on within-group behaviour in the longer term. I discovered that group level within-group affiliation was only affected in the hour after exposure to conflict, but individual social relationships were affected into the longer term, up to two days after exposure. Unlike other studied populations, banded mongooses reduced within-group affiliation and aggression, and these changes differed between males and females, and between younger and older mongooses. I found only tentative evidence that intergroup conflict affected group movement or home range use, however, the risk of intergroup conflict affected leadership, with evidence that females lead more successfully in areas of high risk at the edge of the territory, which may indirectly affect movements in the longer term. This thesis gives evidence that intergroup conflict affects behaviour in the longer term, beginning to bridge the gap between evolutionary theory and empirical observations, and highlights that groups do not respond in a heterogeneous way, as different sex and age classes react differently, potentially due to differential costs and benefits. 3 ACKNOWLEDGEMENTS Firstly I would like to thank my supervisors, Mike Cant and Darren Croft, for all of their valuable feedback and guidance throughout my PhD. Their knowledge, patience and enthusiasm for evolutionary and behavioural ecology is a great source of inspiration. Special thanks to Mike for guiding me through my scientific journey and introducing me to the great banded mongoose system. Thanks also to all the members of the KillerMongWasp team, throughout my time working with the group. Extra thanks to Harry Marshall who introduced me to the field site and Ugandan ways of living, and to Faye Thompson for her advice and guidance both in the field and statistically! I would also like to thank my collaborators and co-authors for their help and insight, especially Sam Ellis for his social network know-how. None of this research would have been possible without the help of Francis Mwanguhya, Solomon Kyabulima, Kenneth Mwesige, Robert Businge and Solomon Ahabyona. Thanks to you all for teaching me all about the mongooses, keeping me safe in the field, and telling me all your wild tales of animal encounters! Special thanks to Solomon Kyabulima for your diligence, hard work and incredible mongoose identification skills; without you these experiments would never have worked. Thanks to everyone I met at Mweya during my field work. Thanks for making Mweya a beautiful and friendly place to work, and for throwing some good parties Ugandan style! Thanks to everyone in the Centre for Ecology and Conservation for making it a friendly and inclusive place to work. Thanks especially to my first set of housemates Guill McIvor, Harry Marshall, Sarah Paul, Honey and Louis for 4 welcoming me to Cornwall and making my transition so much easier and more fun! Thanks to all my office mates and lunch pals for science and not-so-science discussions, lunch times would have been much less eventful without you! Thanks to all my Falmouth friends for making Cornish life fun, and always making time for a pint in the Front. Thanks also to my other friends scattered across the country who visited Cornwall, or provided support from afar. Thanks for listening to me chat about mongooses. Thanks to my family for being an unending support. Thanks to my Mum and Dad for supporting me through all of my studies, despite them taking me so far away. I’m so glad that you could come and experience Uganda and finally go on safari. Thanks to my brother Alex for family times, putting me up on my way to conferences, and providing a different perspective. Thanks to my animal family – Tobie and Jim – you made writing up so much easier and provided much needed distraction and cuddles; and Spike and Willow for providing the same in Rutland. Finally, thanks to Henry. You’ve always been so supportive, even when I moved you down to Cornwall with me and promptly left for Uganda! Thanks for offering to proof read my thesis even when it was the last thing you wanted to do. Thanks for looking after me and reminding me that I could do it, especially when I didn’t think I could. I’m really glad you could see Uganda too, and understand why I love field work so much. I could not have done this without you. 5 LIST OF CONTENTS Summary........ ........................................................................................................... 3 Acknowledgements .................................................................................................. 4 List of Contents ........................................................................................................ 6 List of Figures ........................................................................................................ 10 List of Tables .......................................................................................................... 12 Author’s Declaration .............................................................................................. 18 Publications.. .......................................................................................................... 19 Chapter 1: General Introduction ....................................................................... 21 Intergroup conflict and multi-level selection .......................................................... 22 Intergroup conflict in animals ................................................................................ 29 Who participates in intergroup conflict? ............................................................ 33 When and where do intergroup conflicts take place? ........................................ 35 What are the consequences of intergroup conflict? .......................................... 37 Study system: banded mongoose......................................................................... 46 Cooperative breeding ........................................................................................ 48 Intergroup conflict .............................................................................................. 49 Study site and methods ..................................................................................... 51 Aims and outline of the thesis ............................................................................... 53 6 Chapter 2: Effects of simulated intergroup conflict on collective behaviour do not persist into the long-term in banded mongooses ................................... 57 Abstract ................................................................................................................ 59 Introduction ........................................................................................................... 60 Methods ................................................................................................................ 64 Study Site .......................................................................................................... 64 Data Collection .................................................................................................. 65 Statistical Analysis ............................................................................................ 73 Results .................................................................................................................. 75 Immediate reactions .......................................................................................... 75 Behavioural responses ...................................................................................... 76 Discussion ............................................................................................................ 80 Chapter 3: Network-level consequences of intergroup conflict in a cooperative mammal society: redirection of grooming and aggression between sexes......................................................................................................................... 87 Abstract ...............................................................................................................
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