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Consequences of within- and between-group conflict in dwarf mongooses Amy Morris-Drake A dissertation submitted to the University of Bristol in accordance with the requirements for award of the degree of Doctor of Philosophy in the Faculty of Life Sciences January 2021 School of Biological Sciences Word count: 48,443 Summary Conflict is rife in the natural world and is widely recognised as a powerful selective pressure in social evolution. For group-living species, conflict over valuable resources arises between members of the same group (within-group conflict) and with conspecific outsiders (between-group conflict; also called outgroup or intergroup conflict). Traditionally, research on nonhuman animals has focused on the contests that arise within- and between groups; in the former case, there have also been extensive studies of the immediate consequences of aggressive interactions. By contrast, relatively little is known about the short-term consequences of between-group conflict and, for both conflict types, there has been little empirical consideration of the longer-term consequences. In this thesis, I combine detailed behavioural observations, field experiments and long-term life-history data to investigate the consequences of within- and between-group conflict over different timeframes, using wild dwarf mongooses (Helogale parvula) as a model system. First, I show that unresolved within-group conflict can have cumulative behavioural effects and present evidence for delayed post-contest management by bystanders (Chapter Two). In the remaining data chapters, I focus on between-group conflict. I find consequences of outgroup threats in the short-term (in the hour after exposure) for within-group affiliation, foraging and sentinel behaviour (Chapter Three). I then demonstrate that such threats can also influence behaviour over longer timeframes, with carryover effects on affiliation into the following day, and cumulative effects on affiliation, foraging, sentinel and territorial behaviour, as well as body mass, after repeated outgroup encounters across a week. Lastly, I examine the reproductive consequences of between-group conflict and, unexpectedly, find a positive relationship with pup survival. By demonstrating an array of effects over timeframes rarely addressed previously, I help shed light on the more-lasting consequences of social conflict which is important for a greater understanding of how it shapes the lives of group-living species. 1 2 In loving memory of Peter For a life full of adventure 3 4 Acknowledgements From the UK to South Africa, I have been lucky to work with so many incredibly generous and inspirational people. Thank you to everyone for making the last four years an overwhelmingly positive experience. There is one person in particular, to whom I would like to extend the most gratitude for his unceasing support throughout this whole process, Andy Radford. On countless occasions over the past four years his kindness has awed me and his intellect flawed me. I always come away from meetings with Andy feeling a lot clearer about the direction I’m headed, and with a renewed sense of optimism. He is always on hand to offer his expertise and moral support, whether I'm just down the corridor in the department or 10,000 miles away in the field. One of Andy’s many talents is assembling teams of people who share a sense of humour, but also a genuine academic curiosity, ambition and respect for science. In many ways I have him to thank for introducing me to a number of the following people whom I also wish to thank. Julie Kern established the Dwarf Mongoose Research Project (DMRP). For that feat alone I will have eternal respect for her. She has been a great friend and collaborator, and although she stopped working directly with the project in 2017, she has always been there for me when I have had questions, and there have been many! This probably goes some way to explaining why our voice notes are rarely shorter than 10 minutes, sorry about that. Something I have always appreciated from Julie is feeling able to vent during challenging times, and because she understands the work better than anyone, I never have to explain myself. I’m so happy that we managed to coincide our field seasons at the beginning of the PhD. Having Julie and Garth down the road on future trips was also such a treat – thank you for the many catch-ups over G&Ts and large glasses of vino! The same can be said about Charlotte Christensen. Charlotte and I spent our very first field season on the project together back in 2013; an experience that cemented a life-long friendship and a new-found love for fieldwork in Africa. Charlotte is one of those friends who completely understands what I do and why, with a healthy mongoose obsession herself! I feel very lucky to have had her support throughout the PhD process. Charlotte was also the original dwarfie “poo-ninja”, so I also have her to thank for introducing me to that less glamourous side of fieldwork. None of this research would have been possible without Hennie, Eddie and Darnie Yeates. I would like to thank them for allowing us to work on their beautiful reserve. I would also like to extend my thanks to the friendly staff I overlapped with whilst working on Sorabi - Christoffel, Suzelle, James, Justice, Lindiwe and Petunia. From honey badgers raiding our bins and mysterious elephant break ins, to leopard kills, zebra meat collection and numerous flat tyres, we all experienced this rollercoaster together. I would like to extend a huge thank you to the following people whom I overlapped with during three field seasons on the DMRP: Katie Collier, Emma Eastcott, Harry Suter, Alanah Lewis, Tecla Mohr, Izzy Jarrold, Sophie Manson, Jenny Linden, Josh Arbon, Emily Richens, Aidan Byrne, Hannah Rickets, Frazer Bayliss, Emily Grout, Alex Fergusen, Izzy Shan, Natalie Tegtman, Izzy Carpenter and Megan Layton. Thank you for always smiling, despite the lack of sleep, sweaty work and seemingly constant smell of egg! Special thanks go to Jenny Linden, without whom I may still be running the experiment in Chapter Four, to Natalie Tegtman for keeping me sane when it felt like everything was spiralling against me during my final field season, and to Megan Layton for managing the project amazingly through a global pandemic. Megan has also helped with project admin during the final month of writing up, for which I am extremely grateful. 5 I would also like to thank my surrogate family in South Africa for reminding me of the virtues of a good work/life balance. I have many fond memories of braiis and game drives with the following people, and am fully converted to celebrating Christmas in July! Thank you Katie, Andrew, Garth, Robbie, Rusty, Ed, Anka, Andre, Jen, Phil, Ann and Neil. At this point it would be remiss of me not to thank the mongooses! Their quirky characters were a constant source of entertainment. A special thank you to Bean, and the fine supporting cast of Sorabi’s rich wildlife, particularly Big boy, Elvis, Delilah and Gemima - a few of the resident ungulates. Back in the UK I feel privileged to have been part of an inspiring group of people from the Universities of Bristol and Exeter, carrying out research spanning dwarf mongooses, fish and wasps. In 2020 the value of having this cohesive group of like-minded individuals to call on for support has been particularly apparent. Regular research group meetings on Zoom, virtual writing retreats and quizzes have all helped to keep morale up during this challenging time. There are a few people in particular whom I would like to thank for the help they have offered me: Patrick Kennedy is one of the smartest people I have ever met, and when it comes to Excel he appears to have a shortcut for absolutely everything. This has saved me an enormous amount of time. Ines Goncalves has also been a constant source of both technical and emotional support to me. I have an incredible amount of respect and gratitude for them both, as do I for Harry Harding. I look back very fondly on our daily coffee catch ups. One the biggest challenges this year has been keeping calm whilst writing up throughout Covid-19. To this end I would like to thank my yoga instructor, Cae, for her regular sessions on Zoom.