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Durham E-Theses Durham E-Theses Behavioural mechanisms of conict and conict reduction in a wild breeding polygynous pinniped BISHOP, AMANDA,MARIE How to cite: BISHOP, AMANDA,MARIE (2015) Behavioural mechanisms of conict and conict reduction in a wild breeding polygynous pinniped , Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/11146/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk 2 Behavioural mechanisms of conflict and conflict reduction in a wild breeding polygynous pinniped By Amanda M. Bishop School of Biological and Biomedical Sciences Durham University 2015 Submitted for the degree of Doctor of Philosophy Abstract Aggressive interactions arise as a means of resolving access to resources such as food, habitat or mates, but these interactions are often costly in time, energy, or physical damage. Therefore, natural selection favours social systems, spatial organisations and behavioural mechanisms which can balance the trade-offs between conflict and increasing fitness. The diversity of behaviours associated with animal conflict has been investigated in a variety of species; however, rarely are these investigations done in wild systems or with consideration of geographic and intra-seasonal variation in environmental factors, resource availability or social dynamics. Therefore, the aim of this thesis was to examine the behavioural mechanisms underpinning population and individual conflict and conflict reduction at a variety of temporal and spatial scales. I focused my investigation on a particular form of animal contest, male-male breeding aggression, and used the colonial, wild grey seal (Halichoerus grypus) as my model. Specifically, I studied males at Donna Nook, an atypical breeding colony, in order to: (1) update our current knowledge of grey seal breeding systems by comparing the behavioural patterns at recently expanding mainland colony to previously examined, offshore colonies, (2) investigate the information present in a geographically isolated behaviour used in male grey seal conflict, (3) identify the variation in assessment strategies used by individuals in grey seal conflict and conflict reduction, (4) examine how environmental, anthropogenic, and social environments shape individual variation in aggression, and (5) assess the individual variation in decision-making processes such as mating strategies within- and across seasons. Draws were identified as a common conflict outcome for male grey seals, and the acceptance of draws represented a behavioural mechanism which can promote conflict reduction. Assessment strategies relied on individual energetics when costs of conflict were high, but mutual assessment was used in low-cost contexts. Activity budgets were relatively conserved across colonies, and social stability played a key role in mediating conflict. These findings all demonstrate the trade-offs between conserving energy for reproductive activities and expending energy to ensure exploitation of resources. Methodological approaches which accounted for variation in individual partitioning of aggression and reproductive effort within- and between-seasons revealed that the relative importance of dominance as a driver of conflict, the use of specific aggressive behaviours, and broad assessment and mating strategies were all context-dependent at a variety of temporal and spatial scales. Overall, these findings have provided new insights into the evolution of conflict and conflict reduction within polygynous mating systems. This work highlights the importance of incorporating the natural environmental variation and social dynamics into models of individual behaviours. Such approaches not only reveal the plasticity or consistency in how individuals deal with tradeoffs, but they also allow for observing the importance of behavioural mechanisms such as draws, which might have been ‘artificially selected out’ in controlled, laboratory settings. Finally, by using the grey seal breeding system as a model, this work has contributed to our knowledge of this species’ behavioural repertoire, and the role of topography in the evolution of polygyny and aggression in pinnipeds. i Declaration The material contained within this thesis has not previously been submitted for a degree at Durham University or any other university. The research reported here has been conducted by the author unless stated otherwise. © The copyright of this thesis rests with the author. No quotation from it should be published without their prior written consent and information derived from it should be acknowledged. ii Acknowledgements The research presented in this thesis was funded by the Durham Doctoral Studentship, the Peter Evans Trust, and a NERC SEIS-UK equipment loan (#928). It was coordinated by Durham University and the Sea Mammal Research Unit at St. Andrews University. I am grateful to the Association for the Study of Animal Behaviour (ASAB), Ustinov College and the Society for Marine Mammalogy, all of whom provided travel grants that allowed me to present some of this research at various international and local conferences. Thank you to the Lincolnshire Wildlife Trust, particularly Rob Lidstone-Scott and Lizzie Lemon for all your logistic support during fieldwork, for providing local knowledge and for the occasional cuppa tea to warm me up in the hide. Also thanks to the RAF Donna Nook team, particularly Craig and Linda, again for support and logistics. For fieldwork assistance, thank you to Paul Denton and the SEIS-UK team for helping a behavioural scientist quickly get up to speed on geology and seismometers, and for your continued support in-field and during the write-up process. Special thanks also go to James Stewart for your two years of dedicated field assistance. I have no idea why after one year of freezing in a little wooden box (and I mean the one in Mablethorpe) you wanted to subject yourself to it again, but I can’t thank you enough for helping gather valuable data over the years! Thank you to Paddy Pomeroy for your continual support, comments on papers, advice, and affinity for thinking of snazzy titles for papers and posters. I will never forget that ‘the third time is not the charm” when it comes to paper reviews; the “third time is the one that comes before the 4th time”. Thank you to the Isle of May 2014 team, Holly, Matt, Veggie-Matt, Hayley, Joe, Simon and Kimberly: whether you knew it or not, our dinner table conversations, discussions while walking up Palp Brae, or just the general sharing of our respective projects gave me a greater insight into my own work and the bigger pictures of ecology, physiology and evolution at a time when I was struggling to tie up all the strings of thought. To the Durham crew: first, thank you to everyone who has wandered through the doors of the ‘seal lab’ over the years: Charlotte, James, Hani, Sam, Hagen, Amy and Naomi. Through coffee breaks, and being conference buddies, it was a pleasure getting to know and work with you. Especially thank you to Ross Culloch, for showing me the ropes as a new PhD student, for writing my first loop in R, for just chatting over a pint, and for giving me advice on what committees not to join (even if I didn’t listen to you). I’d also like to thank the rest of the Adaptation and Ecology (BEER) group: Dr. Phil Stevens, Dr. Shane Richards, Dr. Martyn Lucas, Dr. Steve Willis and Dr. David Baker for general support over the course of my PhD in statistics, teaching or academic life. To all the masters and PhD students who also came and went over the years, of which there are too many to name: thank you for helping me keep a healthy work-life balance, providing copious amounts of cake, for general advice and support, and for proving that misery really does love company when it comes to statistics. iii To my family, thank you for your continued support regardless of how far away from home I might have wandered this time. (A side thank you to the makers of Skype). Mom and Dad, thank you for instilling in me an appreciation for nature and natural sciences at an early age through camping trips and memberships to zoos and museums. Thank you for always supporting me in the pursuit of my goals. To my siblings, thanks for nodding along patiently when I lapse into talking about seals, and for making time to play Mario kart whenever I did make it home. Thank you to the flight attendants and airline crews who made the constant travelling back and forth from the US to the UK a less horrid experience. And finally, thank you to my supervisor, Dr. Sean Twiss: for taking on a PhD student for a behavioural study with very little prior behavioural experience; for your patient and supportive guidance; for showing me the ropes that first field season; and for constantly helping me in attaining opportunities to further my career through academic endeavours, networking opportunities, teaching experience, and additional fieldwork opportunities. Thank you for helping me take advantage of the opportunity to travel to the Biennial Society of Marine Mammalogy conference in NZ, and for fixing the flight details when it all when wobbly last minute! Thank you for supporting our cause for PhD students to get more teaching experience.
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