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University of Bath PHD Conflict in Context: A critique and investigation into the logic, nature, and evolutionary consequences of competition in animals Raihani, Gina Award date: 2007 Awarding institution: University of Bath Link to publication Alternative formats If you require this document in an alternative format, please contact: [email protected] 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: 06. Oct. 2021 Conflict in Context: A critique and investigation into the logic, nature, and evolutionary consequences of competition in animals Gina Raihani A thesis submitted for the degree of Doctor of Philosophy University of Bath Department of Psychology April 2007 COPYRIGHT Attention is drawn to the fact that copyright of this thesis rests with its author. This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with its author and that no quotation from the thesis and no information derived from it may be published without the prior consent of the author. 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Box 1346 Ann Arbor, Ml 48106-1346 C ontents Conflict in Context: A critique and investigation into the logic, nature, and evolutionary consequences of competition in animals:............................................................................. 1 Contents...................................................................................................................................2 Summary................................................................... 3 Acknowledgements................................................................................................................4 Chapter 1. A review and critique o f the study o f competition in animals .........................5 Chapter 2. Aims, objectives and rationale ........................................................................30 Chapter 3. Background review ...........................................................................................40 1. The evolution o f sexual size dimorphism 2. The evolution o f alternative reproductive behaviours 3. Competition, cooperation and the evolution o f female coalitions in matrilineal social formations Chapter 4. The influence o f sexual selection and male agility on sexual size dimorphism in bustards (Otididae) .....................................................................................58 Chapter 5. Male mating tactics in the American ruby spot damselfly: territoriality, non territoriality and switching behaviour ............................................................................... 71 Chapter 6. Competition, cooperation and the dynamic structuring o f social formations in Spotted Hyenas ................................................................................................................. 91 Chapter 7. Discussion and future directions...................................................................132 Bibliography .......................................................................................................................163 Appendix.The influence of sexual selection and male agility on sexual size dimorphism in bustards (Otididae): Published version.......................................................................204 2 Abstract The conventional logic of competition assumes a general ‘struggle’ between organisms for access to food or mates. As a consequence of this ‘struggle’, it is suggested that the most vigorous and competitive individuals usually win out at the expense of their rivals. Despite the clear advantage certain individuals have over others, fitness gradients are maintained in all natural populations. I address this paradox using two complementary approaches. First, I address gender-biased assumptions embedded in traditional mate choice models, and critically examine the extent to which evolution is driven by male- male competition, which is assumed to favour the most vigorous males. I propose that such androcentric bias exposed by feminist critiques is a symptom of a larger problem posed by the conventional neo-Darwinian paradigm. Therefore, I address some of the problems associated with the conventional logic of competition, and propose an alternative perspective, which may help to dissolve the inconsistencies embedded in our current logic. To complement my theoretical critiques, I explored intraspecific competition in three different animal taxa: Birds, insects, and mammals. My first objective was to investigate the effects of reproductive competition on male morphology and mating behaviour in bustards (Otididae). Then, in the American Rubyspot damselfly (Hetaerina americana ), I explored how reproductive competition influences alternative mating behaviours in males. Finally, I used what has been considered an unusual animal, spotted hyenas (Crocuta crocuta), to expose the problem of rigid male-biased approaches to science, and how they have been responsible for mis-representing a number of animal groups including spotted hyenas. Then, I investigated various forms of competition and cooperation in the female dominance hierarchy with a special focus on the relationship between female social rank and the pattern of female coalition formations, and their evolutionary consequences in terms of female reproductive success and group dynamics. Finally, I integrate these studies into a coherent framework, which focuses on the active ways in which animals respond to and influence diverse contexts of competition. 3 Acknowledgements First of all I would like to thank my main supervisor, Professor Helen Haste, for providing me with the opportunity and support to conduct my PhD in the department of Psychology. Helen’s passion for innovative and original works allowed me to explore my research interests using both conventional and non-conventional methods. I would also like to thank Dr. Alan Rayner, who offers a novel perspective, termed inclusionality, which questions the conventional logic of competition by including space as vital source of evolutionary creativity, in which life forms are “pulled together” as a result of their imperfection, rather than “pushed apart” in their strive towards perfection. This helped me to interpret the nature of competition in my study groups from very diverse perspectives. Dr. Tamas Szekely, my undergraduate supervisor, inspired me to work in the field of animal behaviour, owing to his endless passion for the evolution and ecology of birds. Tamas’s support and friendship throughout my postgraduate research has been of great importance from the beginning to the end of my thesis. I am also very grateful towards Dr. Marion East, who kindly offered to co supervise my PhD. With her help I was able to study spotted hyenas in the wild, which would not have been possible without her support. I extend my gratitude to Alejandro Serrano-Meneses, who offered me the opportunity to work with the American rubyspot damselfly, in which I had the freedom to explore the types of behaviours that contributed to my overall research interests. Dr. Alejandro Cordoba-Aguilar also helped and supported my research interests in damselflies. I also extend my gratitude to my family, without whose unconditional support, I would never have managed to complete this challenge. Finally I would like to thank Joanna Bryson and Dr. John Lazarus for agreeing to examine my thesis. 4 Chapter 1:A review and critique o f the study of competition in animals Gina Raihani 5 My childhood dream, my lifelong wish, would have been fulfilled if it had really been possible to converse with animals. Hediger, 1985 1. The logic of competition 1.1 Natural selection The Darwinian and neo-Darwinian synthesis o f natural and sexual selection Can it, then, be thought improbable, seeing that variations useful to man have undoubtedly occurred, that other variations useful in some way to each being in the great and complex battle of life, should sometimes occur in the course of thousands