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Richarsonmarinera2020phd.Pdf (3.062Mb) EFFECTS OF INTERSPECIFIC INTERACTIONS ON INDIVIDUAL SPECIALISATION Marine Raphaële Amélie Richarson Department of Zoology University of Otago A thesis submitted for the degree of Doctor of Philosophy December 2019 | i “‘Did you know there are certain deep-sea fish that create their own light and feed on lethal volcanic gasses? That's supposed to be impossible. And bats in Australia as large as dogs. And that American coyotes mate for life. And enormous octopi that can squeeze their bodies through holes no larger than a ha'penny coin. Can you imagine that? If only we would stop and look and wonder. And wonder. ‘Do you have a favourite? ‘Not meant to. But mostly the unloved ones. The unvisited ones. The cases that get dusty and ignored. All the broken and shunned creatures. Someone's got to care for them. Who shall it be if not us?” John Logan – The Day Tennyson Died (2016) | ii Abstract In many species, some individuals are specialist users of a subset of the total resources available to their population, a phenomenon known as individual specialisation. This intraspecific variation has long been suspected of shaping population dynamics and has potentially large effects at the ecosystem level. Variation in the degree of individual specialisation can be driven by ecological interactions. The common bully Gobiomorphus cotidianus, a New Zealand native fish, displays a generalised diet at the population level, with moderate levels of individual specialisation. Juvenile perch Perca fluviatilis compete for the same food resources, while larger perch are potential predators. These antagonistic biological interactions may have an impact on the trophic niche and the degree of individual specialisation in bullies. I used surveys of fish communities and cross-sectional diet data to assess individual specialisation in wetland bully populations in relation to ecological factors. Then, in two series of controlled manipulative experiments, I asked how individual specialisation in bullies responded to intraspecific competition, interspecific competition and predation risk from perch. I hypothesised that ecological niches at the population and individual levels would vary according to the composition of the fish community. Finally, I led a collaborative comparative phylogenetic analysis of the literature which compiled indices of individual specialisation in diet across ecosystems and taxa and tested whether individual specialisation was predicted by cumulative measures of human impacts. I anticipated that due to global resource and habitat disruptions, human impacts on individual specialisation would generally be negative. Consistent with the niche variation hypothesis, I found that intraspecific competition led to overall higher intraspecific diet diversity, while interspecific competition led to lower degrees of individual specialisation. Predation risk had | iii a positive effect on dietary, and a negative effect on habitat individual specialisation. However, the direction of these effects changed when interspecific competition occurred along predation risk. At the population level, I recorded niche shifts due to interspecific competition both in habitat use and in diet. Finally, I found negative effects of human activities on individual specialisation in terrestrial and freshwater ecosystems. There was however no evidence of a similar negative relationship in marine ecosystems. This discrepancy in results might be linked to differences in the types of drivers that impact individual specialisation between each ecosystem type. This work underscores the importance of integrating individual variation when evaluating a population’s response to antagonistic biological interactions. | iv Preface & Acknowledgements Individuals matter. This had long been my personal philosophy towards human beings in general. Little did I know that, at the term of four years focusing on the dietary choices of a little fish I had never even heard of, my views on individuality and its importance would encompass, well, every living thing. Individuals matter, because without individuals, there is no community. The flip side of that coin is that there is nothing special about being exceptional. Individuals matter, yes – but the wealth of their differences is nothing when taken in isolation. The value of exceptionalism only appears in the group; it becomes apparent only once individuals start interacting with their congeners, with other species, with their environment. There is no value in exceptionality when it is not expressed within a system where all exceptionalities get a chance to thrive. It is a terribly humbling realisation – or a very reassuring one, depending on where you stand on individualism. And the community shapes individuals. It shackles possibilities beyond body shape and size and abilities and character. That is a terribly depressing realisation – or a very reassuring one, depending on your views on freedom. What I got from it all, however, was a state of quasi-permanent awe about life itself. If all individuals are exceptional, well then, every living being is a miracle in and of itself. Who would have thought that getting more intimate with the ecology of individuals would lead me to gaze upon the world and see little wonders everywhere? *** | v It is difficult to wrap up what a PhD is in a few words. It is a challenge, but only a challenge to yourself. It is world-changing, but the world being changed is your own. It is hard, but only on yourself, and some precious times it is exhilarating – but only to yourself. I did not think I had it in me to keep myself focused on the same topic for four years. And, granted, there were times when I just hated my fish, my bugs, the endless cleaning of the tanks or getting stuck knee-deep – again – in soft, smelly mud. Sometimes I hated it because it was too boring, sometimes I could not believe my luck doing something so exciting. There were highs and lows, a lot of good people, a few great people. There were some struggles and a few victories. There were lessons for the moment, and some lessons for life. During this PhD, I got to understand a little more about the world, and a whole lot more about myself – about what I could handle, about what I could not cope with, about what made me feel whole. It came together because many good people helped me along the way. This project benefitted from the trust and generosity of the New Zealand Te Apārangi through the Marsden fund. Their financial support allowed me to contribute in my humble way to mankind’s ever-growing body of knowledge instead of benefitting from it, for once, and to travel and share my findings with a community of scientists I admire within and beyond New Zealand’s shores. My thanks to the University of Otago and to the Department of Zoology for their logistic and financial support. I feel incredibly privileged to have been welcomed in this beautiful institution. Thank you to my advisory committee, Marc Schallenberg and Robert Poulin, for their support and feedback along the way. I’m also thankful for the many little hands that helped me get through field and lab work: the staff, Nicky McHugh, Kim Garrett, Ken Miller and Nat Lim; and the small army of interns and friends: Yvonne Kalhert, Alex Connolly, Pauline Guilloux, Stef Bennington, Rose-Marie Payet, Gregoire Saboret, Nicky Kerr, Lucian Funnell, Jules Travert, Ludovic Vincent, Raul Costa Pereira, Andrew Veale, and others I might forget… Many thanks to Glen Riley and Te | vi Nohoaka o Tukiauau/Sinclair Wetlands Trust for opening their doors to my helpers and me. Jared and Jane, thanks for your support in my side artistic endeavours; your encouragements meant a lot to me. To the Hot Yoga Dunedin teachers: thank you for keeping me (somewhat) sane in the last stretch (pun intended) of writing. Katherine, Jaz, Matt, Scott, Sven, Lisa – thank you for the chats, the coffees, the support, the advice, the moments of fun. Leida, Pasha, Jenny, Ash, Fasil, you were my little family here. I am missing those of you who aren’t around anymore, and I’m loving those of you who are. To Travis, my patient, supportive, awesome supervisor; you welcomed me in your lab and in your home, and you were the best mentor any student could wish for. Merluches, mes chéries, vous me manquez tous les jours. Ici j’essaie de me rendre heureuse, mais sans votre compagnie c’est un peu plus dur. Rose, je te promets que lors de ton prochain séjour, tu n'auras pas à photographier des poissons ! Et vous, ma famille, amours de ma vie - vous ne comprenez pas toujours pourquoi je fais ce que je fais, mais vous me donnez la liberté de le faire. Cela n’a pas de prix. Now, let’s talk about fishes. | v i i Table of Contents Abstract ........................................................................................................... ii Preface & Acknowledgements ......................................................................... iv Table of Contents .......................................................................................... vii List of tables and figures ................................................................................. x 1 GENERAL INTRODUCTION ................................................................................ 1 Thesis background ........................................................................................... 2 Biological models ............................................................................................ 6 Thesis structure .............................................................................................
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