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Thesis Template The Interactive Effects of Phenotype and Environment on Dispersal by Celina Beatrice Baines A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto © Copyright by Celina Beatrice Baines 2019 The Interactive Effects of Phenotype and Environment on Dispersal Celina Beatrice Baines Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto 2019 Abstract Dispersal is the movement of organisms between habitat patches that creates the potential for gene flow. This movement of individuals and their genes across space impacts the dynamics of metapopulations and metacommunities. For example, dispersal influences the risk of metapopulation extinction, and affects the coexistence of species with competitors, predators, and parasites. Dispersal therefore influences local and regional species richness and community composition. Dispersal is a complex behaviour characterized by high levels of variation between patches and among individuals. The central question of my dissertation is, what produces this variation? I addressed this using three approaches: quantitative review, theoretical modeling, and empirical investigations. In chapter 2, I conducted a meta-analysis of body size-dependent dispersal. I found that size had a positive effect on dispersal distance, but its average effect on emigration and immigration did not differ from zero. However, there were high levels of heterogeneity in this effect within each stage of dispersal. In chapter 3, I developed an individual based model of how dispersal evolves to respond to both body condition and population density. I found that dispersal evolved to be a positive threshold function of density, and the value of this threshold depended on condition. In chapters 4-6, I used the backswimmer, Notonecta undulata, as a model empirical system to explore the effects of phenotype and the environment on ii dispersal. In chapter 4, I conducted a mark-release-recapture study which found that dispersal probability was determined by the interaction between population density, body mass, and sex. In chapter 5, I conducted an experiment which demonstrated that habitat conditions experienced during development have carryover effects on dispersal in later life stages. Finally, in chapter 6, I investigated the effects of parasites on dispersal. I found that risk of parasitism induced dispersal in healthy backswimmers, but infected backswimmers had low dispersal ability. This impacted dispersal in natural environments: parasites eliminated density dependence in backswimmer dispersal. My thesis provides novel information about how dispersal is influenced by phenotype, the environment, and their interaction. These results alter our understanding of the consequences of dispersal for biological processes, including the spatial distribution of biodiversity. iii Acknowledgements There are many, many people who have taught me, helped me, and mentored me during my PhD, and who have generally made my experience more enjoyable. First, I want to thank Shannon McCauley. I first met Shannon as a first-year undergraduate, when I worked as her summer field assistant. Shannon is a wonderful and enthusiastic teacher, and she really sparked a passion in me for ecological research. When I had the opportunity to do a PhD in her lab, I jumped at it, and I’m so grateful I did. Shannon has been a wonderful supervisor; she has guided me through the challenges of ecological research and academia, and supported my intellectual growth and career development. I’m so lucky to have had her as an advisor, and to count her as a close friend. I also want to thank my supervisory committee. Locke Rowe has been my advocate since I started working in his lab as an undergraduate. He has always believed in me more than I believe in myself, and has challenged me to reach for higher research and career goals. I’m so grateful for his support. I want to thank Ben Gilbert for providing helpful and insightful comments and advice throughout my PhD, and for answering my stats questions. Helen Rodd has provided invaluable support both through her role on my supervisory committee, and as graduate chair. I want to thank her for her insightful comments, for sharing the notes she takes during committee meetings, for providing career advice, and for helping me with a million and one things that have smoothed my path through grad school. I have had the great pleasure of doing research at the Koffler Scientific Reserve, and I want to thank everyone there for all they have done to make the reserve run so well. I especially want to thank Stephan Schneider who has gone above and beyond to help me conduct experiments, has built the crazy things I need for research (including flight tunnels and ponds), and has made the reserve an efficient, fun, and safe place to do research. Steph has also rooted for me every step of the way, and I’m so grateful he’s my friend. I also want to thank John Stinchcombe, Jenn English, John Jensen, Tobias Mankis, and Olivera Joksimovic for logistical (and sometimes physically strenuous) support at the reserve. There are many people to thank for keeping the department running smoothly, and making my life easier in countless ways. At UTM, I want to thank Carolyn Moon, Stephanie do Rego, Amy Yeung, and Bob Liange. And in the EEB department, I want to thank Kitty Lam, Olivera Joksimovic, and again Helen Rodd. iv Many people provided help in the lab and field during my PhD. Ilia Ferzoco has been an amazing research partner. She has made invaluable contributions with her diligent observations on the natural history of backswimmers, and she has spent countless hours helping me in the field. Much of the work in my thesis could not have been possible without her. I also want to thank Shantel Catania, Ariana Longley, David Soliman, Quan Le, Tharusha Wijewardena, Sarah Hasni, Rokhsar Rezaee, Arjan Banerjee, Julienne Bonoan, Racquelle Mangahas, Samantha Hasbum, Bansari Patel, Rosemary Martin, Dachin Frances, Hugh Traquair, and Betty Dondertman for their help in the field and the lab. I also want to thank my collaborators on the theoretical dispersal modeling project, Justin Travis and Greta Bocedi, for being fantastic teachers. I was lucky to have amazing labmates to share the grad school experience with. Huge thanks to Ilia Ferzoco, Rosemary Martin, Dachin Frances, Sarah French, Rosalind Murray, and Chris Searcy. I can’t imagine better people to work with, laugh with, and complain to. Special thanks to Ilia for being my number one pep-talker and confidence booster, I’ve appreciated the support so much. Shout outs to Ros, for making me laugh every day and being my Toronto soul-sister, and to Rosie for making everything fun, and making every day more beautiful with her insect- themed art. I want to thank my family for all of their love, patience, and support. They have always believed I could do anything, and supported me even when I chose a career collecting bugs. My parents Betty Dondertman and Frank Baines, and my sister Nicky Baines, have encouraged and helped me every step of the way. I’m so grateful for my mom, who encouraged my love of nature and animals, and who even drove out to help me in the field at the end of a season when I was exhausted and needed extra moral support. Finally, I want to thank my partner Hugh Traquair. Hugh has listened to me talk endlessly about bugs, helped me work through problems, and helped me in the field many times, including once during a snow storm. Hugh has provided boundless moral and emotional support through all the highs and lows of research and academia. Life is better for sharing it with you. v Table of Contents Acknowledgements ........................................................................................................................ iv Table of Contents ........................................................................................................................... vi List of Tables ...................................................................................................................................x List of Figures ................................................................................................................................ xi List of Appendices ....................................................................................................................... xiii Chapter 1 General Introduction .......................................................................................................1 1.1 Overview ..............................................................................................................................1 1.2 What is dispersal and how do we study it? ..........................................................................1 1.3 Causes of dispersal ...............................................................................................................4 1.3.1 Ultimate causes of dispersal.....................................................................................4 1.3.2 Proximate causes of dispersal ..................................................................................7 1.4 Consequences of dispersal ...................................................................................................9 1.4.1 Consequences of dispersal for ecological dynamics ................................................9 1.4.2 Consequences of dispersal for evolutionary
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