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Historical Contingencies in the Ecology and Evolution of Species Diversity

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Historical Contingencies in the Ecology and Evolution of Species Diversity Historical Contingencies in the Ecology and Evolution of Species Diversity by Rachel M. Germain 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 Rachel M. Germain 2016 Historical contingencies in the ecology and evolution of species diversity Rachel M. Germain Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto 2016 Abstract Ecologists have long-sought to explain the high diversity of species in biological communities, given that classic theory predicts that diversity is limited by available niche space. In recent years, ecologists have looked towards ‘historical contingencies’, the persistent effects of past ecological and evolutionary processes, as possible mechanisms that maintain diverse communities, either by relaxing the constraints of niche availability or by adding temporal dimensions to species’ niches. In this thesis, I use field and greenhouse experiments to explore three ways in which historical contingencies manifest in annual plant communities. First, my work on maternal effects shows that abiotic (ch. 2) and biotic (ch.3) conditions in the maternal generation have diverse effects on offspring phenotypes across an assemblage of species. Because species differences in environmental responses can facilitate coexistence, these studies suggest that maternal effects could act as a form of niche differentiation, and motivate future research to clarify their influences on coexistence outcomes. Second, I performed, to my knowledge, the first experimental decoupling of dispersal limitation and environmental sorting in ii a natural landscape by manipulating entire seed pools of annual plants (ch. 4). In doing so, I was able to identify the pervasive and scale-specific influences of dispersal limitation that constrain species distributions in plant communities. Lastly, I used competitive trials to identify macroevolutionary divergence in competitive interactions among species (ch. 5), and found evidence that divergence is contingent on historical competitive interactions in ways that are consistent with character displacement. In sum, my dissertation work has expanded our understanding of (i) the number of potential niche dimensions that might allow species to differentiate, (ii) how this differentiation can arise over evolutionary time, and (iii) the interplay of current and historical conditions in the maintenance of species diversity, and the timescales over which they play out. iii Acknowledgments I have a long list of friends and colleagues to thank who have helped me complete my PhD and keep my sanity in the process. First and foremost, I am indebted to my advisor Benjamin Gilbert. It is rare to find an advisor who is the combination of a brilliant mentor, an enthusiast of Ben Stiller movies c. 2000, and willing to shovel (and re-shovel)10 tonnes of soil off of a university driveway when his student makes poor life choices. Choosing a PhD advisor is one of the most important and most difficult decisions that a young scientist can make, and I truly believe that working with Ben was one of the best decisions I have made for myself. He has forever altered the way that I think as a scientist and my perspective of academia, and it has been exciting to watch his lab develop and successes accumulate over the past five years. I will deeply miss my talented lab family: Natalie Jones, Tess Grainger, Denon Start, and Kelly Carscadden. Natalie is the sister I never had; she has sat ~4ft from me at all times during the last five years, and I am glad to have shared many of life’s big moments with her. Her successes have led her to more xeric pastures living the dream at UC San Diego. Tess has taught me a lot about life and how to live it (buying eggs on a Tuesday); I am going to miss our amped up science discussions and mutual desire to derail book clubs. Denon is someone that I am both inspired and annoyed by, but I will admit that that his projects are pretty rad (he’s too young to know what that means). Kelly is pun-master extraordinaire; I admire her attention to detail, command of the literature, and quirky sense of humor. Each of these folks will go on to do amazing things in life, and I look forward to being their colleague in the future. In addition to the members of my lab, I have made a number of great friends who have offered comradery, advice, feedback, support, and commiseration in different ways throughout my time here. This includes, in no particular order, Alex DeSerrano, Adam Cembrowski, Franus Hauser, Jenn Coughlan, Rebecca Batstone, David "Jello" Timerman, Michael Foisy ♥, Mitch Daniel, Luke Rogers, Thomson-lab Paul, Kyle Turner, Stephen DeLisle, not Luke Frishkoff (I haven’t known him long enough), and many others. A special shout out to Alex DeSerrano, who I first met in 2011 while admiring her use of fish cartoons as a font on a research poster. Grad school is full of ups and downs but it is great friends and fun that keep things moving. iv The Department of Ecology & Evolutionary Biology at the University of Toronto is truly exceptional, and I am glad to have had the opportunity to carry out my graduate work here. I am grateful for the time and insight of past and present members of my thesis committee, Jason Weir, John Stinchcombe, and Marc Cadotte. Helen Rodd, Don Jackson, and Locke Rowe are champions in the improvement of graduate student life in their positions within EEB and SGS. I also thank my office neighbor and close friend of Ke$ha, Donald L. Mahler, for his tolerance and for including me in his lab meetings. There are many more faculty and graduate students to thank who have contributed to my intellectual and interpersonal development. There are a number of folks at UC Davis and the University of Toronto who have helped me in the field, laboratory, greenhouse, or administratively. From my time in California, I thank Sharon Strauss and Susan Harrison for introducing me to a new study system, as well as their lab members for project feedback and plant identification. At the University of Toronto, I have been helped by Kitty Lam, Bruce Hall, Andrew Petrie, Bill Cole, Jim Dix, Jill Wheeler, Jenn English. There is also a small army of undergraduate assistants to thank, most notably Chris Blackford, Alanna Leale, Abdul Yossof, Yvonne Chan, and Alicia Chou. I am grateful for my early mentors at the University of Guelph’s Department of Integrative Biology, Andrew MacDougall, Chris Caruso, Hafiz Maherali, and Mark Sherrard, for their dedication to undergraduate research training. I am particularly thankful to Chris and Hafiz, who welcomed me into their labs despite my lackluster grades and helped me develop the skills to be successful in graduate school, and to Andrew MacDougall, whose laidback enthusiasm for community ecology and field research compelled me to pursue it in graduate school. I would not be where I am today without having worked under their guidance at a critical juncture in life. Last but not least, I could not have completed this work without the support of my mum and brother, Marie Germain and Adam Germain, as well as my partner, Alex Kushnir. Alex has put up with me through late nights, bad moods, weekends in the greenhouse, “insufficient funds”, long fieldwork absences, etc. As someone outside of academia, his perspective of how bizarre the day-to-day of ecology and graduate school is (“daycare for adults”) reminds me not to take myself too seriously and to enjoy the ride. It has been fun exploring Toronto together, and I look forward to new adventures on the west coast with our three cats, who are to blame for any typographical errors in this thesis. v This research was funded by NSERC, Sigma Xi, the Botanical Society of America, and fellowships provided by the University of Toronto and the Department of Ecology and Evolutionary Biology. vi Table of Contents Acknowledgments........................................................................................................................... ii Table of Contents .......................................................................................................................... vii List of Figures ..................................................................................................................................x List of Appendices ....................................................................................................................... xiv Chapter 1 General Introduction .......................................................................................................1 Conceptual background ...............................................................................................................1 Thesis overview ..........................................................................................................................3 Maternal effects of past environments .................................................................................4 Dispersal histories in a metacommunity ..............................................................................5 Historical interactions and macroevolution .........................................................................6 Synthesis of research objectives ..........................................................................................7 Literature cited ............................................................................................................................8
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