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Energy Flow and Food Web Ecology Along a Hydroperiod Gradient

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Energy Flow and Food Web Ecology Along a Hydroperiod Gradient Energy Flow and Food Web Ecology Along a Hydroperiod Gradient by Tiffany A. Schriever A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Ecology and Evolutionary Biology University of Toronto © Copyright by Tiffany A. Schriever 2012 Energy Flow and Food Web Ecology Along a Hydroperiod Gradient Tiffany A. Schriever Doctor of Philosophy Ecology and Evolutionary Biology University of Toronto 2012 Abstract Identifying the ecological mechanisms that determine food web structure is critical for understanding the causes and consequences of diversity. The objective of this thesis was to identify the mechanisms structuring aquatic food webs across environmental gradients from a multi-level perspective (individual to ecosystem) using integrative methodology and field experiments to test classic ecological theory. My results demonstrate support for the dynamic constraints hypothesis, which predicts habitats with greater disturbance should have shorter food chains, but are not consistent with the ecosystem size hypothesis that predicts larger ecosystems have longer food chains. Insect and amphibian richness increased with increasing pond size and hydroperiod, indicating that insertion of new consumers into pond communities was driving variation in food-chain length. A multivariate analysis testing the influence of physicochemical variables on food-web characteristics revealed that hydroperiod and pond area had a strong influence on amphibian and invertebrate assemblages, trophic diversity and 15N range. Food- chain length did not respond strongly to any one variable, but instead responded weakly to multiple environmental variables, suggesting interacting influences on food-web structure. Conversely, the trophic niche of amphibian larvae was not influenced by pond hydroperiod, but did exhibit ontogenetic diet shifts. Populations of amphibian larvae with broader niche widths ii also had increased individual variation, supporting the niche variation hypothesis. In addition, I assessed whether species diversity influenced the magnitude of cross-habitat resource flow between aquatic and terrestrial habitats via emerging aquatic insects, metamorphosing amphibians, and litter deposition. Deposition into ponds far exceeded carbon exported via insect and amphibian emergences. We found a negative relationship between resource flux and the diversity of amphibians and insects, which contradicts the general pattern of positive biodiversity-ecosystem function relationships. My research strongly suggests environmental variation is a key process in shaping food-web structure and function and that multiple methodologies are needed to understand temporal and spatial dynamics of aquatic ecosystems. iii Acknowledgments My first and biggest thank you goes to Devin Bloom. I can’t possibly express in words how grateful I am for all his love, support, and pep-talks. I surely would not have kept any sliver of sanity or had any fun without him around. He has been my greatest supporter, best friend, and editor for too many years to count. I wish I was half as great of a scientist as he is. Thank you and I love you! I owe an enormous thank you to my advisor, Prof. Dudley Williams. He took a chance and gave me the opportunity to become part of his legacy. Dudley gave me the freedom to develop as an aquatic biologist through his guidance and financial support through the years. I am grateful for the quick turnaround and thorough edits of manuscripts and positive feedback. I owe him a big thank you because he is responsible for my newfound love of invertebrates. I am saddened that I am one of the last members of the Williams’ lab, but humbled to have been part of it. I am thankful for my PhD advisory committee, Rudy Boonstra, Bob Murphy, and Marc Cadotte. I appreciate all their helpful comments and discussion throughout the different stages of my program. A big muddy, pond-smelling hug for all those who labored in the field with me: Jennifer Arblaster, Katherine Bannar-Martin, Devin Bloom, Shauna Bloom, Kristen Brochu, Monica Candelaria, Kirsten Comberford, Mark Conboy, Jake Cowper Szamosi, Klara Jaspers-Fayer, Maria Modanu, Stephen Pynn, David Stitt, Caroline Tucker, and Tristan Willis. I am thankful for Fiona Munro who walked around the ponds numerous times with a heavy Trimble GPS unit and gave me my first introduction to GIS. I loved working in the ponds all day with such wonderful company! Caroline, I’m sorry for the nights lacking sleep and bug bites. Kirsten thanks for pointing out all the spiders even though you were not thrilled to see them – everywhere! Kat, Maria, Kristen, Pynn and Devin, you are all workhorses with strong backs and big smiles. Thank you again and again! I could not have processed as many samples, sorted as many invertebrates, or dried and ground as many amphibians as I did without the amazing help of: Sarah Booth, Siao Ryan Yang, Ruby Sambi, Gaayathiri Jegatheeswaran, Maryam Mahmood, Gagan Gill, David Stitt, Katie Keogh, Kristen Beck, and Christen Dschankilic. Some of these students stuck with me for multiple years iv and even after graduating! I wish you all the best of luck in your academic and life endeavors. Thank you so much for all your countless hours of unpaid work. Sarah and Jenn you are amazing, intelligent, and creative biologists. I can’t wait to see what you bring to the field. Thank you for letting me practice playing the role of advisor on you! I want to extend a special thanks to Marc Cadotte and his lab members, Caroline, Lanna, Nick, Kelly, and Robin, for welcoming me into their joyful lab. I enjoyed the meetings, get-togethers, birthday cakes and the candy that was always available. Marc you have been kind, extremely helpful, and a very important mentor to me. I thank Brian Crother, my master’s supervisor, who continually provides spirited support and encouragement. Nate Lovejoy, thank you for writing a reference letter for my new post-doctoral position. I am very grateful for the Toronto Zoo staff and Malcolm Campbell for providing an enjoyable and enlightening way to spend my teaching hours. My time at UTSC was made tremendously better by the friendship of Emily, Maria, Kristen, and Megan. My lab mates: Catherine, Kevin, Julie, Sarah, Jessica, Oksana, and Judith provided guidance, support, and good times. I will miss the walks in the valley, lunch on the patio, and your smiles. I have enjoyed my time in Toronto, although I have a love-hate relationship with Toronto. Toronto has been a fun city to have called home for five years, but I will be happy to leave the traffic behind. I love the friends I’ve met, of whom I will miss greatly. My newly acquired family of Newfoundlanders: Dave, Steve, Kirsten, and Danielle (yes, you are guilty by association) were always there to provide awesome food, wine, laughs, and encouragement. Karen and Hernán I am blessed to have you as friends. Until we all meet again! Lastly, I am very grateful for a family that has provided support and encouragement through the many tiring years of graduate work. Mom and Dad, South Dakota will always be my home regardless of where I live. I am grateful for the Blooms as they have always supported me, pushed me to be confident, and provided entertainment. My heart is continuously filled with love from my family and for that I thank you all for giving me the strength to pursue my dreams. If I forgot someone, please forgive me. So many people have helped in some way and I am sincerely grateful to all of you. v Chapter Acknowledgments This thesis is comprised of four papers that are either submitted, in review, or in preparation for publication in peer-reviewed scientific journals (Chapters 2 – 5). Co-author Marc Cadotte provided statistical expertise, editorial comments, and consultation on research topic (Chapter 4). Chapter 1 provides the framework for this dissertation research and Chapter 6 summarizes the major findings and importance of this work in the field of Ecology. Schriever, T.A. and Williams, D. D. Accepted. Ontogenetic and individual diet variation in amphibian larvae across an environmental gradient. Freshwater Biology Schriever, T.A. and Williams, D. D. In review. Duration matters, size does not in pond food webs: a study of the dynamic constraints and ecosystem size hypotheses. Oecologia Schriever, T.A., Cadotte, M. W. and Williams, D. D. In preparation. How hydroperiod and species richness affect the balance of resource flows across aquatic-terrestrial habitats. Ecosystems Schriever, T. A. In preparation. Food webs in relation to environmental and community variation: a multivariate approach. vi Table of Contents Contents Acknowledgments.......................................................................................................................... iv Chapter Acknowledgments ............................................................................................................ vi Table of Contents .......................................................................................................................... vii List of Tables ................................................................................................................................. xi List of Figures ............................................................................................................................... xii List of Appendices ....................................................................................................................... xvi 1 Introduction .................................................................................................................................1
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