Mechanisms Structuring Dragonfly Communities Across Canopy and Forest Cover Gradients
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Mechanisms Structuring Dragonfly Communities across Canopy and Forest Cover Gradients by Sarah Kathryn French 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 Sarah Kathryn French 2018 Mechanisms Structuring Dragonfly Communities across Canopy and Forest Cover Gradients Sarah Kathryn French Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto 2018 Abstract Forest regrowth, which is occurring across North America, may have large impacts on the structure of larval dragonfly assemblages. My thesis examined how habitat selection, movement behaviour, and species-sorting shape aquatic larval dragonfly assemblages in response to landscape heterogeneity, including forest cover in the landscape and canopy cover over ponds. The mechanisms structuring aquatic communities across environmental gradients are often difficult to distinguish, so I used multiple approaches to assess the contributions of different mechanisms to these observed patterns. Firstly, I tested the role of adult habitat selection versus larval species-sorting in structuring larval dragonfly assemblages across a canopy gradient using artificial aquatic habitats (i.e. mesocosms) placed across gradients of artificial shading and of natural canopy cover. Adult dragonflies preferentially visited aquatic mesocosms situated under open canopy. I tested for species-sorting effects by placing larvae in mesocosms across a natural canopy cover gradient, but found no evidence of differential mortality or growth. Secondly, I examined the effects of forest cover in the terrestrial landscape on adult dragonfly movement. Adults showed preferences for open field environments rather than closed forest environments, ii moving marginally more often in fields, more often towards fields, and flying longer in fields. However, adults’ direction of movement also depended on what environments they were released in, and their flight times were species-specific. Lastly, I investigated how forest cover in the landscape and pond canopy cover affected the distribution of larval dragonflies, as well as their reassembly in ponds following a drying event. Both local (pond canopy cover, permanence, and area) and landscape (pond connectivity based on distance and amount of forest cover between ponds) heterogeneity affected species diversity and recovery following a drying event. My thesis work provides new insights into how aquatic communities are structured by forest and canopy cover, and how they might respond to forest regrowth. iii Acknowledgments Firstly, thank you to Shannon McCauley for her endless enthusiasm and support, strong mentorship, and for fostering such a fantastic and collaborative group of colleagues. I am incredibly grateful for her patience as I navigated my way through a new field, and for taking a chance on a student who fell asleep during their first interview/car trip together to visit a field station. The McCauley lab group has been a continuous source of support and inspiration. Thank you to Shantel Catania, for her contagious passion for aquatic invertebrates and natural history, and for her assistance in the field during my first year of experiments. Thank you to Dachin Frances, my partner in crime from the beginning, for her help in the field, expertise in the lab, and constant encouragement to the very end. Thank you to Celina Baines, Rosemary Martin, Rosalind Murray, Ilia Ferzoco, and Christopher Searcy, for their unwavering support, friendship, and feedback throughout my degree. I am extremely grateful for the mentorship of Helen Rodd, and her guidance on my committee, and of Helene Wagner, who has helped me to cultivate a love for teaching. Thank you to Benjamin Gilbert, for his constructive feedback on my committee over the years, and to Becky Raboy and Peter Kotanen for their helpful comments on my thesis during my appraisal. A special thanks to Carolyn Moon and Stephanie do Rego for all their administrative help and for fielding an endless barrage of questions. An enormous thanks goes to Stephan Schneider for all his assistance at the Koffler Scientific Reserve, including the construction of field equipment, and for his keen knowledge of and excitement for the outdoors. Funding for my projects was graciously provided by the departments of Ecology and Evolutionary Biology at the University of Toronto, and Biology at the University of Toronto Mississauga, the University of Toronto’s School of Graduate Studies, a Queen Elizabeth II/Pfizer Graduate Scholarship in Science and Technology, a Natural Sciences and Engineering Research Council of Canada Scholarship, and the American Museum of Natural History. Thank you to the Koffler Scientific Reserve for their research support. iv Thank you to the entire UTM graduate community for making my time in the department so enjoyable and entertaining, and specifically Audrey Reid, Aaron LeBlanc, Connor Fitzpatrick, Krystal Nunes, Chris Wong, Ruth Rivkin, Molly Hetherington-Rauth, Mark MacDougall, and Amy Wong, for their incredible friendship throughout my PhD. I am also greatly appreciative of the ongoing support from my family and friends, particularly from the mother/mother-in-law babysitting dream team during my last stages of thesis writing. Lastly, thank you to Shawn French for your love and encouragement, assistance in impromptu snowy field adventures, and for being my sounding board. A special thanks to Desmond, who has been a constant source of joy and who has been extremely patient with me during the writing stage, and to my thesis fetus and now-emerged little larva, Thea, who greatly improved my sleeping habits while writing, perhaps a little more so than necessary, and who thankfully waited until after my defense to make her entrance. v Table of Contents Acknowledgments ...................................................................................................................... iv Table of Contents ....................................................................................................................... vi List of Tables ............................................................................................................................. ix List of Figures ............................................................................................................................. x List of Appendices ....................................................................................................................xii Chapter 1 General Introduction ................................................................................................... 1 Community Assembly ............................................................................................................ 1 1.1 Habitat Selection ............................................................................................................. 1 1.2 Forest Cover Change ....................................................................................................... 3 1.3 Objectives and Research Questions.................................................................................. 5 1.4 Study System ................................................................................................................... 7 1.4.1 Study Taxon ......................................................................................................... 7 1.4.2 Study Location ..................................................................................................... 8 1.5 Goals of Chapters ............................................................................................................ 9 1.5.1 Chapter 2 ............................................................................................................. 9 1.5.2 Chapter 3 ........................................................................................................... 10 1.5.3 Chapter 4 ........................................................................................................... 11 1.6 References ..................................................................................................................... 12 Chapter 2 Canopy Cover Affects Habitat Selection by Adult Dragonflies .................................. 19 Abstract ................................................................................................................................ 19 2.1 Introduction ................................................................................................................... 19 2.2 Materials and Methods .................................................................................................. 23 vi 2.2.1 Artificial Shading Experiment ............................................................................ 25 2.2.2 Natural Canopy Experiment ............................................................................... 28 2.2.3 Larval Performance Experiments ....................................................................... 30 2.3 Results ........................................................................................................................... 32 2.3.1 Artificial Shading Experiment ............................................................................ 32 2.3.2 Natural Canopy Experiment ............................................................................... 32 2.3.3 Larval Performance Experiments ......................................................................