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Replace This with the Actual Title Using All Caps DIFFERENTIAL IMPACTS OF POLLEN QUALITY AND MICROBIAL COMMUNITIES ON GENERALIST AND SPECIALIST BEES VISITING A SHARED FOOD RESOURCE A Dissertation Presented to the Faculty of the Graduate School of Cornell University In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy by Kristen Brochu May 2018 © 2018 Kristen Brochu DIFFERENTIAL IMPACTS OF POLLEN QUALITY AND MICROBIAL COMMUNITIES ON GENERALIST AND SPECIALIST BEES VISITING A SHARED FOOD RESOURCE Kristen Brochu, Ph. D. Cornell University 2018 Abstract: There is mounting evidence for declines in both managed and wild pollinator species, necessitating support for wild bee populations in order to maintain pollination services. Solitary, ground-nesting bees comprise the vast majority of bees in the world; however, these important pollinators are relatively understudied and we know little about how they are impacted by resource availability, pollen quality, and microbial associates. My dissertation research uses plants of the genus Cucurbita as a case study to assess differential health concerns associated with generalist (Bombus impatiens and Apis mellifera) and specialist (Peponapis pruinosa) pollinators on a shared resource. I begin by examining how cucurbit plant chemistry varies across plant tissue types and varieties and how this variation impacts the foraging choices of both generalist and specialist bees in an agricultural system. Next, I assess the physiological costs associated with digesting cucurbit pollen for these generalist and specialist bees. Finally, I explore the microbial community associated with a specialist ground-nesting bee’s brood cells, evaluate how this community changes over space and time, and assess the most likely routes of microbial colonization into brood cells. BIOGRAPHICAL SKETCH Kristen was born on March 11, 1986 to Joanne Labre and Kenneth Brochu. Growing up in Montreal, she has always been fascinated by the natural world. From an early age she loved learning and was relentlessly curious about the world around her. Encouraged by her parents, she completed a BSc in Wildlife Biology at McGill University, where her love of biology was reinforced by excellent field courses and research opportunities. It was at the Macdonald Campus that she discovered her passion for insects and science. Several wonderful mentors fostered her enthusiasm for research, which led her to pursue an MSc in Evolutionary Biology at the University of Toronto. Here she developed international collaborations and was captivated by the diversity she observed during her tropical fieldwork. While her foray into fish biology taught her many valuable skills, her interest in insects won out and she began her PhD in Entomology at Cornell University. Throughout her education, her passion for sharing science with others has only grown with many wonderful opportunities to be involved in science communication and outreach activities. When not travelling or spending quality time with her collecting net, Kristen can usually be found tending to her insect pets, who are the stars of her outreach program. iv In memory of Terry Wheeler, my first mentor v ACKNOWLEDGMENTS It has been an incredible privilege to work with so many wonderful people during the course of my PhD. First, I’d like to thank my advisor, Bryan Danforth, for his support throughout the ups and downs of my various projects. Bryan was always ready to share his amazing wealth of knowledge about bees and I can only aspire to one day know as much. His great enthusiasm for natural history is infectious and I have many fond memories of collecting bees with Bryan. I would also like to thank my special committee, Andre Kessler and Anurag Agrawal, for their invaluable expertise which truly enriched my graduate experience. Andre was critical in helping me to develop my ideas and brainstorming with him was always enlightening, motivating, and best of all, fun. Anurag’s astute insights improved the quality of my work and his talent for seeing the ‘big picture’ perspective helped me to hone my research interests. The Entomology Department as a whole has been a wonderful academic home for the last six years. Thank you to Scott McArt for generously giving me access to his lab space and equipment, and to Nelson Milano for helping me to set up my microcolonies. Collaborating with the McArt lab has been a wonderful experience, both in and out of the darkroom. Thank you to Kyle Wickings for offering helpful advice about soil sampling and lending equipment to help me accomplish it. I loved spending time up in Geneva and learning more about all that’s underfoot. Thank you to Jessica Petersen and Brian Nault for sharing data and ideas about cucurbit pollinators. Thank you to Michael Mazourek for guidance on growing squash and pumpkin and for his wonderful work on other aspects of cucurbit biology. Thank you to Rob Raguso for many wonderful conversations about pollinators and to Anne Hajek for her fantastic suggestions on working with fungi. Thank you to Laura Harrington vi for her advice on professional development and for generally being a great role model. Thank you to Jason Dombroskie for always helping me to ID odd bugs, and for bringing a bit of Canadian culture to Cornell. Thank you to Linda Rayor for introducing me to entomological outreach at Cornell, for helping me to hone my presentation skills, and above all, for all the bugs. I had so much fun taking and TAing the Naturalist Outreach class and continuing to do outreach afterwards! It will continue to be a big part of my commitment to sharing my love of science. Thank you to Cheryl Gombas, Lisa Westcott, Lisa Marsh, Stephanie Westmiller, Amy Arsenault, Rita Stucky, Laurie McCall, and Alicia Caswell for patiently answering all my weird questions, consistently solving strange administrative problems, and just generally making my life so much easier. I don’t know what I would have done without you! Further afield at Cornell, I would like to thank the staff at the Cornell Greenhouse facilities for all their help growing my plants and for their patience with the cucurbit jungle that I cultivated. I would also like to thank the Statistical Consulting Unit for confirming that although my analyses were always more complicated than they should be (because live animals just do weird things), they were statistically correct. Thank you to Susi Varvayanis, Colleen McLinn, Kimberly Williams, and Derina Samuel for making me a better communicator and teacher. I feel so fortunate that I was able to learn from all of you and I look forward to sharing the skills I gained at all my future academic institutions. I want to thank past and present members of the Danforth lab for their support, encouragement, and camaraderie during my time here. Thanks to Shannon Hedtke, EJ Blitzer, Jason Gibbs, Margarita Lopez-Uribe, Mia Park, and Laura Russo for helping me to focus my early research interests and providing wonderful insights into my experiment designs. I would particularly like to thank Elizabeth Murray, Heather Grab, Maria van Dyke, Mary Centrella, Silas Bossert and Kass Urban-Mead for vii helpful comments on my data analysis, results, and manuscripts. It has been a joy to work with all of you and I’m a much better scientist for your feedback. I want to thank everyone who helped me with fieldwork, especially Maria van Dyke without whom my days in the field would have been much less fun and much less productive. You are absolutely wonderful and I will miss our long talks in the dirt. Thank you to all the squash and pumpkin growers in the Finger Lakes region who let me look for bees and dig giant holes in their fields. All the sites I visited provided a beautiful backdrop for so many lovely sunrises. Thank you also to NSERC, Cornell University, the Entomology Department, the Griswold Endowment, the Rawlins Endowment, and the Hatch Grant for funding my projects and sending me to conferences. Thank you to all my friends who have made my time here so enjoyable. Thank you to Laura, Heather, Elizabeth, Silas, Leticia, Maria, Mary, Kalyn, John, Kass, Joanna, Zoe, Anyi, Nelson, Ashley, Tim, and Maria for many fantastic times in Ithaca. I have so many great memories with you all and I look forward to making many more. Thank you to Roger & Carmela for all the movies. Thank you to Val for always coming to visit me. Thank you to Suji, Jackie, Miao, Kin, and Alice for making my visits home so much fun. Thank you to my family for always supporting me and for listening to me talk endlessly about bugs. I would not be here without your encouragement. Your astute questions have always made my presentations so much better. Please keep sending me photos of bugs to ID. A little less blurry if you can. Thank you to my wonderful partner Paul. For everything. viii TABLE OF CONTENTS BIOGRAPHICAL SKETCH…………………………………………………………….. iv ACKNOWLEDGEMENTS………………………………………………………………. vi LIST OF FIGURES……………………………………………………………………... x LIST OF TABLES……………………………………………………………………… xi CHAPTER 1. POLLEN DEFENSES NEGATIVELY IMPACT FORAGING AND FITNESS IN A GENERALIST BEE………………………………………………………………………. 1 CHAPTER 2. CHEMICAL ECOLOGY OF POLLINATOR DIETARY CHOICES……………... 36 CHAPTER 3. COMMUNITY ECOLOGY OF MICROBES IN THE NESTS OF A SOLITARY, OLIGOLECTIC, GROUND-NESTING BEE………………………………………………… 59 ix LIST OF FIGURES CHAPTER 1 FIGURE 1. Cucurbita pollen use in field and lab experiments……………………….. 13 FIGURE 2. Fitness effects on adult B. impatiens fed various diet treatments………... 15 FIGURE S1. Mass loss due to evaporation over time by treatment…………………... 34 FIGURE S2. Effect of average weight on sucrose consumption……………………… 35 FIGURE S3. Effect of treatment on pollen efficiency………………………………… 35 CHAPTER 2 FIGURE 1. Comparison of chemistry across tissue types…………………………….. 43 FIGURE 2. Non-metric multidimensional scaling plot (NMDS) to compare compound bouquet composition in leaves, nectar, petals, and pollen (all cultivars pooled)…….
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