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Thesis Title Goes Here Plant–Pollinator Interactions in a Changing Climate by Jessica Rachel Keenan Forrest 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 Jessica Rachel Keenan Forrest 2011 ii Plant–Pollinator Interactions in a Changing Climate Jessica Rachel Keenan Forrest Doctor of Philosophy Department of Ecology and Evolutionary Biology University of Toronto 2011 Abstract Climate change is shifting the seasonal timing of many biological events, and the possibility of non-parallel shifts in different taxa has raised concerns about phenological decoupling of interacting species. My thesis investigates interactions between climate, phenology, and pollination, using the plants and pollinators of Rocky Mountain meadows as a study system. Interannual variation in timing of snowmelt since the 1970s has been associated with changes in the assemblages of concurrently flowering species in these meadows, suggesting that plant species differ in their phenological responses to climate. Differences between plants and pollinators in responsiveness to changing climate could, in principle, cause early-flowering plants to flower too early in warm years, before pollinators are active. In fact, I found only transient evidence for pollinator deficits in one early-flowering species (Mertensia fusiformis), even in an early-snowmelt year. However, the assemblage of pollinators visiting M. fusiformis does change predictably over the season, with likely consequences for selection on floral morphology in years when pollen is limiting. Hence, early- and late-flowering populations may evolve in response to phenology of the pollinator community. Differences between plant and pollinator phenologies appear to be due to generally lower temperature thresholds for development in plants, combined with microclimate differences between the soil and the above- ground nests of some pollinators. Phenological decoupling between plants and pollinators seems iii possible but unlikely to be catastrophic, since many taxa possess adaptations to temporally variable environments. Nevertheless, for many species, adaptation to novel climates will entail evolutionary change, and species interactions can influence evolutionary trajectories. For species affected by increasing late-summer drought, earlier flowering may be advantageous. However, in laboratory experiments, bumble bees avoid rare, unfamiliar flower types, causing simulated plant populations to fail to adapt to changing conditions. Overall, my work emphasizes the importance of the interplay between species interactions and environmental change. iv Acknowledgments Many people contributed their time, wisdom, expertise, or equipment to the work described here. Without their support, this research would have been impossible. The trap-nest project in particular (Chapter 5) required the help of many people and businesses who donated space, materials, or services for trap-nest construction. I am grateful to many residents of the Crested Butte area for their interest in and help with this project; many trap-nests spent more than three full years in the field, and not one was vandalized (except by cows). My late uncle, John Keenan—master carpenter and lover of nature—let me use his beautiful workshop and helped build the 2008–2009 trap-nests. The Rocky Mountain Biological Laboratory in Gothic, Colorado, was my home and workplace for five summers. Ian Billick (Director), Jennie Reithel (Science Director), and Jessica Boynton (GPS/GIS Technician) provided logistical support and access to a spectacular working environment. Business Manager and kindred spirit billy barr provided entertainment, chocolate, workshop space, and invaluable weather records dating back to 1975. The analysis of co-flowering patterns could not have been conducted without David Inouye’s incomparable long-term dataset from RMBL. David has been the ideal collaborator: encouraging, generous with his data and his many scientific gadgets, and quick to respond to e- mail. The lovely and brilliant Kate Ostevik has my everlasting admiration: her cheerful disposition in the face of biting flies and June snowstorms made her a better field companion than I could ever have hoped for. A list of her virtues would make up a chapter of its own, so I will limit myself to acknowledging the many small improvements she made to my efficiency in the field, and her talent for plant identification, which led her to point out the inconvenient existence of a second Mertensia species in our study area. Several other people played an important role in the development of this thesis. Most of all, I wish to thank my advisor, James Thomson—for knowing the right tool for any occasion; for introducing me to RMBL; for improving my writing; for letting me have the freedom and resources to develop this thesis the way I wanted, for better or worse; and, especially, for understanding what kind of support I needed most. I am tremendously grateful for his faith in v me. Rob Gegear introduced me to the lab in Toronto and the art of bumble bee-wrangling; conversations (well, arguments) with Rob helped shape many of my research ideas. Jane Ogilvie collected bees for me and was a great friend, ally, and collaborator, in Toronto and Colorado. Josie Hughes and Heather Coiner provided sympathy and advice on various aspects of data management and analysis. I am particularly indebted to Josie for teaching me how to efficiently deal with large and messy datasets, and to Heather for teaching me the basics of temperature measurement. I have also benefited from scientific and social interactions with Emily Austen, James Burns, Jessamyn Manson, Nathan Muchhala, Mike Otterstatter, Ali Parker, Jen Perry, Helen Rodd, Barbara Thomson, and Terry Wheeler. Suggestions from my external examiner, Alison Brody, helped improve the thesis. Spencer Barrett, Art Weis, the members of my advisory committee (Peter Abrams, Don Jackson, and John Stinchcombe), and numerous others in the EEB department have made me feel at home in academia and have helped make me a better scientist. Finally, I am grateful to my parents, Margo Keenan and Kenneth Forrest, without whose love I surely would never have made it this far. vi Table of Contents Acknowledgments ........................................................................................................................... v Table of Contents .......................................................................................................................... vii List of Tables ............................................................................................................................... xiii List of Plates ................................................................................................................................ xiv List of Figures ............................................................................................................................... xv List of Appendices ...................................................................................................................... xvii Chapter 1: Context ......................................................................................................................... 1 Species interactions in a changing climate ................................................................................. 2 Climate change in the Rocky Mountains ................................................................................... 4 Pollinators of Rocky Mountain wildflowers .............................................................................. 6 Thesis outline ............................................................................................................................. 7 Chapter 2: Flowering phenology in subalpine meadows: does climate variation influence community co-flowering patterns? ......................................................................................... 11 Abstract ......................................................................................................................................... 11 Introduction ................................................................................................................................... 12 Methods ......................................................................................................................................... 14 Study area ................................................................................................................................. 14 Focal species ............................................................................................................................ 16 Data analysis ............................................................................................................................ 18 Whole community overlap ................................................................................................ 18 Overlap between species pairs .......................................................................................... 21 Results ........................................................................................................................................... 22 Temporal autocorrelation ......................................................................................................... 22 Whole community overlap ......................................................................................................
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