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Ants As Flower Visitors and Their Effects on Pollinator Behavior and Plant Reproduction

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Ants As Flower Visitors and Their Effects on Pollinator Behavior and Plant Reproduction Ants as flower visitors and their effects on pollinator behavior and plant reproduction by Adam Richard Cembrowski A thesis submitted in conformity with the requirements for the degree of Masters of Science Ecology and Evolutionary Biology University of Toronto © Copyright by Adam Cembrowski 2013 Ants as flower visitors and their effects on pollinator behavior and plant reproduction Adam Cembrowski Masters of Science Ecology and Evolutionary Biology University of Toronto 2013 Abstract Ants regularly visit flowers, but they may decrease plant reproductive success by competing with pollinators or damaging reproductive structures. However, how ants may exert these costs needs further clarification. In Chapter 1, I review the literature, finding that flower-visiting ants often have neutral effects on plant fitness. In Chapter 2, using artificial flowers with male and female function, I investigate how interference competition between flower-visiting Myrmica rubra ants and Bombus impatiens bumblebees changes pollen analogue movement patterns. Ant presence and scent significantly reduced pollen analogue donation and reception because bees avoided flowers with ant cues. In Chapter 3, to assess the frequency of palynivory among ants, I conducted an acetolysis survey of 75 neo- and paleo-tropical ant species. Ants consistently contained low numbers of pollen grains, suggesting opportunistic pollen consumption may be widespread. Altogether, ant visitation may be costly, but the mechanism depends on the plant, pollinator and ant identities. ii Acknowledgments Science is never an individual undertaking. Numerous people made the completion of this thesis possible. First and foremost, Megan Frederickson gave me the freedom to pursue research outside of her realm of expertise, was extremely patient my ever-changing interests and supported the decisions I made in my degree. My committee also deserves strong thanks. James Thomson was instrumental in my studies, giving me lab space, supplies and, most importantly, time and guidance when I had questions about anything pollination, bee or flower related. Ben Gilbert gave statistics help and was helpful with coding questions. As well, thank you to my examination committee, Spencer Barrett and Helen Rodd. The Frederickson and Thomson lab groups helped me immeasurably, giving encouragement, guidance, and reality checks when I went too far off track. Thank you to Kyle Turner, Lina Arcila Hernandez, Kirsten Prior, Jane Ogilvie, Alison Parker, Eric Youngerman, and Rebecca Batstone. Kyle especially helped me in all regards both inside and outside of school and was, without question, the best field roommate and officemate I could have had. Several volunteers, work study students and our lab tech fed ants, washed artificial flowers and counted tiny objects, saving me countless hours, particularly Harry Rusnock, Shannon Meadley Dunphy, Margaret Thompson, and Jackie Day. I would also like to highlight Marcus Guorui Tan in particular for his work collaborating with me and running trials when I was in out of town and being unwaveringly positive about our research. Our experiment would not have been nearly as successful without him. Several people in the department and associated with the lab provided feedback, jokes and friendship, without which grad school would have been a depressing place. So thank you Natalie Jones, Rachel Germaine, Kelly Carscadden, Alex de Serrano, Jordan Pleet, Aaron Hall, Susana Wadgymar, Emily Austen, Eddie Ho, Amanda Gorton, Dorina Szuroczki, Jenn Coughlan, Jon Sanders, and Gabe Miller. People not in the department and both inside and outside of ecology, made my time here possible and more enjoyable than otherwise. Thanks to the Tuner family for taking me in during holidays, Maureen Murray and Matt Mazowita for everything, and my old supervisor Colleen Cassady St. Clair for helping me get where I am. iii Lastly, my family did an astounding amount for me in letting me cultivate my love of biology, buying me books, letting me keep strange animals, and getting me outdoors. So, thank you to my brothers John and Mark, my dad George, and my mom Kay. And to my partner Katherine, for her unbelievable patience, both in regards to my academic life and in her willingness to adapt hers for mine, thank you. This could not have happened without you. “Vor Allem sind aus der Reihe der flügellosen Insecten die weitverbreiteten flügellosen Ameisen sehr unwillkommene Gäste der Blüthen. Und dennoch sind gerade sie nach dem Nectar der Blüthen in hohem Grade lüstern…” “Of all the wingless insects it is the widely dispersed ants that are the most unwelcome guests to flowers. And yet they are the very ones which have the greatest longing for nectar…” A. Kerner 1878, Flowers and their Unbidden Guests iv Table of Contents Acknowledgments.......................................................................................................................... iii List of Tables ................................................................................................................................ vii List of Figures .............................................................................................................................. viii Chapter 1 Ants as flower visitors and their effects on pollinator behavior and plant reproduction ................................................................................................................................1 Abstract ......................................................................................................................................1 Introduction ...............................................................................................................................2 1. Ant interactions with flowers and floral visitors ...................................................................3 2. Ants as pollinators ................................................................................................................12 3. Floral defenses against ants ................................................................................................14 4. Future directions ..................................................................................................................20 References ................................................................................................................................24 Tables .......................................................................................................................................39 Chapter 2 Ants and ant scent reduce bumblebee pollination of artificial flowers .........................54 Abstract ....................................................................................................................................54 Introduction .............................................................................................................................55 Methods ....................................................................................................................................57 Results ......................................................................................................................................62 Discussion .................................................................................................................................63 Acknowledgements ..................................................................................................................66 References ................................................................................................................................68 Figures ......................................................................................................................................73 Chapter 3 Not just for the bees: pollen consumption is common among tropical ants .................75 v Abstract ....................................................................................................................................75 Introduction .............................................................................................................................75 Methods ....................................................................................................................................77 Results ......................................................................................................................................78 Discussion .................................................................................................................................79 Acknowledgements ..................................................................................................................82 References ................................................................................................................................83 Tables .......................................................................................................................................86 Figures ......................................................................................................................................91 Concluding Remarks ......................................................................................................................93 Copyright Acknowledgments ........................................................................................................96 vi List of Tables Table 1.1 List of studies of flower-visiting ants that consumed floral rewards
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