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THE EVOLUTION and REPRODUCTIVE ECOLOGY of OENOTHERA (ONAGRACEAE) by Kyra Neipp Krakos

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THE EVOLUTION and REPRODUCTIVE ECOLOGY of OENOTHERA (ONAGRACEAE) by Kyra Neipp Krakos Washington University in St. Louis Washington University Open Scholarship All Theses and Dissertations (ETDs) 1-1-2011 The volutE ion And Reproductive Ecology Of Oenothera (Onagraceae) Kyra Krakos Washington University in St. Louis Follow this and additional works at: https://openscholarship.wustl.edu/etd Recommended Citation Krakos, Kyra, "The vE olution And Reproductive Ecology Of Oenothera (Onagraceae)" (2011). All Theses and Dissertations (ETDs). 601. https://openscholarship.wustl.edu/etd/601 This Dissertation is brought to you for free and open access by Washington University Open Scholarship. It has been accepted for inclusion in All Theses and Dissertations (ETDs) by an authorized administrator of Washington University Open Scholarship. For more information, please contact [email protected]. WASHINGTON UNIVERSITY Division of Biology and Biomedical Sciences Program in Evolution, Ecology and Population Biology Dissertation Committee: Peter H. Raven, Chairperson Peter C. Hoch Kenneth M. Olsen P. Mick Richardson Barbara Schaal Alan R. Templeton THE EVOLUTION AND REPRODUCTIVE ECOLOGY OF OENOTHERA (ONAGRACEAE) By Kyra Neipp Krakos A dissertation presented to the Graduate School of Arts and Sciences of Washington University in partial fulfillment for the degree of Doctor of Philosophy August 2011 Saint Louis, Missouri ABSTRACT OF THE DISSERTATION Evolution and Reproductive Ecology of Oenothera (Onagraceae) by Kyra N. Krakos Doctor of Philosophy Division of Biology and Biomedical Sciences Program in Evolution, Ecology, and Population Biology Washington University, St. Louis 2011 This dissertation describes the role of pollination in the floral diversification of Oenothera with an integration of both ecological and phylogenetic approaches. Oenothera (Onagraceae) is a model system for studying plant reproductive biology. It provides excellent examples of shifts in reproductive traits such as pollination and breeding system, features that have been important in angiosperm diversification. These systems are evolutionarily labile; they easily shift between different states. These different reproductive traits may shift in a concerted fashion; therefore, a more comprehensive approach to understanding the evolution of these plant systems simultaneously addresses shifts in pollination and breeding system. Using 54 species of Oenothera, I first collected detailed data describing the pollination systems, breeding systems, and floral traits associated with pollinator rewards; and second I determined the ii phylogenetic structure, evolutionary history and relationships among these species. Finally, in that phylogenetic context, I examined the timing and position of transitions in reproductive traits and consider how these traits are associated with pollination and breeding systems. My results offer new insights regarding the specialization of pollination systems and the predictive power of pollination syndromes. I find that specialization in pollination is not accurately characterized by visitation rates alone, and that considering functional groups of visitors to the flowers provides the most informative characterization of pollination systems. I also find that pollination syndromes do not sufficiently or accurately describe these pollination systems. My results also clarify phylogenetic relationships in the genus Oenothera, determine that there have been 13 independent transitions to self-compatibity, and provide the first phylogenetic tree for subsection Kneiffia. I find that pollination and breeding system do not correlate consistently with floral traits, and do not show an association with each other. Finally, I find that the transitions in the reproductive traits reveal a complex and diverse pattern in which shifts in floral traits occur prior and post a transition in pollination system. I also document an example of a rare transition from a generalized pollination system to a specialized pollination system. The placement of floral trait transitions with regards to pollinator shifts suggests selective pressures in floral traits that are predictable and follow transitions to novel dominant pollinator groups, rather than changes in pollination system. iii Acknowledgements A project of this size requires an army of people to complete, and I have received overwhelming support and help in this research. I thank all the members of my committee for all of their support and investment in my education and research. I thank Alan Templeton for his guidance in the analyses, as well as his brilliant teaching. I thank Ken Olsen for his insightful questions, and also for being an excellent example of how to teach at a University. I thank Mick Richardson for all of his support with MOBOT and for his enthusiasm for student research at MOBOT. I thank Barbara Schaal for her guidance, her leadership, her generosity in providing a lab for my research, and her very personal touch in her advocacy for students. I thank Peter Hoch for his hours of assistance with this project, his support and time with Students in the Garden, and for those critical first conversations about this work. Finally, I thank my advisor, Peter Raven, for his support and guidance, for his insightful knowledge of Oenothera, for integrating me into the broader network of botanical researchers, and for inspiring me with his love of science. The resources, staff, and researchers at the Missouri Botanical Garden have been integral to this work. They provided logistical support, plant identifications, logistical support, and access to Oenothera collections and data. I express big appreciation for all that MOBOT has done for me, and that they do for the community. An amazing network of organizations, land owners, and botanists all contributed to the extensive fieldwork necessary to this dissertation. I thank Nels Holmberg, Tim Smith, Martha Younkin, Tom Nagel, James Trager, Gloria and Ron Hoggard, George Yatskievych, Marty Kemper, Marc Phipps, Linda Wallace, Bill Caire, Mike Powell, iv Martin Terry, Patrick Conner, Dave and Judy Dodgen, Matthew Albrecht, Theo Witsell, Cindy Osborne, Carol Ann McCormick, Richard LeBlond, Alan Weakley, Katherine Kennedy, and Anna Strong. A special thanks to Barbara Schaal and her lab for generously supporting and funding the molecular work done in this dissertation. I would also like to thank Allan Larson for the use of his lab facilities. I also give a big thank you to Mike Dyer and the greenhouse staff for letting me plant what proved to be aggressive Oenothera and caring for my plants for years. Logistical support was provided by Missouri Dept of Conservation, Illinois Dept of Conservation, Nature’s Conservancy, Shaw Nature Preserve, Selman Living Labs, Center for Plant Conservation, Great Meadows National Wildlife Refuge, and Harvard University Herbarium. Thank you to Mike Arduser, Phil Kroenig, and Richard Heitzmann for insect identification. I would like to thank the following agencies for funding this work: National Science Foundation (DDIG 0910202), the Howard Hughes Foundation, American Women in Science, Botanical Society of America, Washington University in St. Louis, Sigma Xi, Prairie Biotic Research Inc., Missouri Native Plant Society, Webster Groves Nature Society, and the Missouri Botanical Garden. I want to thank my lab mate, office mate, co-founder of Students in the Garden, and all around partner in crime, Nicole Miller-Struttman. She is an exemplary pollination biologist and provided critical insights as this work began. I owe an enormous debt of gratitude to my colleague and friend, Joshua Reece. None of the molecular work would have been possible without his endless patience, training, and support. His guidance, v expertise, and friendship are invaluable. Guidance, insightful comments, and support were provided by James Beck, Rob Raguso, Marc Johnson, Elena Kramer, Emily Wood, Chuck Davis, Catherine Cardelus, Russ Blaine, Warren Wagner, Sarah Heyman, Brian Allen, Genevieve Croft, Nick Griffin, Kate Waselkov, Nic Kooyers, Peter Bernhardt, Boris Igic, Rachel Levin, Jill Miller, and David Bogler. I had the privilege of mentoring many students who helped with this work. I give a special thank you to my field assistants, Scott Fabricant and Estelle Huang, who braved the wilds in search of Oenothera me; devoted hours to lab work, and worked impressively on their own Oenothera-based projects. I also thank Rebecca Outman, Amy Hawkins, Amelia Hetherington, and Assata Thompson for their vital contributions in the field and lab. I express a heartfelt thank you to all the faculty and staff of Washington University in Saint Louis. They are a wonderful group of people who gave unending support and kindness. I have felt privileged to be a graduate student at this institution. Finally, and with all my heart, I thank my family. I thank my dear husband, Joshua Krakos, for always cheering me on, our lovely and patient daughters, Elizabeth and Katherine, and my brave son Jack. Jack began kindergarten the day I began this doctorate; he is, and always will be, a much better scientist than I am. This dissertation would not have happened without my mother, Elizabeth Neipp, who traveled with me every field season, carrying new born babies, counseling the undergraduates, and being a good sport about whatever crazy place I was chasing plants. Every scientist needs a mother like her. vi Table of Contents Dissertation Abstract…………………………………………………………………......ii Acknowledgments………………………………………………………………………..iv List of Tables……………………………………………………………..……………..viii
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