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Moth Pollination, Low Seed Set, and Vestigialization of Attractive Floral Traits in Abronia Umbellata (Nyctaginaceae)

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Moth Pollination, Low Seed Set, and Vestigialization of Attractive Floral Traits in Abronia Umbellata (Nyctaginaceae) MOTH POLLINATION, LOW SEED SET, AND VESTIGIALIZATION OF ATTRACTIVE FLORAL TRAITS IN ABRONIA UMBELLATA (NYCTAGINACEAE) by Laura Ann Doris Doubleday A thesis submitted to the Department of Biology in conformity with the requirements for the degree of Master of Science Queen’s University Kingston, Ontario, Canada August, 2012 Copyright © Laura Ann Doris Doubleday, 2012 Abstract Flowering plants display remarkable phenotypic diversity, especially in reproductive structures, much of which is thought to be associated with pollination by animals. Pollination syndromes are collections of floral traits (e.g. flower colour, shape, odour) that are associated with a plant attracting particular functional groups of animal pollinators. We explored the extent to which traits associated with the moth pollination syndrome translated into pollination by moths in the Pacific coast dune endemic Abronia umbellata and found mixed results: in one year of study, there was no difference in seed set by day- vs. night-pollinated inflorescences, but in another year of study, night-pollinated inflorescences set significantly more seed than those pollinated during the day. We integrate this work with tests of pollen and resource limitation of seed production and with seed set surveys of natural populations to address proximate and ultimate causes of low seed set, finding low rates of pollinator visitation, high pollen limitation of seed production in all populations studied, and no evidence of endogenous resource limitation of seed production. We propose that “excess” flowers may be functionally male, serving to increase outcross siring success. The transition from self-incompatibility and obligate outcrossing to self-compatibility and predominant selfing is the most common evolutionary transition among the flowering plants and traits associated with outcrossing may become reduced across such shifts, potentially through the action of natural selection, especially if pollinators are also herbivores, or if the signals that pollinators use to locate flowers are also used by herbivores. We examined the reduction of attractive visual and olfactory floral traits in A. umbellata across a shift from outcrossing to selfing and found a reduction of all floral traits considered. We found that floral volatile emissions were reduced more strongly than flower size or floral display (number of flowers per inflorescence), but there was no evidence of an ecological cost associated with conspicuousness: we did not find reduced leaf herbivory among selfers relative to outcrossers. ii Co-Authorship Chapter 2: Moth pollination and extremely low seed set in a Pacific coast dune plant Co-authors: Sara Lynskey (Queen’s University undergraduate) and Dr. Christopher Eckert. Laura Doubleday and Chris Eckert designed the pollinator observations, pollinator exclusion experiments, and pollen supplementation experiments and co-designed the endogenous resource manipulation experiments with Sara Lynskey, who also participated in the execution of the experiments in Chapter 2. Laura Doubleday supervised and participated in the execution of all field activities and analyzed the pollinator observations, pollinator exclusion experiments, pollen supplementation experiments, and the resource reallocation section of the endogenous resource manipulation experiments. Chris Eckert analyzed the within- and among-inflorescence endogenous resource manipulation experiments and the seed set survey. Laura Doubleday wrote the manuscript and Chris Eckert edited it. Chapter 3: In fragrante delicto: vestigialization of attractive floral traits across a transition from outcrossing to selfing in Abronia umbellata Co-authors: Dr. Robert Raguso (Department of Neurobiology and Behavior, Cornell University) and Dr. Christopher Eckert Rob Raguso taught Laura Doubleday how to collect and analyze floral volatiles, assisted with the gas chromatography-mass spectrometry (GC-MS) analysis presented in Chapter 3, and suggested first steps to take in analyzing the chemical data. He also encouraged the behavioural assays with Hyles lineata (Chapter 3) and provided the animals, materials, and space to perform them. Laura Doubleday collected floral volatile samples in the field and glasshouse, prepared them for GC- MS and analyzed them. Laura Doubleday and Chris Eckert designed the floral morphology surveys and Laura Doubleday supervised and participated in their execution in the field. Laura Doubleday analyzed all data and wrote the manuscript, which Chris Eckert edited. iii Acknowledgements Many people contributed to this thesis. Angela Boag, Danielle Denley, Benjamin Gresik, and Sara Lynskey were stalwart members of some of the finest field crews ever to grace California’s coastal dunes. Dave Hubbard, Jenny Dugan, and Cristina Sandoval were gracious and generous hosts in the Santa Barbara area. Scott Hodges, Nathan Derieg, Jenny Dugan, Mark Page, Neil Tsutsui, and Virginia Emery shared their labs with us in times of need. Benjamin Gresik, Genevieve Rochon-Terry, Michelle Jacobs, Meagan King, Celina Willis, and Bradley Jackson opened thousands of anthocarps, earning themselves both fruit-opening calluses and honour. Anne Lizius, John Viengkone, Dale Kristensen, and Paul Cooper greatly contributed to the success of the glasshouse endeavour. Rob Raguso was a fantastic mentor and was extremely generous with his time, space, and encyclopedic knowledge of floral scent chemistry. It was a pleasure to interact with his lab group: Ha Nguyen, Holly Summers, Joaquín Goyret, and Michael Wolfin. I have learned a lot from Chris Eckert while working on this project: to remember the big picture, to be precise, and, most importantly, where to find the best donuts. Adriana Lopez- Villalobos, Andy Wong, Anna Hargreaves, Sara Dart, and Stephanie Greer were awesome labmates: great sounding boards, fonts of knowledge, and pretty friendly people too! The last words of thanks are due to my family and friends – I couldn’t have done this without you. iv Table of Contents Abstract ............................................................................................................................................ ii Co-Authorship ................................................................................................................................ iii Acknowledgements ......................................................................................................................... iv Chapter 1 General Introduction ....................................................................................................... 1 1.1 What is pollination? ............................................................................................................... 1 1.2 Animal-pollination ................................................................................................................. 2 1.3 Attracting pollinators ............................................................................................................. 2 1.4 Analyzing floral volatiles ....................................................................................................... 4 1.4.1 Collecting floral volatiles ................................................................................................ 4 1.4.2 Adsorbing and desorbing volatiles .................................................................................. 5 1.4.3 Gas chromatography ....................................................................................................... 6 1.4.4 Mass spectrometry .......................................................................................................... 7 1.5 Thesis objectives .................................................................................................................... 7 1.6 Literature Cited ...................................................................................................................... 7 Chapter 2 .......................................................................................................................................... 9 2.1 Abstract. ................................................................................................................................. 9 2.2 Introduction .......................................................................................................................... 10 2.3 Methods ............................................................................................................................... 16 2.4 Results .................................................................................................................................. 23 2.5 Discussion ............................................................................................................................ 26 2.6 Acknowledgements .............................................................................................................. 33 2.7 Literature Cited .................................................................................................................... 34 Chapter 3 ........................................................................................................................................ 52 3.1 Abstract ................................................................................................................................ 52 3.2 Introduction .......................................................................................................................... 53 3.3 Methods ..............................................................................................................................
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