Floral Anthesis Rate and Pollen Limitation in Delphinium Glaucum and Their Consequences for Female Fitness
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University of Calgary PRISM: University of Calgary's Digital Repository Graduate Studies The Vault: Electronic Theses and Dissertations 2016 Floral Anthesis Rate and Pollen Limitation in Delphinium glaucum and their Consequences for Female Fitness Cameron-Inglis, Hazel Cameron-Inglis, H. (2016). Floral Anthesis Rate and Pollen Limitation in Delphinium glaucum and their Consequences for Female Fitness (Unpublished master's thesis). University of Calgary, Calgary, AB. doi:10.11575/PRISM/26188 http://hdl.handle.net/11023/2980 master thesis University of Calgary graduate students retain copyright ownership and moral rights for their thesis. You may use this material in any way that is permitted by the Copyright Act or through licensing that has been assigned to the document. For uses that are not allowable under copyright legislation or licensing, you are required to seek permission. Downloaded from PRISM: https://prism.ucalgary.ca UNIVERSITY OF CALGARY Floral Anthesis Rate and Pollen Limitation in Delphinium glaucum and their Consequences for Female Fitness by Hazel Letitia Cameron-Inglis A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE GRADUATE PROGRAM IN BIOLOGICAL SCIENCES CALGARY, ALBERTA MAY, 2016 © Hazel Letitia Cameron-Inglis 2016 Abstract Phenological mismatch between plants and their pollinators and/or interspecific competition for pollination can cause insufficient pollen quantity and/or inadequate pollen quality to limit seed production and siring. Inflorescence display size, a dynamic part of pollinator attraction, influences pollinator behaviour and movement. Thus, a rapid increase in display size as plants begin flowering caused by initially rapid flower anthesis should promote recruitment of pollinators to newly flowering individuals and species. Tests of two assumptions of this hypothesis for Delphinium glaucum revealed: 1) declining anthesis rate during a plant’s flowering period is an intrinsic characteristic of inflorescence architecture, rather than a response to internal resource economy; and 2) pollinator limitation early during the flowering period reduced ovule fertilization and elevated autogamous self-mating. These results suggest that rapidly increasing initial display size largely contributed to the quality, rather than the quantity of female mating. Consequences for male outcross success remain to be assessed. ii Acknowledgements “It takes a village” is a proverb expressing the belief that it takes a community of people to raise a child. I draw a parallel here in that this thesis would not have come to fruition without the guidance, aid and friendship of many people both within and outside of the academic community. This thesis is the culmination of some of the most challenging and rewarding years of my life. Thank-you to everyone who has helped me along this journey. In particular, I am profoundly grateful to my supervisor, Dr. Lawrence Harder, for his guidance and support. Lawrence has challenged, inspired and trained me to think carefully and critically, to write and speak precisely, to interact with others kindly and patiently, and to conduct and assess research objectively. Thank you Lawrence, for the opportunities you have given me and for your valued mentorship. Additionally, I am indebted to my supervisory committee members, Dr. Sean Rogers and Dr. Mary Reid, as well as other professionals, for their contributions to my research. Sean and Mary, along with my external examiner, Dr. David Goldblum contributed insightful comments and advice on my writing and/or research. Moreover, Sean shared lab space, equipment, and molecular knowledge; Dr. Ed Yeung allowed me to use his equipment for pollen enumeration; John Swann aided with insect identification; John Buckley erected an electric fence around my study plants; and the Biogeoscience Institute supplied weather data from the Barrier Lake Research Station. Many thanks are also owed to my lab mates, fellow students and lab collaborators for their assistance and social support. Dr. Mason Kulbaba and Dr. Wan-Jin Liao developed DNA extraction protocols and/or Delphinium glaucum microsatellites used in iii this study. Further, Mason gallantly mentored me on molecular techniques and boosted my morale during troublesome field and lab work. Caitlin Tomaszewski tirelessly and cheerfully assisted me with seed and pollen enumeration and DNA extractions. Dr. Takashi Ida shared plant community phenology data for my study population. Lauren Sawich was a motivating force of reassurance and cheer throughout the analysis and writing process. These individuals, along with other current and past lab members: Illona Clocher, Lisa O’Donnell, David Robinson, and Paul Simpson, shared valuable information, resources, feedback or encouragement. Similarly, I am grateful to several individuals outside my lab: Ella Bowles, Brandon Allen, Stevi Vanderzwan, Leanna Lachowsky, Yan Liu, Sam Robinson and Riley Waytes, for friendly and illuminating conversations relating to pollination, evolution, or molecular techniques. Above all, I am extremely thankful for my friends and family. My friends Robin Butler, Erin Leson and Jon Pellett maintained my sanity and focus by providing encouragement, empathy and creative distractions. My parents, Evelyn and Gordon, and my in-laws, Mike and Audrey, steadfastly supported me emotionally and materially. My wonderful children, Erik and Jamie, understandingly sacrificed their time with me during long field, lab and writing days. And my partner in life, love and eclectic hobbies, Joel Antler, supported me daily by selflessly providing for our needs, constructively listening to my thoughts, wittily encouraging me to laugh, and patiently reminding me to breathe. Finally, the National Sciences and Engineering Research Council of Canada funded this research through Dr. Lawrence Harder’s Discovery and Accelerator Grants and through my Postgraduate Scholarship M Award. iv Table of Contents Abstract ............................................................................................................................... ii Acknowledgements ............................................................................................................ iii Table of Contents .................................................................................................................v List of Tables .................................................................................................................... vii List of Figures and Illustrations ....................................................................................... viii CHAPTER ONE: INTRODUCTION ..................................................................................1 1.1 Pollinator attraction, floral display dynamics and their contributions to plant mating ......................................................................................................................1 1.2 Pollinator limitation and interspecific competition for pollination services ..............5 1.3 Objectives ..................................................................................................................7 CHAPTER TWO: PHENOLOGICAL MISMATCH BETWEEN A PLANT AND ITS POLLINATORS AND ITS CONSEQUENCES FOR SEED PRODUCTION ...........................................................................................................9 2.1 Introduction ................................................................................................................9 2.2 Materials and Methods .............................................................................................12 2.2.1 Study species and site ......................................................................................12 2.2.2 Display dynamics ............................................................................................14 2.2.3 Pollen, seed and plant enumeration .................................................................15 2.2.4 Pollinator visitation .........................................................................................16 2.2.5 Herbivory .........................................................................................................17 2.2.6 Data analysis ....................................................................................................18 2.3 Results ......................................................................................................................20 2.3.1 Pollinators ........................................................................................................20 2.3.2 Phenologies ......................................................................................................20 2.3.3 Pollen receipt and potential donors .................................................................22 2.3.4 Reproductive output ........................................................................................23 2.4 Discussion ................................................................................................................27 2.4.1 Phenological mismatch ....................................................................................27 2.4.2 Limits on seed production ...............................................................................28 CHAPTER THREE: DO ARCHITECTURAL EFFECTS OR RESOURCE DEPLETION GOVERN DECLINING ANTHESIS RATES WITHIN INFLORESCENCES? ..............................................................................................31