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Population Genetics and Breeding Ecology of the Rare Clonal Shrub, Spiraea Virginiana (Rosaceae)

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Population Genetics and Breeding Ecology of the Rare Clonal Shrub, Spiraea Virginiana (Rosaceae) Population Genetics and Breeding Ecology of the Rare Clonal Shrub, Spiraea virginiana (Rosaceae) A dissertation submitted to the Graduate School of the University of Cincinnati in partial fulfillment of the requirements for the degree of Doctor of Philosophy In the Department of Biological Sciences of the College of Arts and Sciences by Jessica R. Brzyski M.S. Biology Georgia Southern University, December 2006 Committee Chair: T. M. Culley, Ph.D. i ABSTRACT Two of the most prevalent reasons cited for the decrease in species abundance are the loss and modification of habitat and the impact from invasive species. Riparian species face both of these challenges, being in a habitat that experiences abundant water flow modifications and experiencing a degree of disturbance which is often desired by invasive species. As a result, riparian habitat contains high levels of biodiversity, and also a high frequency of rare species. Therefore, the goal of my research was to identify genetic and reproductive factors that may be hindering population growth of rare riparian species. Spiraea virginiana can be classified as a characteristic riparian shrub, is considered rare throughout its natural range, and it is suggested to be negatively impacted by competition with the invasive S. japonica. Using this study species, I examined the extent of both clonal growth and sexual reproduction, and seed germination potential using field and laboratory methods. Genetic analyses show that S. virginiana is highly clonal and populations are isolated from one another. Genetic data also indicate that S. virginiana is likely to have been rare for an extended time period rather than recently so. Possibly as a consequence of long-term isolation, sexual reproduction is minimal and there is evidence to suggest that the self-incompatibility system is breaking down. When sexual reproduction does occur and seeds are produced, their viability and germination rates are low, being significantly lower than those of S. japonica. I conclude that the prolonged survival of S. virginiana has been the result of prolific clonal propagation. Although clonality allows the species to persist and expand, it greatly diminishes the adaptive potential of the species. The lack of mates in a self-incompatible system, or the ability to self, combined with low germination rate also contributes to the rarity of this species. Since many riparian species are both clonal and self-incompatible, the information and management recommendations provided through this research could also be applied to these species of similar life history to benefit restoration efforts. ii Copyright 2011 By Jessica R. Brzyski iii ACKNOWLEDGMENTS Thank you to my graduate advisor, Dr. Theresa Culley, for all of her advice and guidance. I really appreciate the freedom she gave me to develop my own project. Thank you for accepting a person with virtually no genetic knowledge, and teaching me the genetic techniques and appreciation I needed to accomplish this research. Anyone who can take a field ecologist and turn her into a geneticist must be a great advisor. In addition, my Research Advisory Committee: Dr. Eric Maurer, Dr. Stephen Matter, Dr. Steve Rogstad, and Sarena Selbo of the U.S. Fish and Wildlife Service, were all instrumental to the project and all provided valuable assistance in the construction and implementation of this research. Thank you all for your knowledge and experience, edits and comments, and general assistance throughout this research which has furthered my professional and personal development. Due to the rarity of my study organism, I needed many permits to collect the samples necessary for this project and for that I thank U.S. Fish and Wildlife Service, National Park Service, Daniel Boone National Forest, OH Department of Natural Resources, TN Department of Environment and Conservation, and the Great Smoky Mountain National Park. The people in charge of permits were all extremely helpful and without them I would not have been able to do this research at all. The assistance I received in the field was indispensable. I could not have done field sampling without the help of Martin McAllister (OH Department of Natural Resources), Andrea Bishop (TN Natural Heritage Program), Deb White and Tara Littlefield (KY Natural Heritage Program), Marie Kerr (Big South Fork NRRA), and Janet Rock (Great Smoky Mountains National Park), who all took time out of their busy schedules to show me where my shrub lived. They are some of the most knowledgeable and friendly people I have ever met who were not only helpful, but good company as well. iv Thank you to The Arnold Arboretum at Harvard University for allowing me to conduct breeding experiments on their S. virginiana collection, while also providing me with funds to do it through the Deland Award. I particularly want to thank Michael Dosmann for his assistance in setting up this relationship, and Abby Hird, who was indispensable for the pollination study, from putting on pollination bags, to assisting in hand pollinations in the rain, to getting me to the airport on time. I am very grateful for additional funding sources, which came from the Catherine H. Beatty Fellowship by the Garden Club of America, the University Research Council, and Wendel, Weiman, Benedict Award from the University of Cincinnati. Lastly, thank you to the people that are my support system. I am infinitely grateful to Dan Wetzel, my best friend and soon-to-be husband, who provided me with advice, guidance, and stress relief along the way. My parents, Madeline and Ronald Brzyski, and my brother Michael, have supported me endlessly in everything I do. My family made me who I am today and I hope I make you proud. v TABLE OF CONTENTS Abstract........................................................................................................................................... ii Acknowledgments.......................................................................................................................... iv List of Tables ................................................................................................................................ vii List of Figures.............................................................................................................................. viii Chapter 1: General Introduction ......................................................................................................1 Chapter 2: Isolation and characterization of microsatellite markers in the rare clonal plant, Spiraea virginiana (Rosaceae)...........................................................................................10 Chapter 3: Genetic variation and clonal structure of the rare, riparian shrub Spiraea virginiana (Rosaceae)..........................................................................................................................20 Chapter 4: Capacity for sexual reproduction in the rare, riparian shrub Spiraea virginiana (Rosaceae)..........................................................................................................................55 Chapter 5: The effect of substrate and stratification in germination rate of a native rare shrub compared to a non-native congener ...................................................................................79 Chapter 6: Managing rare plant species: how identifying when rarity occurred can inform management strategies.....................................................................................................100 Chapter 7: General Conclusion....................................................................................................132 Appendix 1: Diagrams of Spiraea virginiana populations and their genotypes..........................140 vi LIST OF TABLES Chapter 2 Table 1. Characterization of 11 microsatellite primer pairs developed in Spiraea virginiana. Table 2. Results from analyzing primers in three populations of Spiraea virginiana. Chapter 3 Table 1. Descriptive statistics averaged across loci for populations of Spiraea virginiana. Table 2. Genetic description of clonal structure within populations for Spiraea virginiana. Table 3. Average Pgen and Psex values for each genotype within each population of Spiraea virginiana taking into account FIS and number of ramets for each multilocus genotype (MLG) within each population. Table 4. Population assignment results, with confidence intervals (CI) as calculated from BayesAss+. Chapter 4 Table 1. Descriptive statistics of Spiraea virginiana at the self-incompatibility locus. Table 2. Allele frequencies for the self-incompatibility alleles identified within eight populations of Spiraea virginiana. Table 3. Pollination treatments of Spiraea virginiana that produced fruits with viable seed. Chapter 5 Table 1. Main effects of time to germination and germination success were analyzed separately between Spiraea virginiana and S. japonica using Kruskal-Wallis nonparametric test. Chapter 6 Table 1. Results from previous research conducted on Spiraea virginiana. Table 2. List of populations of Spiraea virginiana that were collected, including the state where the population is located, population name, the number of subpopulations that were present within each population, and the number of subpopulations that contained unique alleles. Table 3. Pair-wise comparisons of population differentiation (θ) for Spiraea virginiana. Table 4. Pair-wise migration rates of sampled populations, with clones removed, of Spiraea virginiana.
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