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Genetic Variation and Ecological Differentiation Between Two Southern Utah Endemics: U.S

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Genetic Variation and Ecological Differentiation Between Two Southern Utah Endemics: U.S Genetic variation and ecological differentiation between two southern Utah endemics: U.S. federally threatened Townsendia aprica and a closely related congener, T. jonesii var. lutea (Asteraceae: Astereae) by Linda Jennings B.Sc, The University of British Columbia, 2000 A Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science in The Faculty of Graduate Studies Botany . .^ivr The University of British Columbia March 2005 © Linda Jennings, 2005 Abstract Townsendia aprica Welsh & Reveal (Asteraceae) is one of 17 rare plants in Utah receiving protection under the U. S. Federal Endangered Species Act. Townsendia aprica (last chance Townsendia) is an herbaceous perennial, narrow endemic, occurring along the Colorado Plateau in southeastern Utah. Recent discoveries have increased its known range from roughly 301 square kilometers at the time it was first listed to 4186 square kilometers. Many of these populations comprise fewer than 100 individuals. It was presumed that small population sizes together with geographical isolation would lead to low levels of genetic variation and inbreeding within the different populations, as a result of genetic drift and restricted gene dispersal. With the use of enzyme electrophoresis, nine informative loci were resolved to gain insight into the level and pattern of genetic variation within T. apnea's range. These patterns were compared with those of another narrow endemic that is a close congener, Townsendia jonesii var. lutea. Analysis of inter- and intra-taxon genetic distances for T. aprica and T. jonesii var. lutea provides important insights into the consequences of rarity, as well as into the genetic and ecological distinctness of the two narrowly endemic taxa. The two taxa have very similar, relatively high values estimated for genetic variation parameters based on isozyme analysis of nine loci (%P > 70, A > 2.0, and He > 0.30), with much of the genetic variation being found within populations (FST = 0.060 for T. jonesii var. lutea and FST = 0.100 for T. aprica). No significant inbreeding found was in most populations of either taxon. Forty percent of the variation in genetic distance between populations of T. jonesii var. lutea could be explained by geographical distance, where as only about 10% of genetic variation in T. aprica's could be explained by physical distance. Bottlenecks were detected in more than half of each taxon's populations. The results also reveal that populations of each taxon are more similar to one another (FPT = 0.14), than to populations of the other taxon (FST = 0.21). Significant differences were found in characteristics of the soils on which the two taxa occur, though so far, all measurable morphological characters are overlapping in these two taxa. TABLE OF CONTENTS ABSTRACT II TABLE OF CONTENTS IV LIST OF FIGURES ; V LIST OF TABLES VI ACKNOWLEDGEMENTS VII CHAPTER 1 INTRODUCTION 1 1.1 RARITY, ENDEMISM AND UTAH 1 1.2 CONSERVATION GENETICS , 4 1.3 TOWNSENDIA : 7 1.4 TOWNSENDIA APRICA AND T. JONESII WAR. LUTEA 9 1.5 OBJECTIVES 15 CHAPTER 2 MATERIALS AND METHODS 16 2.1 FIELD SAMPLING 16 2.2 LAB METHODS 21 2.2.1 Pollen size andstainability 21 2.2.2 Electrophoretic procedures 21 2.2.3 Soil procedure 23 2.2.4 Pappus measurement procedure 23 2.3 ANALYSIS 24 2.3.1 Isozyme analysis 24 2.3.2 Genetic analysis between T. aprica and T.jonesii var. lutea 28 2.3.3 Soil analysis 28 CHAPTER 3 RESULTS 28 3.1 BREEDING SYSTEM IN TOWNSENDIA APRICA 28 3.2 GENETIC DIVERSITY IN TOWNSENDIA APRICA AND T. JONESII VAR. LUTEA 30 3.3 RELATEDNESS OF TOWNSENDIA APRICA AND T. JONESII VAR. LUTEA 39 CHAPTER 4 DISCUSSION 44 4.1 BREEDING SYSTEM 45 4.2 LEVELS AND PATTERNS OF GENETIC DIVERSITY 48 4.3 RELATEDNESS CONCLUSIONS 55 4.4 FUTURE DIRECTIONS 57 4.5 CONSERVATION CONSIDERATIONS 58 LITERATURE CITED 60 APPENDIX I TOWNSENDIA APRICA HERBARIUM VOUCHERS EXAMINED 67 APPENDIX H GEOGRAPHICAL DISTANCE 69 APPENDIX III ISOZYME BUFFER RECD7ES 70 APPENDIX IV ALLELE FREQUENCIES 71 APPENDIX V REPRESENTATIVE PHOTOS OF EACH TAXON 73 iv LIST OF FIGURES Figure 1.1. Phylogenetic relationships of the species of Townsendia and a summary of habit and the presence of apomixis in the species. From Beaman, 1957 8 Figure 1.2. Map of Utah showing the geographic range of T. aprica and T.jonesii var. lutea in relation to the northwest boundary of the Colorado Plateau 10 Figure 2.1. Townsendia aprica and T.jonesii var. lutea sites collected in Utah in 2001 and 2002. 17 Figure 3.1. Neighbor-joining tree using Cavalli-Sforza and Edwards chord distances for T. aprica 36 Figure 3.2. Neighbor-joining tree using Cavalli-Sforza and Edwards chord distances for T. jonesii var. lutea 37 Figure 3.3. Graph of regression analysis between pairwise FST and geographical distance between populations for T. aprica 38 Figure 3.4. Graph of regression analysis between pairwise FST and geographical distance between populations for T.jonesii var. lutea 38 Figure 3.5. Neighbor-joining tree using Nei's distances (1978) between T. aprica (1-13) and T. jonesii var. lutea (14-20) 41 v LIST OF TABLES Table 2.1. Townsendia aprica localities and collection information for this study, along with additional information available for each population 19 Table 2.2. Townsendia jonesii var. lutea localities and collection information for this study, along with additional information available for each population 20 Table 2.3. Electrode buffer systems including modifications in pH and run time 22 Table 2.4. Resolved enzymes with each buffer system used 22 Table 3.1. Estimated pollen stainability of Townsendia aprica individuals 29 Table 3.2. Enzymes and loci number resolved 30 Table 3.3. Genetic variability parameters for populations of Townsendia aprica 32 Table 3.4. Genetic variability parameters for populations of Townsendia jonesii var. lutea 32 Table 3.5. Wright's F-statistics (Fis, FIT, FST) for all isozyme loci examined and their means for T. aprica and T. jonesii var. lutea, respectively 33 Table 3.6. Cavalli-Sforza and Edwards (1967) chord distance pairwise comparisons of 13 populations of 71 aprica 35 Table 3.7. Cavalli-Sforza and Edwards (1967) chord distance pairwise comparisons of 7 populations of T. jonesii var. lutea 35 Table 3.8. Nei's genetic distance (1972) (below diagonal) and identity (above diagonal) for all pairwise comparisons between T. aprica (1-13) and T. jonesii var. lutea (14-20) 40 Table 3.9. Soil characteristics comparing T. aprica (N = 23) and T. jonesii var. lutea (N = 7). ..42 Table 3.10. Ray and disk pappus measurements of T. aprica and T. jonesii var. lutea. 43 Table 1.1. All known herbarium vouchers for Townsendia aprica 67 Table 1.2. List populations, sites and herbaria and their acronyms 68 Table II. 1. Geographic distances (km) between the different sampled populations of T. aprica (1- 13) and T. jonesii var. lutea (14-20) 69 Table II.2. Geographic distances (km) between the different sampled populations T. jonesii var. lutea (14-20) 69 Table III. 1. Running buffer recipes used for both T. aprica and T. jonesii var. lutea 70 Table III.2. Extraction buffer recipe used for leaf tissue of T. aprica and T. jonesii var. lutea.... 70 Table IV.1. Allele frequencies of Townsendia aprica 71 Table IV.2. Allele frequencies of Townsendia jonesii var. lutea 71 vi ACKNOWLEDGEMENTS Without a doubt the first person who I am totally indebted to is my supervisor, Jeannette Whitton (a.k.a. The Gambler). She is someone who took a big chance with me, to allow me to utilize my drive to the fullest and gave me the support (money and emotional) needed to make this project and my experiences the best I could possibly hope for. She is a one of a kind "rare" supervisor and I will be forever thankful for this experience with her. I also want to thank my committee members, Sally Aitken and Fred Ganders for taking such an interest in this project. I want to thank Sally for giving her time to look over my analysis and results with a fine tooth comb, asking the critical questions, and to pushing me those extra steps to better understand rare species and population genetics. I want to thank Fred Ganders for taking time to look over my data early on and for helping me to understand how closely linked breeding systems and population genetics really are, and for encouraging me to put all this data into the larger context of speciation. I also want to thank the Whitton lab crew: Gina Choe for her crazy sense of humor, and support over dinner and beer and Chris Sears for taking an interest in T. aprica by continuing on• going studies of this special little plant. I would also like to thank Debbie Clark for spending time with me in the field showing me localities and being the "go to gal" when I needed information from the different US federal agencies about T. aprica. I would also like to add to this list Tom Clark (NPS), Bob Campbell (USFS), Lori Armstrong (BLM), and Larry England (USF&W) for giving me the permits to allow this project to happen. My personal thanks goes out to my mom and sister (Theresa), who call me and write me on a weekly basis to see how I'm doing, if I need anything, and to remind me of how proud they are of me. I also must include my immediate family: Timmie and Ophilia, my cats, without whom I might need an alarm clock and Beckett and Woofie, my dogs, who force me to take "walkies" or pay the consequences.
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