Ecological genetics of Penstemon in the Great Basin, U.S.A. BY ANDREA T. KRAMER B.A., Macalester College, 2000 THESIS Submitted as partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biological Sciences in the Graduate College of the University of Illinois at Chicago, 2009 Chicago, Illinois To my husband, Brian Kramer, whose unconditional love, support, and sense of humor were beyond invaluable throughout the course of this thesis. ACKNOWLEDGEMENTS I am indebted to the many individuals who contributed to the research and writing phases of this thesis. I would first like to thank my advisor, Dr. Mary Ashley, and thesis committee members (Dr. Jeremie Fant, Dr. Kay Havens, Dr. Hank Howe, and Dr. Roberta Mason-Gamer) whose support and encouragement allowed me to take on new challenges and achieve my goals throughout this process. This work was supported by an EPA STAR Fellowship. I owe particular thanks to Kay Havens and Peggy Olwell, who helped make this a reality and who continually inspire me to think big and do more, and to Jeremie Fant, whose guidance over the past six years has been invaluable and whose wonderful drawings I gratefully include here. I am also indebted to the many students, staff and volunteers at the Chicago Botanic Garden who contributed their time and enthusiasm during the field work and data collection components of this research, including Rebecca Tonietto, Charlie Flower, Lara Jefferson, Oscar Herrera, Jeb Moyer, Bianca Rosenbaum, Emily Yates, Marian Hoffher, and Krissa Skogen. And I extend my thanks to Evan Kramer as well as the staff at Utah Botanical Center, Idaho Botanical Garden, Boise State University, Bureau of Land Management, U.S. Forest Service, and Zion and Great Basin National Parks for their assistance during the field work components of this research. Finally, many friends and family members provided both technical and emotional support during both the data collection and writing phases of this thesis, including my parents, Barb and Alan Tietmeyer, my mother and father-in-law, Sara Rubel and Eric Kramer, and my husband, Brian Kramer, without whom none of this would have been possible. To each of the above, I extend my deepest appreciation. ATK TABLE OF CONTENTS CHAPTER PAGE 1. USING ECOLOGICAL GENETICS TO GUIDE ECOLOGICAL RESTORATION IN THE GREAT BASIN................................................................................................................................. 1 1.1 Introduction .................................................................................................................... 1 1.2 What is ecological restoration?....................................................................................... 1 1.3 What is ecological genetics?........................................................................................... 2 1.4 What ecological genetics tells us about natural plant populations ................................. 2 1.5 The role of ecological genetics in ecological restoration ............................................... 4 1.6 A particular need for ecological genetics to guide restoration in the Great Basin ......... 7 1.7 References .....................................................................................................................10 2. LANDSCAPE GENETICS IN THE GREAT BASIN: INFLUENCES OF DISTANCE, TOPOGRAPHY AND POLLINATORS...................................................................................................... 14 2.1 Introduction...................................................................................................................14 2.2 Materials and Methods...................................................................................................17 2.2.1 Study sites ......................................................................................................17 2.2.2 Collections .....................................................................................................18 2.2.3 Molecular data ...............................................................................................20 2.2.4 Analysis...........................................................................................................21 2.3 Results............................................................................................................................23 2.3.1 Descriptive statistics of loci ...........................................................................23 2.3.2 Descriptive statistics of species and populations ...........................................25 2.3.3 Population genetic structure...........................................................................26 2.4 Discussion.....................................................................................................................33 2.5 References.....................................................................................................................39 3. QUANTITATIVE AND MOLECULAR GENETIC DIVERGENCE WITHIN AND AMONG SKY ISLANDS IN THE GREAT BASIN: A COMPARISON AMONG PENSTEMON SPECIES .................. 42 3.1 Introduction .................................................................................................................. 42 3.2 Methods ........................................................................................................................ 45 3.2.1 Study populations and common garden sites ................................................ 45 3.2.2 Quantitative trait measures ............................................................................ 48 3.2.3 Site characterization ...................................................................................... 49 3.2.4 Data analysis.................................................................................................. 50 3.3 Results .......................................................................................................................... 52 3.3.1 Common garden survival............................................................................... 55 3.3.2 Differences in quantitative traits by garden, species and population ............ 57 3.3.3 Partitioning quantitative traits ....................................................................... 63 3.3.4 Climatic measures as predictors of measured traits ...................................... 65 3.3.5 Heritability and QST in seed germination requirements................................. 66 3.4 Discussion..................................................................................................................... 67 3.4.1 Quantitative divergence and climate ............................................................. 67 TABLE OF CONTENTS (continued) CHAPTER PAGE 3.4.2 Quantitative and molecular divergence ......................................................... 69 3.4.3 Quantitative trait plasticity ............................................................................ 71 3.4.4 Implications for ecological restoration .......................................................... 71 3.5 References .................................................................................................................... 73 4. INBREEDING AND OUTBREEDING DEPRESSION IN CROSSES SPANNING GEOGRAPHIC AND GENETIC DISTANCES: A COMPARISON OF TWO PENSTEMON SPECIES.......................... 77 4.1 Introduction .................................................................................................................. 77 4.2 Methods ........................................................................................................................ 80 4.2.1 Study species ................................................................................................. 80 4.2.2 Study populations .......................................................................................... 82 4.2.3 Crossing design ............................................................................................. 85 4.2.4 Seed germination ........................................................................................... 87 4.2.5 Greenhouse study .......................................................................................... 89 4.3 Results .......................................................................................................................... 90 4.3.1 Cross success ................................................................................................. 90 4.3.2 Seed germination ........................................................................................... 95 4.3.3 Greenhouse study ........................................................................................ 100 4.4 Discussion................................................................................................................... 102 4.4.1 Inbreeding and outbreeding depression in cross success............................. 102 4.4.2 Fitness measures in seed germination requirements.................................... 105 4.4.3 Outbreeding depression in competitive greenhouse environments ............. 106 4.4.4 Implications for restoration ......................................................................... 107 4.5 References .................................................................................................................. 109 VITA............... ............................................................................................................................