Dissertation Ecology and Plant
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DISSERTATION ECOLOGY AND PLANT DEFENSE OF TWO INVASIVE PLANTS, HYOSCYAMUS NIGER AND VERBASCUM THAPSUS Submitted by Christa E. Fettig Graduate Degree Program in Ecology In partial fulfillment of the requirements For the Degree of Doctor of Philosophy Colorado State University Fort Collins, Colorado Spring 2016 Doctoral Committee: Advisor: Ruth A. Hufbauer John K. McKay Andrew P. Norton Julie A. Savidge Copyright by Christa Elizabeth Fettig 2016 All Rights Reserved ABSTRACT ECOLOGY AND PLANT DEFENSE OF TWO INVASIVE PLANTS, HYOSCYAMUS NIGER AND VERBASCUM THAPSUS Understanding the factors that drive non-native plant populations to succeed in a new range and the ecological and biological differences that set introduced populations apart from their native counterparts can provide insight into ecological and evolutionary processes, as well as information crucial to management. In this dissertation, I present research on two different plant species that have been introduced to North America, both of which can now be found across the United States and throughout Canada. Chapters 1 and 2 focus on Hyoscyamus niger (black henbane, Solanaceae), a poisonous and state-listed noxious weed. In chapter one I experimentally evaluate whether introduced populations in the western United States are annual or biennial. Both of these life cycles are found in the native range, and have a clear genetic basis. I experimentally manipulated vernalization (a cold treatment for 19 weeks), and find that plants in the introduced range are biennial. Vernalization is critical for bolting and flowering to occur within a growing season. Interestingly, given enough time in a greenhouse setting, 26 percent of plants that were not vernalized were able to flower. This is unlikely to happen in nature, however, as warmer regions without a cold period to naturally vernalize plants are typically lacking sufficient resources (e.g. adequate water or space) for this species. Chapter two aims to understand basic biological and ecological characteristics of black henbane in the introduced range, which lays the groundwork for additional ecological and evolutionary research on this species and will also help direct ii appropriate management practices. In a greenhouse experiment, I test the effects of selfing and outcrossing. In field populations, I measure reproductive output, the size of seed banks of introduced populations, the viability of seed collected over four years, patterns of mortality, and fluctuation in the size of 15 populations. Black henbane is self-compatible, and capable of producing copious seed, and generating large seed banks in naturalized populations. Seeds remain viable for multiple years which may contribute to the dynamic fluctuations of field population sizes that were observed over four years. Populations are generally ephemeral, with high mortality at the rosette stage. Chapter 3 is focused on resistance and tolerance to herbivory, and how they might vary between ranges as well as within individual plants as predicted by optimal defense theory. Optimal defense predicts that defenses are allocated to different tissues based on their value to the plant. I use Verbascum thapsus (common mullein, Scrophulariaceae) to evaluate resistance to both a specialist and a generalist herbivore among plants from the native and introduced range and among leaves of different ages. I also measure tolerance to defoliation by simulating three levels of herbivory and evaluating the regrowth of above and below ground biomass. Both native and introduced mullein plants are highly defended against specialist and generalist herbivores, with high levels of both resistance and tolerance. In accordance with optimal defense theory, young leaves are more highly defended than older leaves. iii ACKNOWLEDGMENTS I’d like to first thank my graduate committee members, Ruth Hufbauer, Andrew Norton, John McKay, and Julie Savidge for their time and mentorship. They offered direction with my research and dissertation, neither of which would have been possible without their assistance and commitment. Ruth Hufbauer, my major advisor, guided me through the creative process that is science. Her door has always been open and our discussions have shaped me as a student and a scientist. I am positive that I have become a better teacher because I was given the opportunity to learn from Ruth as her teaching assistant. Ruth’s encouragement and support for my research and teaching have been indispensable on this journey through graduate school. There are many wonderful people who have contributed to the projects and experiments throughout this dissertation. The cooperation of Colorado, Montana, and Wyoming state and county weed supervisors as well as land owners made studying black henbane possible. Chrissy Alba, Jason Fettig, and Emily Freeland provided great company and help in the field. My parents, Paula and Gordon Hardy, helped with field data collection and made summer data collection trips memorable adventures that I will cherish for a life-time. Janet Hardin, Jennifer Matsuura, and a group of undergraduate students assisted me in the greenhouse, and many friends, family, and colleagues from across the country collected seeds from common mullein. I also thank Andrew Norton, Jim zumBrunnen, and Stacey Endriss for statistics consultations and advice on data analyses. Our extended lab group, including past and present members of the Blumenthal, Hufbauer, Norton, and Ode labs, provided thoughtful discussions and feedback on papers and presentations. The Colorado Agricultural Experiment Station, the USDA, the iv Department of Bioagricultural Sciences and Pest Management, the Graduate Degree Program in Ecology, and the William M. Brown Professional Development Award provided funding for this research. I’ve shared time in Ruth’s lab with some really wonderful scientists and friends. I have created treasured memories with Chrissy Alba, Stacey Endriss, Mike Koontz, Sabrina Kumschick, Peter Leipzig-Scott, Steve Rauth, Marianna Szucs, and Hannah Wilbur. Chrissy, Stacey, and Hannah, in particular, have been sources of emotional support and joy. I am also lucky to have shared time at Colorado State with Myron Bruce and Wyatt Williams. I thank all of these individuals for their laughter and friendship. My husband, Jason, has been field help and greenhouse help, my excel specialist, and therapist. He has been my greatest support and cheerleader on this journey. I have learned so much about ecology and myself through this process, and I can’t imagine being where I am without him next to me. v TABLE OF CONTENTS ABSTRACT.................................................................................................................................. ii ACKNOWLEDGMENTS............................................................................................................. iv CHAPTER 1: INTRODUCED NORTH AMERICAN BLACK HENBANE (HYOSCYAMUS NIGER) POPULATIONS ARE BIENNIAL.................................................................................. 1 CHAPTER 1 OVERVIEW........................................................................................................ 1 MANAGEMENT IMPLICATIONS......................................................................................... 2 INTRODUCTION..................................................................................................................... 3 Study system......................................................................................................................... 4 METHODS................................................................................................................................ 5 Statistical analyses................................................................................................................ 7 RESULTS.................................................................................................................................. 8 DISCUSSION.......................................................................................................................... 10 TABLES AND FIGURES....................................................................................................... 14 LITERATURE CITED........................................................................................................... 19 CHAPTER 2: REPRODUCTIVE STRATEGY, PERFORMANCE, AND POPULATION DYNAMICS OF THE INTRODUCED WEED BLACK HENBANE (HYOSCYAMUS NIGER L.).................................................................................................................................... 23 CHAPTER 2 OVERVIEW...................................................................................................... 23 INTRODUCTION................................................................................................................... 24 Study system....................................................................................................................... 26 METHODS.............................................................................................................................. 28 Reproduction...................................................................................................................... 28 Field performance............................................................................................................... 29 Seed ecology......................................................................................................................