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(Tsuga Canadensis (L.) Carrière) in Minnesota a THESIS SUBM

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(Tsuga Canadensis (L.) Carrière) in Minnesota a THESIS SUBM The Genetic Diversity and Conservation Potential of Eastern Hemlock (Tsuga canadensis (L.) Carrière) in Minnesota A THESIS SUBMITTED TO THE FACULTY OF UNIVERSITY OF MINNESOTA BY Emily Kate Ellingson IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE Stan C. Hokanson, Co-advisor James M. Bradeen, Co-advisor September, 2017 © Emily Kate Ellingson 2017 Acknowledgements I would first like to thank my main advisor, Dr. Stan Hokanson, for seeing potential in me as a graduate student and providing the opportunity, resources, guidance, and understanding that have been instrumental in my success. I would like to thank my co- advisor, Dr. Jim Bradeen, for his continual support and encouragement, as well as the rest of my committee, Dr. Andy David, Dr. Andrew Simons, and Dr. Matt Clark, for their helpful review and comments on this thesis. I’d also like to thank Steve McNamara for being a mentor and a resource for all things propagation, and David Stevenson for providing me with opportunities to learn more about curation. For technical support, I would like to thank Jack Tillman, Dr. Matt Clark, Dr. JiJY Sooska- nguan and Dr. Erin Treiber for providing endless lab expertise and advice, as well as humor and understanding. I would not have had seedling data were it not for arborist Louise Levy, who skillfully collected eastern hemlock cones from precarious locations, and Tacy Sickeler, who took care of the many resulting seedlings. I would like to acknowledge a multitude of partners, including the Minnesota Department of Natural Resources and individuals Mark Cleveland, Rebecca Holmstrom, and Ethan Perry who were instrumental in helping me locate eastern hemlock in Minnesota. I am thankful to the organizations that I worked with, such as the Eloise Butler Wildflower Garden, Glensheen Mansion, and the Minnesota Landscape Arboretum. I would also like to acknowledge collaborator Dr. Robert Jetton and his team at North Carolina State University for their advice and help with collections, as well as Dr. Collin Hobbs at Huntington University, whose doctoral work on eastern hemlock was instrumental in helping me develop and optimize lab protocols. Lastly, I will be forever grateful for my family, friends, and fellow graduate students for their support and friendship over the last few years. i Dedication for my family ii Abstract Eastern hemlock (Tsuga canadensis (L.) Carrière) is a slow-growing and long-lived conifer in the Pinaceae family. Its range extends from Nova Scotia west into Wisconsin and Minnesota and south along the Appalachian Mountains, Northern Georgia, and Alabama with outlier populations along the western range limits in Minnesota, Ohio, Indiana, Georgia, Alabama, Kentucky, and Tennessee. Eastern hemlock is a foundation species across its range that has transformational effects on its surrounding ecosystem. As of 2013, eastern hemlock has been listed as near threatened due to the presence of an invasive insect, the hemlock woolly adelgid (Adelges tsugae Anaand.), which is destroying populations in the eastern United States. Eastern hemlock has historically existed in Minnesota in disjunct and marginal populations and it is listed as endangered in the state. Additionally, trees of known native provenance at the Minnesota Landscape Arboretum were collected from a now extirpated population near Mille Lacs Lake, Minnesota and there are additional trees of unknown provenance in state and municipal parks and public gardens. The objectives of this research were to understand the propagation potential and genetic diversity of native and unknown provenance eastern hemlock in Minnesota with the aim of using this information to inform conservation strategies. Field site visits revealed that there are less than 40 known native mature eastern hemlock trees in Minnesota, with scattered seedlings and saplings. Information on individual trees and herbarium specimens including details on height, diameter at breast height (DBH), location, and notes on tree health, are included in the supplementary spreadsheet Appendix B. Using previously published microsatellite markers (SSRs) derived from eastern hemlock, we observed inbreeding in disjunct Minnesota native trees when compared with trees in the main range. Hemlock Ravine was the most genetically distinct from all other sites sampled, as were native origin trees iii at the Minnesota Landscape Arboretum. Interestingly, neither of these two sites were similar to the other Minnesota disjunct site, West Duluth. The West Duluth trees were more genetically similar to populations sampled in Wisconsin and Michigan. Seedlings grown from native Minnesota trees also displayed inbreeding. From paternity analyses, we found that trees at the Minnesota Landscape Arboretum are potentially outcrossing with non-native trees. Additionally, trees in Minnesota of both native and non-native origin can be propagated successfully via seed. Trees at the Minnesota Landscape Arboretum in particular are amenable to seed propagation, but had little overall success when propagated vegetatively, with the exception of a singular accession (MLA19). These discoveries can be used to inform conservation practices in Minnesota. We recommend that land managers continue in situ preservation of sites across Minnesota and continue ex situ maintenance of eastern hemlock trees in parks and gardens. We also recommend that land managers focus on native Minnesota trees when sourcing material for propagation, planting, and seed-banking in national and local repositories. iv Table of Contents Page Table of Contents .......................................................................................................... v List of Tables ............................................................................................................... vii List of Figures ............................................................................................................... ix CHAPTER 1: Literature Review of Eastern Hemlock (Tsuga canadensis (L.) Carrière): Perspectives from its Northwestern Range Limit Introduction and History ....................................................................................... 1 Ecology ................................................................................................................3 Horticulture ........................................................................................................ 14 Conservation ...................................................................................................... 18 Objectives .......................................................................................................... 22 Tables ................................................................................................................ 24 Figures ............................................................................................................... 25 CHAPTER 2: The Genetic Diversity of Native and Cultivated Eastern Hemlock (Tsuga canadensis (L.) Carrière) in Minnesota Introduction ........................................................................................................ 27 Materials and Methods ....................................................................................... 31 Results ............................................................................................................... 37 Discussion ......................................................................................................... 41 Conclusions ....................................................................................................... 49 Tables ................................................................................................................ 50 Figures ............................................................................................................... 65 CHAPTER 3: Propagation Methods and Landscape Management Recommendations for Eastern Hemlock (Tsuga canadensis (L.) Carrière) in Minnesota Introduction ........................................................................................................ 69 Materials and Methods ....................................................................................... 71 v Results ............................................................................................................... 76 Discussion ......................................................................................................... 77 Recommendations ............................................................................................. 81 Conclusions ....................................................................................................... 85 Tables ................................................................................................................ 87 Figures ............................................................................................................... 98 Literature Cited .......................................................................................................... 105 Appendix A ................................................................................................................ 118 Appendix B ......................................................................... Supplementary Spreadsheet vi List of Tables Page Table 1-1: Origin and conservation status of Tsuga species and hybrids (Farjon, 2013) .............................................................................................................................24
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