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Phylogeography and Trematode Parasitism of Oreohelix Land Snails in Southern Alberta

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Phylogeography and Trematode Parasitism of Oreohelix Land Snails in Southern Alberta PHYLOGEOGRAPHY AND TREMATODE PARASITISM OF OREOHELIX LAND SNAILS IN SOUTHERN ALBERTA ZACHARIAH WILLIAM DEMPSEY BSc, Biological Sciences, University of Lethbridge, 2014 A Thesis submitted to the School of Graduate Studies of the University of Lethbridge in partial fulfillment of the requirements for the degree MASTER OF SCIENCE Department of Biological Sciences University of Lethbridge LETHBRIDGE, ALBERTA, CANADA © Zach W. Dempsey, 2017 PHYLOGEOGRAPHY AND TREMATODE PARASITISM OF OREOHELIX LAND SNAILS IN SOUTHERN ALBERTA ZACHARIAH WILLIAM DEMPSEY Date of Defence: June 07, 2017 Dr. Theresa Burg Associate Professor Ph.D. Co-supervisor Dr. Cameron Goater Professor Ph.D. Co-supervisor Dr. Hester Jiskoot Associate Professor Ph.D. Thesis Examination Committee member Dr. Robert Laird Associate Professor Ph.D. Thesis Examination Committee member Dr. Kathleen Weaver Assistant Professor Ph.D. External, Thesis Examination Committee University of La Verne California, U.S.A. Dr. Tony Russell Associate Professor Ph.D. Chair, Thesis Examination Committee ABSTRACT Modern studies in phylogeography integrate many once-disparate scientific fields. This study investigated terrestrial mountain snails, Oreohelix spp., in southern Alberta using DNA markers and the recent emergence of the trematode parasite Dicrocoelium dendriticum. Large-bodied snails in Cypress Hills (CH) and the Rocky Mountains (RM) formed three clades within the species complex O. subrudis. One was geographically widespread, one was restricted to one region in the RM, and one was restricted to CH. Small-bodied snails in CH were determined to be O. cooperi, a rare Oreohelid thought to be imperilled in the western U.S.A. Phylogeographic analyses determined that snails likely colonized and came into contact in CH due to its glacial history. There was significant spatial and seasonal variation in the prevalence of D. dendtriticum in all three lineages of Oreohelix present in CH. Isolation and geological history have played an important role in determining Oreohelid biogeographical patterns in this region. iii ACKNOWLEDGEMENTS This project started as an independent study in the last year of my undergraduate degree. By the time I graduated, it became clear that the ecology, biodiversity, and parasitology of terrestrial snails in Alberta was much more complex and fascinating than anyone had realized. Dr. Theresa Burg and Dr. Cameron Goater agreed to take me on as a Masters student to investigate further, and for that I am grateful. This study has altered the course of my life for the better, and it would not have been possible without the gracious support of my two supervisors. My supervisors have inspired me with the passion they hold for their respective fields – molecular ecology and parasitology. I would also like to thank my committee for their valuable feedback. Both Dr. Robert Laird and Dr. Hester Jiskoot have provided a multi-disciplinary perspective vital for projects such as mine. I would like to thank Dr. Kathleen Weaver for acting as my external. I have cited her work for so long, it was an honour to finally meet her. I would also like to thank Dr. Tony Russell for being the chair of my thesis examination committee. I would also like to acknowledge the various organizations that have funded my work over the past three years. N.S.E.R.C., Alberta Conservation Association Grants in Biodiversity, and the University of Lethbridge have all provided the resources that have allowed me to pursue this research. Funding is always a concern, and the generosity of these organizations have made my research possible. Thanks also to Parks Canada for putting up with me crawling around Waterton looking for snails, and thanks to the kind staff of Cypress Hills Interprovincial Park for providing lodging during my field seasons. I would like to get personal for a moment, as academic life does not exist in a vacuum. Primary among those I wish to thank is Sarah. Looking back, it seems so iv improbable that we ever met at all. Despite our busy schedules over the past few years, I know I am happiest with Sarah. She has supported me without fail from the very beginning. Thank you Sarah for being the most kind, wonderful, amazing, loving, and reliable partner I could ever ask for. I can’t wait for our future together. I would like to acknowledge mom and dad for fostering my love of science, and for always raising me to the best of their abilities. I know I was a handful at times (mostly in Foremost), but they never stopped being good parents. I would like to thank my sister Sarai for her insight, she is wise beyond her years. She may have been a foil to my childhood, but I miss her now that we live far apart. I would also like to thank my extended family for their support. Thanks to Grandma and Grandpa for all of the home- cooked meals and support, thanks to Nana and Papa for getting back in touch with us and visiting from so far away, thanks to Brandon and Crystal for the great movie and game nights, and thanks to Keenan and Kalan for being super fun. Thanks also Sarah’s family. Sarah’s mom and grandma have been very kind and supportive throughout my research. I would like to acknowledge my lab mates, who have been very influential to me. Thanks to Rachael Adams, Colin MacFarlane, Ashley Curtis, and Brad van Paridon for helping me out in the lab when I first got started. Thanks to Arturo Aanaya Villalobos for being a great field assistant. Thanks to Ashley Jensen for teaching me a bit about birds and birding despite my complaints, thanks to Cat Welke for being such a cheerful presence in the lab, thanks to Kathrin Munro for hosting great wine and cheese nights, thanks to Libby Natola for ensuring there’s never a dull moment in the lab, and thanks to Marie Prill for always filling the lab with good-natured laughter. I will never forget the zany escapades of the lab. v I would like to acknowledge the instructors and students of BIOL2000. At first I found teaching duties to be a chore, but the more I taught, the more I enjoyed it. Now I could see myself teaching for a living. This has been in no small part to the other instructors of BIOL 2000, the Principles of Genetics. Quintin Steynen, Helena Danyk, Randall Barley and Joanne Golden have all inspired me to always teach to best of my abilities. Thanks to Kurt Clarke and Jan Tuescher for sharing that journey with me. I see myself in my students, and I would also like to thank them for grounding me in the classroom and reminding me why I like to teach. Thanks also to the members of Pathfinder Society for keeping me sane in these trying times. I will never forget the half-baked antics of The Foxy Five or The Outsiders. Thanks to Ballto for showing me how to give a character personality and voice. Thanks to Josh for showing me how to make a character strong and well-rounded. Thanks to Mark for sharing his 1st edition old-school D&D wisdom and classic dungeoneering skills. And thanks to Russell for showing me what it means to be a patient GM and a gracious player. All of us have shared stories of adventure and shenanigans over the past two seasons of PFS, and I will never forget them. And finally, I would like to thank you, reader. Perhaps you’re one of the few who are obligated to read this, or maybe you just pulled this thesis off the shelf or downloaded it from the university. Regardless, thank you for taking the time to consider this work. I hope that when you close this book or file, you hold a little more knowledge and a little more wonder for terrestrial snails, invasive parasites, and sky islands than before you started. Nothing would make me happier. vi TABLE OF CONTENTS ABSTRACT iii ACKNOWLEDGEMENTS iv LIST OF TABLES ix LIST OF FIGURES x LIST OF ABBREVIATIONS xi CHAPTER 1: General Introduction 1 1.1 General Background 1 1.2 Model system 4 1.2.1 Oreohelix 4 1.2.2 Dicrocoelium dendriticum 6 1.3 Selection and development of molecular markers 7 1.4 Thesis objectives 8 1.5 Predictions 10 1.6 References 12 CHAPTER 2: Distribution of the land snails Oreohelix cooperi Binney, 18 1858, in southern Alberta, Canada 2.1 Abstract 19 2.2 Introduction 20 2.3 Methods 23 2.3.1 Study site 23 2.3.2 Sampling protocol 24 2.3.3 Sequencing and analyses 24 2.3.4 Morphological characteristics 27 2.4 Results 27 2.4.1 Sequencing and analyses 27 2.4.2 Morphology 28 2.5 Discussion 29 2.6 Acknowledgements 34 2.7 References 35 CHAPTER 3: Phylogeography of Oreohelix land snails in southern 44 Alberta and Saskatchewan 3.1 Abstract 45 3.2 Introduction 46 3.3 Methods 48 3.3.1 Study areas and sampling 48 3.3.2 DNA extraction and amplification 50 3.3.3 Data analyses 51 3.4 Results 52 3.4.1 COI 52 3.4.2 ITS2 54 3.4.3 Morphological characteristics 54 3.5 Discussion 55 3.5.1 COI overview 55 3.5.2 ITS2 and cytonuclear discord 57 3.5.3 Glacial history 58 vii 3.5.4 Glacial refugia and colonization 60 3.5.5 PIM vs CSM 63 3.5.6 Contemporary patterns 64 3.5.7 Conclusions and future directions 65 3.6 Acknowledgements 66 3.7 References 67 CHAPTER 4: Spatio-temporal patterns of emerging larval liver fluke 81 (Dicrocoelium dendriticum) in three sympatric species of land snails in southern Alberta, Canada 4.1 Abstract 82 4.2 Introduction 83 4.3 Methods 86 4.3.1 Study area 86 4.3.2 Design of host survey 87 4.3.3 Dissections 87 4.3.4 Molecular diagnostics 88 4.3.5 Analyses 89 4.4 Results 89 4.4.1 Characteristics of host populations 89 4.4.2 Infection patterns 90 4.5 Discussion 91 4.6 Acknowledgements 96 4.7 References 97 CHAPTER 5: General Discussion 107 5.1 Conservation implications and future directions 107 5.2 Phylogeography implications and future directions 109 5.3 Epidemiology of larval D.
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