PHYLOGEOGRAPHY OF THREE HIGH LATITUDE RESIDENT CORVIDS: CLARK’S NUTCRACKER (NUCIFRAGA COLUMBIANA), EURASIAN NUTCRACKER (NUCIFRAGA CARYOCATACTES), AND GRAY JAY (PERISOREUS CANADENSIS) KIMBERLY MARGARET DOHMS Bachelor of Science Honours, University of Regina, 2001 Masters of Science, University of Regina, 2009 A Thesis Submitted to the School of Graduate Studies of the University of Lethbridge in Partial Fulfilment of the Requirements for the Degree DOCTOR OF PHILOSOPHY Department of Biological Sciences University of Lethbridge LETHBRIDGE, ALBERTA, CANADA © Kimberly M. Dohms, 2016 PHYLOGEOGRAPHY OF THREE HIGH LATITUDE RESIDENT CORVIDS: CLARK’S NUTCRACKER (NUCIFRAGA COLUMBIANA), EURASIAN NUTCRACKER (NUCIFRAGA CARYOCATACTES), AND GRAY JAY (PERISOREUS CANADENSIS) KIMBERLY MARGARET DOHMS Date of Defense: November 30, 2015 Dr. T. Burg Associate Professor Ph.D Supervisor Dr. A. Iwaniuk Associate Professor Ph.D Thesis Examination Committee Member Dr. C. Goater Professor Ph.D Thesis Examination Committee Member Dr. D. Logue Assistant Professor Ph.D Internal Examiner Dr. K. Omland Professor Ph.D External Examiner University of Maryland, Baltimore County Baltimore, Maryland Dr. J. Thomas Professor Ph.D Chair, Thesis Examination Committee DEDICATION For the owl and the pussy cat. (And also the centipede.) iii GENERAL ABSTRACT High latitude resident bird species provide an unique opportunity to investigate patterns of postglacial and barrier-mediated dispersal. In this study, multiple genetic markers were used to understand postglacial colonization by and contemporary barriers to gene flow in three corvids. Clark’s nutcracker (Nucifraga columbiana), Eurasian nutcracker (N. caryocatactes), and gray jay (Perisoreus canadensis) are year-round resident northern hemisphere passerines with ranges encompassing previously glaciated and unglaciated regions and potential barriers to dispersal (e.g. mountain ranges). Using mitochondrial DNA control region sequences, we found limited geographic genetic structure and one glacial refugium in nutcrackers, contrasting with seven distinct genetic groups and five refugia for gray jays. Nuclear microsatellite markers revealed additional and contrasting patterns of near-panmixia in nutcrackers and multiple hierarchical breaks in gray jays. Genetic patterns are explained by differences in natural history traits, specifically food preferences, for these species. iv THESIS ACKNOWLEDGEMENTS Science is a way of life. Science is a perspective. Science is the process that takes us from confusion to understanding in a manner that's precise, predictive and reliable - a transformation, for those lucky enough to experience it, that is empowering and emotional. Brian Greene As I write these acknowledgements and reflect upon this project, I could never have predicted how true Greene’s words would become. I suggest, though, that the scientist is not the only one that goes through this journey; she also takes a lot of people along for the ride. For all of those who were there for part or all of this process, I want to express my immense gratitude. Thank you to my supervisor, Theresa Burg, for taking on a student with so little phylogeography knowledge and encouraging me to make this project my own. This was not always an easy journey for either of us, but I am ultimately grateful that we both stuck with it. Thank you for continuing to believe that I could and would complete this thesis. Thank you to my supervisory committee, Andrew Iwaniuk and Andrew Hurly, for your excellent questions and feedback throughout this process. Special thanks to Cameron Goater for providing me with a second lab home and stepping in as Andy’s replacement on my committee upon his retirement. My experience at the University of Lethbridge will not soon be forgotten for the influence it has had on me as a biologist; my thanks to all of you for being a part of that. v Thank you to my field assistants, Leah, Heather, and Kayleigh, who made every dawn chorus just a little easier to get up for and accompanied me in my mad desire to eat well while we lived out of Scarlett’s canopy. Massive thanks to my labmates, Brendan (the sweetest, surliest wombat and friend I could ever ask for), Paulo (thanks for the mouse!), John (many late night chats, much music, and scotch), and so many others who shared that special time that is grad school. To my “little lab brother”, Michael B., the source of many bear hugs and constant lively debates: I promise not to hold your love of Harper against you, too much. Endless thanks to Melissa, my academic wife, for everything, especially for making me feel like part of your Lethbridge family. Thank you, Sarah, for being my insomniac work friend, rainbow sock buddy, and endless source of inspiring bird media. My kindred (lady grad student) spirits, Lori P., Janet N., Jennifer D., Alice M., Sue B., Kimberly G., Leslie S., Tanya S., Elly K., and Mikaela D.: thank you for sharing in the struggles and the moments that defined this trip we all shared; your existence in my life reminds me to keep fighting the good fight. To my parents and my Baba: thank you for always telling me I could do anything I wanted for a career and for believing in me, even when you were really unsure about my decisions. Thank you for continuing to ask me if I was “done yet” and, “when [I would] finally stop going to school”; even though I may have seemed annoyed by your reminders, they kept me from packing it all in on a few occasions. I think this is the day you were waiting for...probably. Guess what, vi Mom? I am going to be a doctor, just not quite the type you thought; at least I can still help you ID all those backyard birds. Dad, thank you for always reminding me to check my oil and fluids, keep my engine tuned, and be creative and frugal in solutions to any of my problems, mechanical and otherwise. Baba, thank you, again, for fueling another grad degree with perogies and unconditional love. Thank you to my “little” sister, Angela, for trying to understand my biologist brain and always greeting that with dark humour and baking. It is not an easy brain to figure out, yet you keep trying, stubborn Dohms girl. To those who I met along this journey that left an impression on me, thank you: Cuatro and Gracie (for the heart opening, the music, and the Colorado love), Darroch (for bird nerd talk, advice, so much coffee, and Gros Morne home place), Marc (for the music and intense nerding out sessions), Jackson (for the Waskesiu adventures and the Tour de Fish), and Robin in San Bernadino, California (for all of the local help and continued cheerleading). Thank you, Kevin and Jeremy, for your sage post-doc words, adventurous spirits, and always open doors. Thank you, Mike, for all of the birding and other adventures, for encouraging me to move to Vancouver, and for introducing me to a community that would become my home. To my chosen Vancouver family and community: I cannot properly express my gratitude to you for providing me with a physical and heart home. To Cole: how do I even begin to thank you for all of your encouragement, care, vii communication, music, and perfect sunny side up eggs? Your appearance in my life was unexpected and a total game changer. Never stop holding me to the standards you do. And last, but never least: to Jordan, my BFF, who has been there for this whole trip. I could go on for chapters about all of the ways you have supported and helped me through this process. This has been a “Slow Show”, for certain, and your tough love, kindness, patience, and reality checks have been there every step of the way. If it were possible, all of you would be authors on this thesis. This science, this “way of life”, would not have happened without you. viii TABLE OF CONTENTS Dedication ..................................................................................................................................... iii GENERAL ABSTRACT ............................................................................................................. iv THESIS ACKNOWLEDGEMENTS ......................................................................................... v LIST OF TABLES ....................................................................................................................... xii LIST OF FIGURES .................................................................................................................... xiii LIST OF ABBREVIATIONS, ACRONYMS, AND SYMBOLS ........................................ xiv Chapter 1: General introduction ................................................................................................. 1 1.1 Phylogeography .................................................................................................................. 1 1.2 Barriers to gene flow .......................................................................................................... 2 1.2.1 Behavioural/natural history ....................................................................................... 2 1.2.2 Physical/geographic .................................................................................................... 3 1.2.2.1 North America ........................................................................................................ 4 1.2.2.2 Eurasia ..................................................................................................................... 4 1.3 Pleistocene glaciations ......................................................................................................