University of Calgary PRISM: University of Calgary's Digital Repository Graduate Studies The Vault: Electronic Theses and Dissertations 2020-12-11 Indigenous Knowledge and Biomarkers of Physiological Stress Inform Muskox Conservation in a Rapidly Changing Arctic Di Francesco, Juliette Di Francesco, J. (2020). Indigenous Knowledge and Biomarkers of Physiological Stress Inform Muskox Conservation in a Rapidly Changing Arctic (Unpublished doctoral thesis). University of Calgary, Calgary, AB. http://hdl.handle.net/1880/112840 doctoral thesis University of Calgary graduate students retain copyright ownership and moral rights for their thesis. You may use this material in any way that is permitted by the Copyright Act or through licensing that has been assigned to the document. For uses that are not allowable under copyright legislation or licensing, you are required to seek permission. Downloaded from PRISM: https://prism.ucalgary.ca UNIVERSITY OF CALGARY Indigenous Knowledge and Biomarkers of Physiological Stress Inform Muskox Conservation in a Rapidly Changing Arctic by Juliette Di Francesco A THESIS SUBMITTED TO THE FACULTY OF GRADUATE STUDIES IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY GRADUATE PROGRAM IN VETERINARY MEDICAL SCIENCES CALGARY, ALBERTA DECEMBER, 2020 © Juliette Di Francesco 2020 Abstract Glucocorticoids play a key role in energy regulation and are mediators of the physiological stress response in mammals. Their concentrations are commonly measured in wildlife to understand the effects of environmental changes and anthropogenic disturbances, but their use is associated with multiple challenges and there is a need for species-specific validation. Muskoxen (Ovibos moschatus) are an essential part of the Arctic ecosystem, where they have a strong economic, nutritional, and sociocultural value for Indigenous communities. Recent population declines and mortality events suggest that muskoxen may be threatened by the multiple environmental changes and associated stressors to which they are increasingly exposed. Overall, I sought to establish fecal glucocorticoid metabolites (FGM) and qiviut (woolly undercoat) cortisol as biomarkers of physiological stress in muskoxen, and to apply these tools together with Indigenous knowledge (IK) to explore potential causes and patterns of physiological stress in wild muskoxen. Through two repeated pharmacological challenges in captive muskoxen, I showed that qiviut cortisol and FGM levels accurately reflect long-term (over the period of the hair’s growth) and short-term changes in circulating cortisol, respectively. I also demonstrated that changes in circulating cortisol are not reflected in qiviut in the absence of growth and highlighted variations across body regions, significant differences in qiviut segments over time, and differences between shed and unshed qiviut. Additionally, I documented IK which provided novel insights on the potential stressors of muskoxen and their specific importance. Finally, I identified important factors influencing qiviut cortisol (sex, geographical location, season, and year), and found associations between qiviut cortisol and marrow fat and lungworm intensity. Findings were interpreted in part collaboratively with IK holders. This work has advanced our understanding of glucocorticoid deposition and stability in hair, and of the limitations and challenges associated with hair glucocorticoid interpretation. It has highlighted the multiple benefits of incorporating IK in wildlife endocrinology studies and provided a framework for doing so. Finally, identifying factors associated with qiviut cortisol is a key step to simultaneously investigating the causes and consequences, both at the individual and population levels, of physiological stress in muskoxen. i Preface This thesis consists of manuscripts that have been published in peer-reviewed journals, submitted for publication, or are intended to be submitted for publication. Juliette Di Francesco designed this study, collected the data, analyzed the data, interpreted the results, and wrote the papers with guidance from her primary supervisor, her thesis committee, and other collaborators. All co-authors provided important technical and intellectual support while revising the papers. Written permission for reproduction in their entirety of the scientific articles included in this thesis has been obtained from the publishers and all co- authors. Published manuscripts Chapter 2 – Di Francesco J, Navarro-Gonzalez N, Wynne-Edwards K, Peacock S, Leclerc L-M, Tomaselli M, Davison T, Carlsson A, Kutz S (2017) Qiviut cortisol in muskoxen as a potential tool for informing conservation strategies. Conservation Physiology, 5(1):cox052. Submitted manuscripts Chapter 3 – Di Francesco J, Hanke A, Milton T, Leclerc L-M, Kugluktuk Angoniatit Association, Gerlach C, Kutz S. Documenting Indigenous knowledge to identify and understand the stressors that affect muskoxen (Ovibos moschatus). ARCTIC (submitted on September 25th, 2020; currently under review). Chapter 4 – Di Francesco J, Mastromonaco GF, Rowell JE, Blake J, Checkley SL, Kutz S. Fecal glucocorticoid metabolites reflect hypothalamic–pituitary–adrenal axis activity in muskoxen (Ovibos moschatus). PLoS ONE (submitted on September 9th, 2020; currently under review). Chapter 5 – Di Francesco J, Mastromonaco GF, Checkley SL, Blake J, Rowell JE, Kutz S. Qiviut cortisol reflects hypothalamic–pituitary–adrenal axis activity in muskoxen (Ovibos moschatus). General and Comparative Endocrinology (submitted on September 8th, 2020; currently under review). Manuscripts intended to be submitted Chapter 6 – Di Francesco J, Kwong GPS, Deardon R, Checkley SL, Mastromonaco GF, Mavrot F, Peacock S, Leclerc L-M, Kutz S. Intrinsic and extrinsic factors associated with increased qiviut cortisol in wild muskoxen (Ovibos moschatus). Intended to be submitted to Conservation Physiology or PeerJ. ii Acknowledgements I am extremely grateful to my supervisor, Dr. Susan Kutz, for giving me the opportunity to undertake this unique PhD research. Thank you for everything that you taught me, for letting me develop with a lot of liberty this amazing project with a lot of freedom while providing constant and invaluable guidance at every step of the way, and for always encouraging me. I also owe a great deal to my committee members, Drs. Gabriela Mastromonaco, Sylvia Checkley, and Craig Gerlach, for their availability and for providing me invaluable feedback and guidance both during the development of my PhD project and the analyses of the results. Thank you for your kindness, mentorship, and support throughout my PhD. This graduate experience has been extremely enriching both professionally and personally, and this has a lot to do with all of you. Drs. Steven Cooke and Mary Pavelka took the time to read my work: I am grateful for the stimulating discussions we had during my defense. This work also benefitted from the input of several major collaborators, both at the University of Alaska Fairbanks and at the University of Calgary. Special thanks go to Drs. Jan Rowell, John Blake, Katherine Wynne-Edwards, Grace Kwong, Melanie Rock, Karin Orsel, and Rob Deardon for their invaluable contributions. Numerous people participated in this project and helped me with sample collection in the field or processing in the lab. I am particularly grateful to all the staff at the Robert G. White Large Animal Research Station of the University of Alaska Fairbanks (Sarah Barcalow, Thalia Souza, Hanna Sfraga, Carla Wiletto, Christine Terzi, Megan Roberts, Jean Rein, Claire Kepner, and Charles Ashlock), Christine Gilman and Patricia Medd at the Endocrinology Laboratory of the Toronto Zoo, Kamala Sapkota, Felix Nwozu, Ruokun Zhou, my summer students (Akaysha Envik, Leslie Bottari, and Mélanie Meyer), and everyone who helped with the tedious qiviut sorting. A special thank you also to all the UCVM staff for their kindness and support (Joy Punsalan, Abir Bachir, Katrine Maurer, Robert Forsyth, and Kasia Wodjyla). An important part of this project involved field work in the community of Kugluktuk, Nunavut. I am deeply grateful to all the hunters and community members who took part in muskox sample kit collection and participated in the interviews. This research would also not have been possible without support from the Kugluktuk Angoniatit Association, and particularly from Amanda Dumond, Larry Adjun, and Darlene Hokanak, or from the Government of Nunavut, especially Lisa-Marie Leclerc, Terry Milton, Kevin Methuen, Russell Akeeagok, and Allen Niptanatiak. It has been a real pleasure collaborating with all of you. I also thank Kugluktuk High School, who gave me the opportunity to do science outreach activities, and particularly Michael Valk, Haydn George, Brett MacCallum, and Rohan Hollingsworth. It was a very fun and enriching experience. Finally, I thank all the students I had the opportunity to teach and all the other wonderful people I got the chance to meet during my stays in the community, and especially Matt Stadnyk, Ronald Ladd, and Savannah Rose Hiko. The communities of Ulukhaktok, Ekaluktutiak, Paulatuk, and Sachs Harbour also participated in this project. My thanks go to the Government of the Northwest Territories, to the local Hunters and Trappers Committees/Organizations, and to all the harvesters who took part in sample kit collection. This project would not have been possible without the multiple funding agencies who supported it: the Morris Animal Foundation, the