The Impact of Climate Change on the Flowering and Fruiting Phenology of Arctic Plants in Nunavut, Canada
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The impact of climate change on the flowering and fruiting phenology of Arctic plants in Nunavut, Canada by Zoe Panchen A thesis submitted to the Faculty of Graduate and Postdoctoral Affairs in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biology Carleton University Ottawa, Ontario 2016 Zoe Panchen Abstract Phenology is the timing of nature’s seasonal events. Ambient temperature plays a key role in phenology and hence, as the climate warms, phenology will likely change. This thesis studied the impact of Arctic climate change on Arctic plant flowering and fruiting phenology in Nunavut, Canada. To establish a baseline for current plant phenology, the first question asked was ‘How does flowering phenology vary across Nunavut?’. Contrary to what might be expected, plants at a more northerly location flower earlier or at the same time and for a shorter duration than conspecifics at a more southerly location. Observations of vast differences in flower abundance in three consecutive and climatically-contrasting years highlighted the challenges of reproductive success with weather extremes associated with contemporary climate change given that Arctic plants require three plus years to complete the sexual reproductive cycle. Finally, three methods, employing long-term phenology monitoring, historical phenological records and an elevation gradient, combined with temperature records, were used to ask the questions: ‘How have temperatures in Nunavut changed?’, ‘How have Nunavut Arctic flowering and fruiting times responded to climate change?’ and ‘What is the predicted temperature- sensitivity of Arctic plants to rising temperatures of climate change?’. Annual temperatures in Nunavut are rising faster than the global average. However, in contrast to temperate regions where spring temperatures are rising the most, monthly temperatures in late summer, autumn and winter are rising significantly in Nunavut. Later-flowering species have advanced flowering times more than early-flowering species and seed dispersal times have advanced more than flowering times. Flowering time temperature- sensitivity is species specific and Nunavut region specific with mid-summer-flowering ii species more sensitive than early- and late-flowering species and Nunavut Arctic archipelago plants more sensitive than Nunavut mainland conspecifics. That Arctic plants’ reproductive phenological events are temperature-sensitive is a good news story suggesting that they will respond to climate change and possibly experience greater reproductive success. Interspecific and inter-regional variation in phenological temperature-sensitivity suggests Arctic plant ecological community structure will alter with climate change but differentially across Nunavut. iii Preface This thesis contains five data chapters. Chapters 2, 3 and 4 are published and Chapters 5 and 6 are in review at refereed journals. I am the sole author of Chapters 3 and 6, while Chapters 2, 4 and 5 were co-authored with my thesis advisor, Dr. Root Gorelick. For all five data chapters I conceived the ideas, designed the studies, collected the data, ran the analysis and wrote the manuscripts. Dr. Root Gorelick provided guidance in his advisory role and editorial suggestions for the manuscripts. All published chapters are included in their entirety in this thesis with minor modification to correct grammar and typos, and provide minor clarifications requested by the thesis examining committee. Chapter 2 is in press as PANCHEN, Z.A. and R. GORELICK. 2016. Canadian Arctic Archipelago conspecifics flower earlier in the High Arctic than the mid-Arctic. International Journal of Plant Sciences. DOI 10.1086/687984. Reprinted in this thesis with permission from The University of Chicago Press Chapter 3 is published as PANCHEN, Z.A. 2016. Arctic plants produce vastly different numbers of flowers in three contrasting years at Lake Hazen, Quttinirpaaq National Park, Ellesmere Island, Nunavut, Canada. Canadian Field-Naturalist 130: 56–63. Reprinted in this thesis with permission from Ottawa Field-Naturalists’ Club. Chapter 4 is published as PANCHEN, Z.A. and R. GORELICK. 2015. Flowering and fruiting responses to climate change of two Arctic plant species, purple saxifrage (Saxifraga oppositifolia) and mountain avens (Dryas integrifolia). Arctic Science 1: 45– 58. DOI 10.1139 /AS-2015-0016. Reprinted in this thesis with permission from Canadian Science Publishing. iv Chapter 5 is in review at Ecology and Evolution as PANCHEN, Z.A. and R. GORELICK. Prediction of Arctic plant phenological sensitivity to climate change from historical records. Chapter 6 is in review at Polar Biology as PANCHEN, Z.A. Space for time: Arctic plant phenological responses to climate change, substituting an elevation gradient as a proxy for the rising temperatures of climate change. v Acknowledgements I would like to thank Dr. Root Gorelick for guiding me through all the ups and downs that is a Ph.D.; his pearls of wisdom were timely, insightful and helpful. I would also like to thank my thesis committee members, Dr. Christopher Burn and Dr. Lynn Gillespie, for their sage advice; their extensive Arctic expertise was valuable in building my knowledge on Arctic climate and Arctic plants and preparing me for my field seasons. I thank Dr. David Inouye and Dr. Brett Stevens for their roles as external examiners of my thesis and for their thoughtful questions during the defense. I feel fortunate and privileged to have had the opportunity to visit and work in Nunavut and I thank the Hamlets and Hunter and Trapper Organisations of Grise Fiord, Iqaluit and Resolute for supporting my research on Inuit land. Over my three years of Arctic field work, my field assistants, Sophia Jain- Schlaepfer, Orla Osborne, Emma Micalizzi, Joan Makaroff, Teresa Tufts and Carly Casey did a stellar job of assisting me with data collection, enduring sometimes less than ideal weather and bugs, and counting more flowers than they would care to remember. At Carleton University I received support and advice from many faculty and peers and I’d like to particularly thank Andrew Simons, Derek Mueller, Joe Bennet, Richard Webster, Tom Hossie, Dani Fraser, Lindsey Derraugh and Felipe Dargent. Outside of the University I received advice and help from Bea Alt, Jennifer Doubt at the Canadian Museum of Nature, Jason Carpenter at Nunavut’s Arctic College, Paul Smith at NWRC, Kyra St. Pierre at the University of Calgary and staff at the National Herbarium of Canada (CAN), Agriculture Canada Vascular Plant Herbarium (DAO), Herbier Louis- Marie (QFA) and University of Toronto at Mississauga Herbarium (TRTE). Some of vi those listed above and the Primack lab, Jeff Saarela, Paul Sokoloff, Johann Wagner and several anonymous reviewers took the time to review some of my manuscripts and I thank them for improving the manuscripts and helping me become a better writer. I received funding from many sources without which, I would not have been able to conduct field work in one of the remotest regions of the Arctic. I would like to thank the following for their financial support: Scholarships: Arctic Institute of North America (Jennifer Robinson Memorial Scholarship); Canadian Federation of University Women (Beverley Jackson Fellowship); Carleton University; Natural Sciences and Engineering Research Council (NSERC); and Ontario Graduate Scholarship (OGS) Program. Grants and travel bursaries: Carleton University; Northern Scientific Training Program (NSTP); and Polar Continental Shelf Program (PCSP). In kind support: Parks Canada’s Nunavut Field Unit. This research was conducted under Nunavut Department of Environment Wildlife Research Permits WL 2013-039 and WL 2014-021; Nunavut Territorial Parks Park Use Permits 2013-01 PU and 2014-02 PU; and Parks Canada Research and Collections Permits QUT 2013-13959 and QUT-2014-16198. Finally, but not least, I would like to thank Chris Macknie for his support and encouragement to the ‘perpetual student’. vii Table of Contents Abstract .............................................................................................................................. ii Preface ............................................................................................................................... iv Acknowledgements .......................................................................................................... vi List of Tables …………………………………………………………………..………..xi List of Figures .................................................................................................................. xv List of Appendices .......................................................................................................... xxi 1 Chapter: Introduction ................................................................................................ 1 1.1 Arctic climate and climate change in the Canadian Arctic ............................................. 3 1.2 Plant reproductive phenology in a changing climate ...................................................... 6 1.3 Arctic plant reproductive phenology and temperature ................................................... 9 1.4 Thesis Overview ........................................................................................................... 12 2 Chapter: Canadian Arctic Archipelago conspecifics flower earlier in the High Arctic than the mid-Arctic ............................................................................................. 16 2.1 Abstract .......................................................................................................................