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Constraints on reproductive success of northern fulmars in the Canadian High Arctic by Mark Laurence Mallory A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Biology Carleton University Ottawa, Ontario November 2008 © 2008, Mark Mallory Library and Bibliotheque et 1*1 Archives Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A0N4 Ottawa ON K1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-47484-6 Our file Notre reference ISBN: 978-0-494-47484-6 NOTICE: AVIS: The author has granted a non L'auteur a accorde une licence non exclusive exclusive license allowing Library permettant a la Bibliotheque et Archives and Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par Plntemet, prefer, telecommunication or on the Internet, distribuer et vendre des theses partout dans loan, distribute and sell theses le monde, a des fins commerciales ou autres, worldwide, for commercial or non sur support microforme, papier, electronique commercial purposes, in microform, et/ou autres formats. paper, electronic and/or any other formats. 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Canada Constraints on reproductive success of northern fulmars in the Canadian High Arctic by Mark Laurence Mallory A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Biology Carleton University Ottawa, Ontario November 2008 © 2008, Mark Mallory Abstract The main goal of my thesis was to explore variation in breeding schedules and reproductive success of a seabird, the northern fulmar (Fulmarus glacialis), as a guide to understanding constraints and adaptation of this species for breeding in the Canadian High Arctic. Seabirds occupy top-level trophic positions and are common over much of the marine area of Arctic Canada. Such species should be sensitive to environmental change which is predicted to be considerable and rapid in the Arctic. However, we lack even basic information to predict how environmental changes may affect these birds and other Arctic wildlife. The key physical factor influencing the breeding ecology of High Arctic fulmars is the extent and timing of break-up of sea ice, which influences timing of peak marine food availability. Fulmars breeding in the Canadian High Arctic might benefit from an earlier break-up of sea ice, as predicted by many climate change models. Compared to fulmars nesting farther south at colonies across the North Atlantic or in Alaska, fulmars in the Canadian High Arctic initiated breeding later and had a relatively compressed and synchronized breeding schedule. Furthermore, they arrived at the colony with larger endogenous nutrient reserves, had a longer exodus from the colony prior to egg-laying, took longer incubation shifts, and had lower reproductive success compared to fulmars breeding at southern colonies. High Arctic fulmars also entered primary molt after breeding was completed (or had failed), iii presumably because they needed to offset the High energetic demands of molting and breeding in the Arctic. Despite my finding obvious constraints on Arctic breeding by fulmars, there was still considerable phenotypic variation in sizes of breeding birds and in reproductive behaviours at High Arctic colonies, which seemed somewhat paradoxical. Future work should address how ocean climate change influences the timing of available marine food resources relative to traditional breeding times, as this will likely provide selection against that variation. IV Acknowledgements I submitted my M.Sc. thesis when my son was only a few months old, literally typing up chapters with one hand on an old "386", while holding him with the other hand. It took a long time before I entertained thoughts of doing another degree, but here I sit, 17 years later, completing and submitting my Ph.D. thesis, just prior to my son entering university. Needless to say, this dissertation was a long time in the making, and I have many people to thank for their patience and support in allowing me to finally achieve this goal. First, I thank my supervisor and friend Mark Forbes, who has been patient, supportive and insightful throughout this study. This is actually the second permutation of a doctoral project that I started with Mark, who understood my changing employment situation and allowed me to switch projects along the way. His infectious enthusiasm and stream-of-consciousness teleconferences guided this project into important areas that I had not envisioned at the start. Given that Mark now has four students affiliated with Arctic projects, I think I have almost convinced him to refocus his entire research program to Arctic issues ... almost. Mark, my sincere thanks for everything. I conducted this study while an employee of the Canadian Wildlife Service, and I owe a huge debt of thanks to four people in particular who went to bat for me to get and keep this project going, at a time of uncertain funding for government environmental research. Kevin McCormick, Chief of the Northern Conservation v Division (now retired), knew that I wanted to do a Ph.D. when he hired me in 1999, and it was his casual reference during my second annual appraisal that actually prodded me into this work. Kevin asked, "Weren't you thinking of doing a Ph.D. before you moved to the Arctic?" Somewhat dumbfounded (as I had never spoken with him about this), I responded with a very professional sounding, "Um, yeah", and he quickly added "Well, why isn't that one of the goals on your appraisal?" I had not encountered this sort of strong, forward-thinking managerial support previously. Once I climbed back into my chair, the project(s) began to take shape. Appropriately, Kevin required sound scientific justification for the research, which I provided, and then Gerry McKeating (retired Director, Prairie and Northern Region) and his successor Bill Gummer (also now retired), both provided the necessary approvals. Finally, Siu-Ling Han (not retired!), Head of the Eastern Arctic Section, became my supervisor part-way through the study, and she has continued the friendship and stalwart support for this work that was exhibited by Kevin, Gerry and Bill. Regrettably, the times are a-changing at Environment Canada and I'm not sure that a project like this could be undertaken now, but I remain unwaveringly thankful for these four managers (and friends) who cleared the path for me to conduct this study. I wish I had a brilliant, theoretical, scientific explanation for why I initiated a study on northern fulmars at a remote and relatively hostile site in the Canadian High Arctic. But, in fact, the real reason is much simpler. In 2002, when I was starting to focus on seabirds in my job, I was working with Grant Gilchrist conducting ivory gull vi surveys on Ellesmere Island. We were ahead of schedule, and decided to get dropped off for a couple of beautiful, bright, 24-hr sunlit days at a location neither of us had ever been, above the cliffs at Cape Vera, Devon Island, to scope out the fulmar colony and check for eiders (Grant's main research beasties). Grant and I had discussed what we thought were the biggest priority knowledge gaps for Canadian Arctic seabirds, and it was down to two candidates: anything to do with black-legged kittiwakes, or anything to do with northern fulmars, both of which were largely unstudied in Canada. While fulmars have the annoying habit of projectile vomiting, we both agreed that the whining calls of kittiwakes were probably intolerable for a full field season. As we sat on the fractured sedimentary cobble of the polar desert above the Cape Vera cliffs, having instant coffee and smoked oysters at about 11 p.m. on that July night, and looking down on thousands of fulmars, I think the decision was made in my mind. Little did I know how the weather could change ... Conducting fieldwork at Cape Vera was pretty challenging. Over the duration of this study, we had one entire cabin, 2 outhouses, and four large, canvas-walled tents either blow away, or get destroyed by polar bears. Then there was the 330 m climb from the base camp to the plateau above the cliffs where one needed to observe the fulmars. Then there was the 100 km/h+ scouring winds and associated flying debris and grit each month. Oh yeah, the crash landing on the sea-ice. And the fog. And the snow. Did I mention the bears? I was also provided data from fulmar observations at an equally appealing site, the oft fog-enshrouded and snowy Prince vii Leopold Island. A flock of wonderful, talented and resilient people worked at these field sites, and each in some way put at least one day into gathering fulmar data that were used in this project.