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Longitudinal Diet Studies of Arctic Whales

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Longitudinal Diet Studies of Arctic Whales LONGITUDINAL DIET STUDIES OF ARCTIC WHALES CORY J. D. MATTHEWS A thesis submitted to the Faculty of Graduate Studies of The University of Manitoba in partial fulfillment of the requirements of the degree of DOCTOR OF PHILOSOPHY Department of Biological Sciences University of Manitoba Winnipeg, MB Copyright © 2014 by Cory J.D. Matthews ii ABSTRACT An animal’s foraging ecology can vary over a range of temporal scales, mirroring seasonal and longer term changes in prey availability, as well as ontogenetic shifts in diet and distribution. Obtaining individual-based, longitudinal diet information through direct observation, however, is logistically challenging for marine mammals that pursue and consume prey underwater, and are often widely distributed. Isotopic profiling along continuously growing tissues like teeth and baleen, which archive dietary inputs at the time of growth in their stable isotope composition, allows for chronological dietary reconstructions over multi-year timespans. This thesis reports longitudinal diet studies of three Arctic whale species, killer whales (Orcinus orca), bowhead whales (Balaena mysticetus) and beluga whales (Delphinapterus leucas), derived from serial isotopic measurements along teeth and baleen. Study objectives varied by species, but general goals were to characterize seasonal, ontogenetic, and/or individual diet variation. Results revealed similar trophic-level diet, but regional spatial separation, among eastern Canadian Arctic/Northwest Atlantic killer whales. However, isotope and tooth wear differences between two individuals and the rest of the sampled whales suggested potential specialisation on sharks, while the other whales likely had diets comprising marine mammals. Small within-tooth isotopic variation indicated individual killer whale diets were stable over periods up to 25 yrs. Cyclic isotopic variation along Eastern Canada-West Greenland bowhead whale baleen was consistent with year-round foraging, although at a reduced rate during winter. Resting zooplankton could be an important food resource outside of periods of peak productivity, and accessibility likely drives winter habitat selection. Isotopic cycling did not differ between female iii and male bowheads, or among age classes, indicating similar seasonal foraging patterns despite reported spatial segregation throughout their summer range. Individual beluga whales from three eastern Canadian Arctic populations varied in timing of ontogenetic diet shifts (i.e. weaning age), as well as overall trophic position, which could reflect size-specific energetic requirements and foraging capabilities (with larger whales able to dive deeper to access larger, more energetically profitable prey). As with killer whales, low within- tooth isotopic variation indicated long-term individual diet stability, although diet shifts at sexual maturity in males were consistent with dispersal from natal groups. Population-specific beluga whale diet trends over a period of several decades likely reflected climate-related expansions of southern forage fish. Collectively, findings of seasonal, ontogenetic, and/or individual diet variation contribute a greater understanding of intrapopulation variation in foraging ecology of these species, and of large-scale structuring of Arctic marine ecosystems. iv ACKNOWLEDGEMENTS My academic advisor, Steve Ferguson, supported my research throughout, always making time to discuss research questions and results, finding funding to explore new ideas, and providing opportunities to attend conferences and workshops. Steve promptly reviewed each chapter of this thesis. Norm Halden was a particularly involved committee member who supervised a large component of my Ph.D. research measuring trace element profiles in teeth. This work was ultimately not included in this thesis, but we generated a lot of data that I hope we can revisit. I would also like to thank committee member Gail Davoren for her time and commitment, which always reminded me to consider where my research fit within a broader ecology context. Rob Stewart (DFO, Winnipeg) was an honorary committee member of sorts whose input in the planning stages of my Ph.D. is greatly appreciated. Rob also reviewed Chapter 5. Thank you to the many Inuit hunters from the eastern Canadian Arctic communities of Arviat, Cape Dorset, Coral Harbor, Grise Fiord, Igloolik, Hall Beach, Iqaluit, Kangiqsujuak, Kugaaruk, Pangnirtung, Pond Inlet, Rankin Inlet, Repulse Bay, and Resolute who collected teeth and baleen specimens in community-based monitoring programs. Blair Dunn (DFO, Winnipeg) arranged shipments and assisted with locating specimens, along with Sheila Atchison and Tara Bortoluzzi (DFO, Winnipeg). Killer whale teeth were generously provided by the Canadian Museum of Nature, the Nova Scotia Museum, the Manitoba Museum, the Royal BC Museum, John Ford (DFO, Nanaimo), Jack Lawson (DFO, St. John’s), and Wayne Ledwell (Portugal Cove, NL). Many people generously shared their lab space and equipment: Rob Stewart and Blair Dunn (wet saw and sectioning equipment); Al Turnock (University of Manitoba) and Rick Wastle (DFO, v Winnipeg) (polishing wheel and sonicator); R. Wastle and Ravinder Sidhu (University of Manitoba) (imaging equipment); and Misuk Yun (University of Manitoba) always found time for me to use the micromill and microbalance. Dave Yurkowski, Brent Young, and Ricardo Ibarra assisted with baleen sample collection, Nick Sader assisted with beluga teeth extraction and cleaning, and Randi Anderson assisted with beluga teeth polishing. Paul Middlestead, Wendy Abdi, and Patricia Wickham (G.G. Hatch Stable Isotope Laboratory, University of Ottawa) analysed all samples for nitrogen and carbon isotopic composition, and shared their expertise and hospitality during several lab visits. Brian Popp and Natalie Wallsgrove (Stable Isotope Biogeochemistry Laboratory, University of Hawai’i) performed amino acid specific isotope analysis of killer whale samples, and reviewed Chapter 2. Richard Heemskerk and William Mark (Environmental Isotope Laboratory, University of Waterloo) performed sulfur isotopic analysis on baleen samples. I would especially like to thank Panseok Yang of the LA-ICP-MS Laboratory at the University of Manitoba for his mentorship spanning two years of trace element data collection, and R. Sidhu for her assistance mapping calcium distribution in beluga teeth using electron microprobe analysis. I was lucky to have one of the world’s beluga ageing experts, Barb Stewart, conveniently located just down the hall, which made for many impromptu visits to her lab to discuss interpretation of beluga dentine growth layers. Barb confirmed age estimates of several specimens, and reviewed Chapter 5. I owe much to the bodies of work by Keith Hobson (Environment Canada, Saskatoon) and Seth Newsome (University of New Mexico), which I referred to often. Rebecca Killick (Lancaster University), Theo Koulis (University of Manitoba), and Marianne Marcoux vi (Université du Québec à Rimouski) advised on data analysis and R code. Andy Foote (Uppsala University) reviewed Chapter 2, and Bernard LeBlanc (DFO, Winnipeg) and Nadine Lysiak (Woods Hole Oceanographic Institution) reviewed Chapter 3. Thank you to my labmates for discussions about Arctic ecology and research, and to Cortney Watt in particular, whose love- hate relationship with stable isotope data often fueled my own. Research funding was provided by the ArcticNet Network of Centres of Excellence of Canada, the Nunavut Wildlife Management Board, and various DFO Canada programs, including Canada’s Species at Risk Act and the Nunavut Implementation Fund. Conference funding was provided by the Stu Innes Award, ArcticNet, the Arctic Research Consortium of the United States, the Society for Marine Mammalogy, and the University of Manitoba’s Departments of Biological Sciences and Graduate Studies and Faculty of Science. Generous scholarship funding was gratefully received from the Duff Roblin Fellowship and the E. Scherer Memorial Scholarship at the University of Manitoba, and the Garfield Weston Foundation. Finally, thank you to my roster of dogsitters: Dale Bially, Elly Chmelnitsky, Kyle Green, Carie Hoover, Marilyn Kullman, Chantal Sawatsky, Brent Young, and Dave Yurkowski. Only once was I ever truly concerned, when Kyle phoned during the first day of a conference to ask which end of the leash goes around the dog’s head. Oh, and that time Elly nervously phoned me in the field about a ‘huge’ vet bill that turned out to be just $146 (Elly now has her own dog … enough said). Your commitment, especially Dale and Elly (who each took care of Maddy for a cumulative four months), allowed me to spend late nights at the lab, attend conferences, conduct Arctic fieldwork, and enjoy the odd visit home. vii TABLE OF CONTENTS ABSTRACT ................................................................................................................................... ii ACKNOWLEDGEMENTS ........................................................................................................ iv TABLE OF CONTENTS ........................................................................................................... vii LIST OF TABLES .........................................................................................................................x LIST OF FIGURES ................................................................................................................... xiii THESIS FORMAT AND MANUSCRIPT CLAIMS ............................................................
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