CETACEAN STUDIES USING PLATFORMS OF OPPORTUNITY Rob Williams A Thesis Submitted for the Degree of PhD at the University of St. Andrews 2003 Chapters 1-5 of this thesis have been updated and subsequently published in multi-authored articles in peer-reviewed journals. If required, citations should be made to these updated articles. Full metadata for this item is available in Research@StAndrews:FullText at: http://research-repository.st-andrews.ac.uk/ Please use this identifier to cite or link to this item: http://hdl.handle.net/10023/2836 This item is protected by original copyright Cetacean studies using platforms of opportunity Rob Williams Thesis submitted in partial fulfillment for the degree of DOCTOR OF PHILOSOPHY in the School of Biology, UNIVERSITY OF ST ANDREWS 1 December 2003. ii ACKNOWLEDGEMENTS Leaving Canada to study whales in the Antarctic and to write about them in Scotland was a wonderful adventure. It was made possible by a lot of people, but mostly because Phil Hammond was precisely the supervisor that I needed to help me to turn my ideas into a project. In the following chapters, people who made specific research projects possible are thanked. Those who made the degree possible are thanked here. I should like to start by thanking Phil Hammond for being such a wonderful advisor, especially when his advice included things that I did not want to hear. He was helped in that process by David Borchers and Val Smith, who served on my advisory committee. David has a way of making statistics accessible. Val has managed to make me care about fish diseases. All are wonderful teachers. Sharon Hedley served in an informal advisory capacity, to the extent of giving me a crash course in distance sampling weeks before my first Antarctic field season. The thesis was improved by valuable comments after a very enjoyable viva with Ian Boyd and Per Berggren. For a project on how to study whales on the cheap, this degree sure was expensive. The three-year cost of foreign-student tuition fees at the University of St Andrews (forget living and travel expenses) exceeded that of oceanfront property near my home in British Columbia, Canada. And it was worth every penny. Several people were instrumental in reducing my costs to study with my mentors at SMRU and CREEM, including: Ted Chamberlin and Carol Finlay (McLean Foundation); Miss Gillam (Newby Trust); Sarah Haney (Canadian Whale Institute – in landlocked Bolton, Ontario); Nancy Mackay (The Russell Family Foundation); Tom and Bonnie Murphy (and their anonymous friend at the Jane Marcher Foundation, with logistical support iii from the University of St Andrews American Foundation, Raincoast Conservation Society and Bill Rossiter of Cetacean Society International); the Overseas Research Scholarship; Julia Samuel (Karten Foundation); Vanessa Williams (Whale and Dolphin Conservation Society); and Dean & Kathy Wyatt (Knight Inlet Lodge). The following organisations provided ship time in the Antarctic: Cheesemans Ecology Safaris; the Mars family; Patrick Shaw and Eric Stangeland (formerly of Marine Expeditions, now with Quark Expeditions); and Victoria Underwood- Wheatley and Jorie Butler-Kent of Abercrombie & Kent and the Abercrombie & Kent Global Foundation. The people at SMRU and CREEM make St Andrews a great place to work. In addition, I owe a thank-you to the following for their help: Erin Ashe, Robin Baird, Lance Barrett-Lennard, Kimberley Bennett, Kevin Bone, David Borchers, Nicola Brabyn, Trevor Branch, Steve Buckland, Louise Burt, Kim and Tony Chater, Cheryl Ciccone, Chris Clark, Nic Dedeluk, Greg Donovan, Graeme Ellis, Mike Fedak, Ari Friedlaender, Sarahbeth Giles, Ailsa Hall, Phil Hammond, John Harwood, Sharon Hedley, Aleta Hohn, Sascha Hooker, Dave Johnston, Diana Krall, Jeff Laake, Clare Last, Russell Leaper, Mike Lonergan, David Lusseau, the MacKenzie family, DJ Mar, Fernanda Marques, Vicky Mars, Stephanie Martin, Jason Matthiopolous, the McCain family, Bernie McConnell, Patrick Miller, Mike Moore, Alexandra Morton, Linda Nichol, Dawn Noren, Charles Paxton, Bob Pitman, Randy Reeves, Hugh Rose, David Rosen, Helen Sharp, Sophie Smout, Peter Stevick, Len Thomas, Dave Thompson, Fernando Trujillo, Paul Wade, Jane Watson, Gae Weber, Terrie Williams, Ben Wilson, Arliss Winship, the Wolyniec family, Simon Wood and Alex Zerbini. David iv Bain, John Ford and Andrew Trites got me started in this discipline, and are my co- authors in Chapter 4. My home on Pearse Island, Johnstone Strait, BC, Canada, is my favourite place to be. But working there can be difficult. The island has no electricity, running water or phone lines. My neighbours are pretty terrific though: this year, they helped me to install a solar panel on my roof, to drill pins in the rock to keep the two-way satellite Internet connection (thanks, Sarah!) from blowing away, and to fix a generator. In fact, neighbour Kate’s laptop is bootstrapping some humpback data as I type this. My friends and family are equally supportive, and I’m lucky to have them. Doug Sandilands even gave me a crash course in GIS mapping, and helped me to create the map in Chapter 5. Kerry Irish, Misty MacDuffee, Peter Ross, Ben Wilson and my friends at Raincoast Conservation Society helped me access libraries close-ish to home, and Misty was embarrassingly helpful with formatting and printing drafts. I want to thank my good friend (Red) Sonja Heinrich, whose first words to me should have been a warning. On our first, chance meeting on a ship in the Antarctic, Sonja asked about my thesis topic. My proposal for Chapter 3 triggered the most polite smirk imaginable, and a very sympathetic, if succinct, evaluation from her: “I highly doubt it.” Her next move? To ask Phil Hammond to supervise her PhD. Smart. And finally, I want to thank my Grade Three teacher, Mrs. Kuss, who taught me, among other things, that science is awfully fun. v ABSTRACT As human impact on marine ecosystems continues to grow, so too does the need for sound conservation and management strategies that are informed by science. Cetaceans, the whales, dolphins and porpoises, epitomise this challenge, because they are hard to study, they have been heavily exploited in the past, and because some of their habitats, behaviours and life-history strategies make them acutely vulnerable to human activities. Unfortunately, research on free-ranging cetaceans in remote areas is costly, and financial resources are limited. The approach used in this thesis to acquire inexpensive quantitative information on cetacean populations and behaviour was to seek out platforms of opportunity. Tourism and environmental education projects provided access to remote areas of importance to cetaceans. The topic was explored in two main areas. First, studies were conducted to investigate the use of ships of opportunity in estimating distribution and abundance, namely of Antarctic baleen whales. The second area of interest was the effect of boats on killer whales in the northeast Pacific. Platforms of opportunity proved valuable for collecting data to model the role of measurement error on abundance estimation. Measurement error was found to be a potential source of bias in four distance estimation experiments. Platforms of opportunity could be used to train observers on protocols, and to learn to use range- finding photogrammetric equipment well before conducting dedicated surveys, which would eliminate this source of bias, as well as estimating abundance in some cases. vi Abundance and distribution of three whale species were modelled using data collected aboard Antarctic tourist ships. Spatial modelling techniques were used to model distribution of minke, fin and humpback whales using line-transect data collected from a survey that could not be randomised. Strong gradients in animal density were predicted, which could be used to inform future surveys. In the meantime, rough estimates of abundance were obtained, and this approach shows promise for other areas where lack of resources makes systematic surveys prohibitively expensive. A government-funded environmental education project provided logistical support for two studies that dealt with effects of boats on killer whale behaviour. One quantified the extent to which a particular style of whalewatching was disruptive to whale behaviour, and commercial whalewatchers agreed to halt this activity. The other found that a protected area conferred benefit to killer whales, even though it protects only a fraction of the whales’ habitat for a fraction of the year. The thesis contains four case studies that illustrate how inexpensive methods may be used to obtain practical quantitative information to aid decision-making about conservation and management of wild cetaceans that interact with (i.e., whalewatching), compete with (i.e., fishing) or are exploited by (i.e., whaling) humans. vii Declarations: i. I, …………………, hereby certify that this thesis, which is approximately …… words in length, has been written by me, that it is the record of work carried out by me and that it has not been submitted in any previous application for a higher degree. date …………… signature of candidate …………………………. ii. I was admitted as a research student in ………. [month, year] and as a candidate for the degree of …………….. in ………. [month, year]; the higher study for which this is a record was carried out in the University of St Andrews between ……….. [year] and ……….. [year]. date …………… signature of candidate …………………………. iii. I hereby certify that the candidate has fulfilled the conditions of the Resolution and Regulations appropriate for the degree of ……………. in the University of St Andrews and that the candidate is qualified to submit this thesis in application for that degree. date …………… signature of supervisor …………………………. iv. In submitting this thesis to the University of St Andrews I understand that I am giving permission for it to be made available for use in accordance with the regulations of the University Library for the time being in force, subject to any copyright vested in the work not being affected thereby.