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15 - Energetic linkages between short-term and long-term effects of whale-watching disturbance on cetaceans

An example drawn from northeast Pacific resident killer whales

from Part III - Ecological dimensions of whale-watching

Published online by Cambridge University Press:  05 April 2014

By
David E. Bain ,
Rob Williams  and
Andrew W. Trites
Edited by
James Higham ,
Lars Bejder  and
Rob Williams
David E. Bain
Affiliation:
University of Washington
Rob Williams
Affiliation:
University of St Andrews, Scotland
Andrew W. Trites
Affiliation:
University of British Columbia
James Higham
Affiliation:
University of Otago, New Zealand
Lars Bejder
Affiliation:
Murdoch University, Western Australia
Rob Williams
Affiliation:
University of St Andrews, Scotland
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Summary

Introduction

Many studies have demonstrated short-term behavioural responses by whales and dolphins in the presence of vessels, but the population-level implications of such changes are poorly understood (Lusseau, 2003, 2004; Bejder et al., 2006a; Lusseau & Bejder, 2007). One means for developing such an understanding is to use a modelling framework such as the Population Consequences of Acoustic Disturbance (PCAD) model. PCAD identifies four levels at which data can be collected, and allows for estimates of modelling parameters at one level to be based on measured data at another level (National Research Council, 2005).

The first level contains short-term behavioural responses, such as those that have been the typical focus of studies on effects of whale-watching. Effects vary within and between species, and include changes in respiration patterns, surface active behaviours, swimming velocity, vocal behaviour, activity state, inter-individual spacing, wake riding, approach and avoidance, and displacement from habitat. Collisions may result in injury or death (Wells & Scott, 1997; Laist et al., 2001). More detailed reviews of vessel effects can be found in Lien (2001) and Ritter (2003).

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Type
Chapter
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Whale-watching
Sustainable Tourism and Ecological Management
 , pp. 206 - 228
Publisher: Cambridge University Press
Print publication year: 2014

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References

Aguilar Soto, N., Johnson, N., Madsen, P.T., Tyack, P.L., Bocconcelli, A. & Borsani, J.F. (2006). Does intense ship noise disrupt foraging in deep-diving Cuvier's beaked whales (Ziphius cavirostris)? Marine Mammal Science 22(3), 690–699.CrossRefGoogle Scholar
Allendorf, F.W., Bayles, D., Bottom, D.L., et al. (1997). Prioritizing Pacific salmon stocks for conservation. Conservation Biology 11, 140–152.CrossRefGoogle Scholar
Andrews, R.D., Pitman, R.L. & Balance, L.T. (2008). Satellite tracking reveals distinct movement patterns for Type B and Type C killer whales in the southern Ross Sea, Antarctica. Polar Biology 31, 1461–1468.CrossRefGoogle Scholar
Au, W.W.L. (1993). The Sonar of Dolphins. New York, NY: Springer.CrossRefGoogle Scholar
Au, W.W.L., Nachtigall, P.E. & Pawloski, J.L. (1999). Temporary threshold shift in hearing induced by an octave band of continuous noise in the bottlenose dolphin. Journal of the Acoustical Society of America 106, 2251.CrossRefGoogle Scholar
Au, W.W.L., Ford, J.K.B., Horne, J.K. & Newman-Allman, K.A. (2004). Echolocation signals of free-ranging killer whales (Orcinus orca) and modeling of foraging for chinook salmon (Oncorhynchus tshawytscha). Journal of the Acoustical Society of America 56, 1280–1290.CrossRefGoogle Scholar
Ayres, K.L., Booth, R.K., Hempelmann, J.A., et al. (2012). Distinguishing the impacts of inadequate prey and vessel traffic on an endangered killer whale (Orcinus orca) population. PLoS ONE 7(6): e36842. .CrossRefGoogle ScholarPubMed
Bain, D.E. (1986). Acoustic behavior of Orcinus: Periodicity, sequences, correlations with behavior, and an automated technique for call classification. In Kirkevold, B. & Lockard, J. (Eds), Behavioral Biology of Killer Whales. New York, NY: Liss, pp. 335–371.Google Scholar
Bain, D.E. (2001). Noise-based guidelines for killer whale watching. Paper submitted to the Wildlife Viewing Workshop. Vancouver, BC.
Bain, D.E. (2002). Acoustical properties of pingers and the San Juan Island commercial gill net fishery. NMFS Contract Report 40ABNF701651 (unpublished). 14 pp. Available from National Marine Fisheries Service, 7600 Sand Point Way, NE 98115, USA.
Bain, D.E. (2007). The relative importance of different vessel types in the immediate vicinity of southern resident killer whales. NMFS Contract Report Number AB133F-F-04-SE-1272. 24 pp.
Bain, D. & Balcomb, K.C. (1999). Population trends of southern resident killer whales (Orcinus orca) from 1960–1999 (unpublished). Status Review Group Workshop, December 1999, Wailea, Maui, HA.
Bain, D.E. & Dahlheim, M.E. (1994). Effects of masking noise on detection thresholds of killer whales. In Loughlin, T.R. (Ed.), Marine Mammals and The Exxon Valdez. New York, NY: Academic Press, pp. 243–256.CrossRefGoogle Scholar
Baird, R.W. (2001). Status of killer whales, Orcinus orca, in Canada. Canadian Field-Naturalist 115, 676–701.Google Scholar
Baird, R.W. & Hanson, M.B. (2004). Diet studies of ‘southern resident’ killer whales: prey sampling and behavioral cues of predation. NMFS Contract Report No. AB133F-03-SE-1070. 12 pp.
Baird, R.W., Hanson, M.B. & Dill, L.M. (2005). Factors influencing the diving behaviour of fish-eating killer whales: Sex differences and diel and interannual variation in diving rates.Canadian Journal of Zoology 83, 257–267.CrossRefGoogle Scholar
Barrett-Lennard, L.G., Ford, J.K.B. & Heise, K.A. (1996). The mixed blessing of echolocation: Differences in sonar use by fish-eating and mammal-eating killer whales. Animal Behaviour 51, 553–565.CrossRefGoogle Scholar
Bejder, L., Samuels, A., Whitehead, H., et al. (2006a). Decline in relative abundance of bottlenose dolphins (Tursiops sp.) exposed to long-term disturbance. Conservation Biology 20(6), 1791–1798.CrossRef
Bejder, L., Samuels, A., Whitehead, H. & Gales, N. (2006b). Interpreting short-term behavioural responses to disturbance within a longitudinal perspective. Animal Behaviour 72, 1149–1158.CrossRefGoogle Scholar
Bigg, M.A. & Wolman, A.A. (1975). Live-capture killer whale (Orcinus orca) fishery, British Columbia and Washington, 1962–1973. Journal of the Fisheries Research Board of Canada 32, 1213–1221.CrossRef
Bigg, M.A., Olesiuk, P.F., Ellis, G.M., Ford, J.K.B. & Balcomb, K.C. (1990). Social genealogy of resident killer whales (Orcinus orca) in the coastal waters of British Columbia and Washington State. International Whaling Commission Special Issue 12, 383–405.Google Scholar
Borrell, A., Bloch, D. & Desportes, G. (1995). Age trends and reproductive transfer of organochlorine compounds in long-finned pilot whales from the Faroe Islands. Environmental Pollution 88, 283–292.CrossRefGoogle ScholarPubMed
Candy, J.R. & Quinn, T.P. (1999). Behavior of adult chinook salmon (Onchorhynchus tshawytscha) in British Columbia coastal waters determined from ultrasonic telemetry. Canadian Journal of Zoology 77, 1161–1169.CrossRefGoogle Scholar
Carretta, J.V., Forney, K.A., Lowry, M.S., et al. (2010). U.S. Pacific Marine Mammal Stock Assessments: 2009. U.S. Department of Commerce, NOAA Technical Memorandum NMFS-SWFSC-453. 336 pp.
Clark, C.W., Ellison, W.T., Southall, B.L., et al. (2009). Acoustic masking in marine ecosystems as a function of anthropogenic sound sources. IWC/SC/61/E10. Submitted to Scientific Committee, International Whaling Commission. 19 pp.
D’Vincent, C.G., Nilson, R.M. & Hanna, R.E. (1985). Vocalization and coordinated feeding behavior of the humpback whale in southeastern Alaska. The Scientific Reports of the Whales Research Institute No. 36, 11–47.Google Scholar
de Swart, R.L., Ross, P.S., Vos, J.G. & Osterhaus, A.D. (1996). Impaired immunity in harbour seals (Phoca vitulina) exposed to bioaccumulated environmental contaminants: Review of a long-term feeding study. Environmental Health Perspectives 104, 823–828.CrossRef
Erbe, C. (2002). Underwater noise of whale-watching boats and potential effects on killer whales (Orcinus orca), based on an acoustic impact model. Marine Mammal Science 18, 394–418.CrossRefGoogle Scholar
Fisheries and Oceans Canada. (2008). Recovery Strategy for Northern and Southern Resident Killer Whales (Orcinus orca) in Canada. Species at Risk Recovery Series. Ottawa: Fisheries and Oceans Canada. ix + 81 pp.Google Scholar
Foote, A.D., Osborne, R.W. & Hoezel, A.R. (2004). Whale-call response to masking boat noise. Nature 428(29 April), 910.CrossRefGoogle ScholarPubMed
Ford, J.K.B. & Ellis, G.M. (2005). Prey selection and food sharing by fish-eating ‘resident’ killer whales (Orcinus orca) in British Columbia. Can. Sci. Advisory Sec. Res. Doc. 2005/041. 34 pp.
Ford, J.K.B., Ellis, G.M. & Balcomb, K.C. (2000). Killer Whales. Vancouver: UBC Press.Google Scholar
Ford, J.K.B., Ellis, G.M. & Olesiuk, P.F. (2005). Linking prey and population dynamics: Did food limitation cause recent declines of ‘resident’ killer whales (Orcinus orca) in British Columbia? Can. Sci. Advisory Sec. Res. Doc. 2005/042. 31 pp.
Ford, J.K.B., Ellis, G.M. & Balcomb, K.C. (2010). Linking killer whale survival and prey abundance: Food limitation in the oceans’ apex predator?Biology Letters 6, 139–142.CrossRefGoogle ScholarPubMed
Gilpin, M.E., Case, T.J. & Ayala, F.J. (1976). Theta-selection. Mathematical Biosciences 32, 131–139.CrossRefGoogle Scholar
Griffin, R.M. & Bain, D.E. (2006). Sound exposure of southern resident killer whales in the southern strait of Georgia. Contract Report to the National Marine Conservation Area Feasibility Study. Vancouver, BC. 23 pp.
Hall, J. & Johnson, C.S. (1972). Auditory thresholds of a killer whale Orcinus orca Linnaeus. Journal of the Acoustical Society of America 51, 515–517.CrossRefGoogle Scholar
Hanson, M.B., Baird, R.W., Ford, J.K.B., et al. (2010). Species and stock identification of prey consumed by endangered southern resident killer whales in their summer range. Endangered Species Research 11, 69–82.CrossRefGoogle Scholar
Hauser, D.D., Logsdon, M.G., Holmes, E.E., VanBlaricom, G.R. & Osborne, R.W. (2007). Summer distribution patterns of southern resident killer whales Orcinus orca: Core areas and spatial segregation of social groups. Marine Ecology Progress Series 351, 301–310.CrossRefGoogle Scholar
Heise, K., Barrett-Lennard, L.G., Saulitis, E., Matkin, C. & Bain, D. (2003). Examining the evidence for killer whale predation on Steller sea lions in British Columbia and Alaska. Aquatic Mammals 29(3), 325–334.CrossRefGoogle Scholar
Higham, J.E.S., Bejder, L. & Lusseau, D. (2008). An integrated and adaptive management model to address the long term sustainability of tourist interactions with cetaceans. Environmental Conservation 35(4), 294–302.CrossRefGoogle Scholar
Hilborn, R., Cox, S.P., Gulland, F.M.D., et al. (2012). The Effects of Salmon Fisheries on Southern Resident Killer Whales: Final Report of the Independent Science Panel. Prepared with the assistance of D.R. Marmorek and A.W. Hall, ESSA Technologies Ltd., Vancouver, B.C. for National Marine Fisheries Service (Seattle. WA) and Fisheries and Oceans Canada (Vancouver. BC). xv + 61 pp. + Appendices.Google Scholar
Holt, M.M. (2008). Sound exposure and Southern Resident killer whales (Orcinus orca): A review of current knowledge and data gaps. U.S. Dept. Commer., NOAA Tech. Memo. NMFS-NWFSC-89, 59 p.
Holt, M.M., Noren, D.P., Veirs, V., Emmons, C.K. & Veirs, S. (2008). Speaking up: Killer whales (Orcinus orca) increase their call amplitude in response to vessel noise. Journal of the Acoustical Society of America 125, EL27–EL32.CrossRefGoogle Scholar
Jefferson, T.A. (1987). A study of the behavior of Dall's porpoise (Phocoenoides dalli) in the Johnstone Strait, British Columbia. Canadian Journal of Zoology 65, 736–744.Google Scholar
Jeffries, S., Huber, H., Calambokidis, J. & Laake, J. (2003). Trends and status of harbor seals in Washington State: 1978–99. Journal of Wildlife Management 67, 207–218.CrossRefGoogle Scholar
Jensen, F.H., Bejder, L., Wahlberg, M., Johnson, M., Aguilar Soto, N. & Madsen, P.T. (2009). Masking of delphinid communication by small vessels. Marine Ecology Progress Series 395, 161–175.CrossRefGoogle Scholar
Kastelein, R.A., Walton, S., Odell, D., Nieuwstraten, S.H. & Wiepkema, P.R. (2000). Food consumption of a captive female killer whale (Orcinus orca). Aquatic Mammals 26, 127–131.Google Scholar
Kastelein, R.A., Kershaw, J., Berghout, E. & Wiepkema, P.R. (2003). Food consumption and suckling in killer whales. International Zoo Yearbook 38, 204–218.CrossRefGoogle Scholar
Kasting, N.W., Adderley, S.A.L., Safford, T. & Hewlett, K.G. (1989). Thermoregulation in beluga (Delphinapterus leucas) and killer (Orcinus orca) whales. Physiological Zoology 62, 687–701.CrossRef
Koski, K.L. (2011). 2010 Soundwatch Program annual contract report. Contract Number: AB133F-07-CN-0221 Tasks 2.1.2 & 2.1.3. 75 pp.
Krahn, M.M., Ford, M.J., Perrin, W.F., et al. (2004). Status review of Southern Resident killer whales (Orcinus orca) under the Endangered Species Act. US Department of Commerce, Vol. NMFS-NWFSC-62.
Krahn, M.M., Hanson, M.B., Baird, R.W., et al. (2007). Persistent organic pollutants and stable isotopes in biopsy samples (2004/2006) from Southern Resident killer whales. Marine Pollution Bulletin 54, 1903–1911.CrossRefGoogle ScholarPubMed
Kriete, B. (1995). Bioenergetics in the killer whale, Orcinus orca. PhD dissertation (unpublished). University of British Columbia, Vancouver, BC.
Kruse, S. (1991). The interactions between killer whales and boats in Johnstone Strait, B.C. In K. Pryor & K.S. Norris (Eds), Dolphin Societies: Discoveries and puzzles. Berkeley, CA: University of California Press, pp. 148–159.Google Scholar
Lachmuth, C.L., Barrett-Lennard, L.G., Steyn, D.Q. & Milsom, W.K. (2011). Estimation of southern resident killer whale exposure to exhaust emissions from whale-watching vessels and potential adverse health effects and toxicity thresholds. Marine Pollution Bulletin 62(4), 792–805.CrossRefGoogle ScholarPubMed
Laist, D.W., Knowlton, A.R., Mead, J.G., Collet, A.S. & Podesta, M. (2001). Collisions between ships and whales. Marine Mammal Science 17, 35–75.CrossRefGoogle Scholar
Lien, J. (2001). The conservation basis for the regulation of whale watching in Canada by the Department of Fisheries and Oceans: A precautionary approach. Can. Tech. Rep. Fish. Aquat. Sci. 2363: vi + 38 p.
Lusseau, D. (2003). The effects of tour boats on the behavior of bottlenose dolphins: Using Markov chains to model anthropogenic impacts.Conservation Biology 17, 1785–1793.CrossRefGoogle Scholar
Lusseau, D. (2004). The hidden cost of tourism: Detecting long-term effects of tourism using behavioral information. Ecology and Society 9(1), 2 [online] URL: .CrossRefGoogle Scholar
Lusseau, D. & Bejder, L. (2007). The long-term consequences of short-term responses to disturbance: Experiences from whalewatching impact assessment. International Journal of Comparative Psychology (Special Issue) 20, 228–236.Google Scholar
Lusseau, D., Bain, D.E., Williams, R. & Smith, J.C. (2009). Vessel traffic disrupts the foraging behavior of southern resident killer whales Orcinus orca. Endangered Species Research 6, 211–221.CrossRefGoogle Scholar
May, R.M. (1980). Mathematical models in whaling and fisheries management. In Oster, G.F. (Ed.), Some Mathematical Questions in Biology, Vol. 13. Providence, RI: American Mathematical Society.Google Scholar
Miller, P.J.O. (2000). Maintaining contact: Design and use of acoustic signals in killer whales, Orcinus orca. PhD dissertation (unpublished). MIT-WHOI.CrossRef
Morton, A.B. & Symonds, H.K. (2002). Displacement of Orcinus orca (L.) by high amplitude sound in British Columbia, Canada. ICES Journal of Marine Science 59, 71–80.CrossRef
Nachtigall, P.E., Pawloski, J.L. & Au, W.W.L. (2003). Temporary threshold shifts and recovery following noise exposure in the Atlantic bottlenosed dolphin (Tursiops truncatus). Journal of the Acoustical Society of America 113, 3425–3429.CrossRefGoogle ScholarPubMed
National Research Council. (2003). Ocean Noise and Marine Mammals. Washington, DC: National Academies Press. 192 pp.Google Scholar
National Research Council. (2005). Marine Mammal Populations and Ocean Noise: Determining when noise causes biologically significant effects. Washington, DC: National Academies Press.Google Scholar
Nichol, L.M. & Shackleton, D.M. (1996). Seasonal movements and foraging behaviour of northern resident killer whales (Orcinus orca) in relation to the inshore distribution of salmon (Oncorhynchus spp) in British Columbia. Canadian Journal of Zoology 74, 983–991.CrossRefGoogle Scholar
Olesiuk, P.F. (2009). An assessment of population trends and abundance of harbour seals (Phoca vitulina) in British Columbia. Canadian Science Advisory Secretariat Research Document 2009/105, 163 pp. Available from .
Olesiuk, P.F., Bigg, M.A. & Ellis, G.M. (1990). Life history and population dynamics of resident killer whales (Orcinus orca) in the coastal waters of British Columbia and Washington State. International Whaling Commission Special Issue 12, 209–243.Google Scholar
Olesiuk, P.F., Nichol, L.M., Sowden, M.J. & Ford, J.K.B. (2002). Effect of the sound generated by an acoustic harassment device on the relative abundance and distribution of harbor porpoises (Phocoena phocoena) in retreat passage, British Columbia. Marine Mammal Science 18, 843–862.CrossRefGoogle Scholar
Olesiuk, P.F., Ellis, G.M. & Ford, J.K.B. (2005). Life history and population dynamics of northern resident killer whales (Orcinus orca) in British Columbia. Canadian Science Advisory Secretariat Research Document 2005/045. .
Osborne, R.W. (1999). A historical ecology of Salish Sea ‘Resident’ killer whales (Orcinus orca): With implications for management. PhD dissertation, University of Victoria, Victoria, BC.
Raverty, S., Zabek, E., Schroeder, J.P., Wood, R., Bain, D.E. & Cameron, C.E. (2007). Preliminary investigation into the microbial culture and molecular screening of exhaled breaths of southern resident killer whales (Orcinus sp) and pathogen screening of the sea-surface microlayer (SML) and sub-surface water samples in Washington State. Poster presented to the Int. Assoc. Aquatic Anim. Med. Conf.
Reddy, M.L., Reif, J.S., Bachand, A. & Ridgway, S.H. (2001). Opportunities for using Navy marine mammals to explore associations between organochlorine contaminants and unfavorable effects on reproduction. Science of the Total Enivronment 274, 171–182.CrossRefGoogle ScholarPubMed
Richardson, W.J., Finley, K.J., Miller, G.W., Davis, R.A. & Koski, W.R. (1995a). Feeding, social and migration behavior of bowhead whales, Balaena mysticetus, in Baffin Bay vs. the Beaufort Sea – Regions with different amounts of human activity. Marine Mammal Science 11, 1–45.CrossRefGoogle Scholar
Richardson, W.J., Greene, Jr, C.R., Malme, C.I. & Thomson, D.H. (1995b). Marine Mammals and Noise. San Diego, CA: Academic Press.Google Scholar
Ritter, F. (2003). Interactions of Cetaceans with Whale Watching Boats – Implications for the Management of Whale Watching Tourism. Berlin: M.E.E.R. e.V., 91 pp.Google Scholar
Rosalind, M.R., Parks, S.E., Hunt, K.E., et al. (2011). Evidence that ship noise increases stress in right whales. Proceedings of the Royal Society B 279, 2363–2368.Google Scholar
Ross, P.S., Ellis, G.M., Ikonomou, M.G., Barrett-Lennard, L.G. & Addison, R.F. (2000). High PCB concentrations in free-ranging Pacific killer whales, Orcinus orca: Effects of age, sex and dietary preference. Marine Pollution Bulletin 40, 504–515.CrossRefGoogle Scholar
Schevill, W. & Watkins, W. (1966). Sound structure and directionality in Orcinus (killer whale). Zoologica 51, 71–76.
Szymanski, M.D., Supin, A.Ya., Bain, D.E. & Henry, K.R. (1998). Killer whale (Orcinus orca) auditory evoked potentials to rhythmic clicks. Marine Mammal Science 14, 676–691.CrossRef
Szymanski, M.D., Bain, D.E., Kiehl, K., Henry, K.R., Pennington, S. & Wong, S. (1999). Killer whale (Orcinus orca) hearing: Auditory brainstem response and behavioral audiograms. Journal of the Acoustical Society of America 106, 1134–1141.CrossRefGoogle ScholarPubMed
Taylor, B.L., Wade, P.R., DeMaster, D.P. & Barlow, J. (2000). Incorporating uncertainty into management models for marine mammals. Conservation Biology 14, 1243–1252.CrossRefGoogle Scholar
Wade, P. & Angliss, R. (1997). Guidelines for assessing marine mammal stocks: The GAMMS Workshop. NOAA Technical Memorandum. NMFS-OPR-12. 65 pp. Available from Alaska Fisheries Science Center, 7600 Sand Point Way, NE, Seattle, WA 98115, USA.
Waite, J.M. (1988). Alloparental care in killer whales (Orcinus orca). Masters thesis (unpublished). University of California at Santa Cruz, Santa Cruz, CA.
Ward, E.J., Holmes, E.E. & Balcomb, K.C. (2009). Quantifying the effects of prey abundance on killer whale reproduction. Journal of Applied Ecology 46, 632–640.CrossRefGoogle Scholar
Washington Department of Fisheries, Washington Department of Wildlife and Western Washington Treaty Indian Tribes. (1993). 1992 Washington State Salmon and steelhead stock inventory. Available from Washington Department of Fish and Wildlife, Fish Program, 600 Capitol Way North, Olympia, WA 98501–1091, USA.
Wasser, S.K, Azkarate, J.C., Booth, R.K., et al. (2010). Non-invasive measurement of thyroid hormone in feces of a diverse array of avian and mammalian species. General and Comparative Endocrinology 168, 1–7.CrossRefGoogle ScholarPubMed
Wells, R.S. & Scott, M.D. (1997). Seasonal incidence of boat strikes on bottlenose dolphins near Sarasota, FL. Marine Mammal Science 13(3), 475–480.CrossRefGoogle Scholar
Williams, R. & Ashe, E. (2007). Killer whale evasive tactics vary with boat number. Journal of Zoology 272, 390–397.CrossRefGoogle Scholar
Williams, R. & Noren, D. (2009). Swimming speed, respiration rate and estimated cost of transport in killer whales. Marine Mammal Science 25, 327–350.CrossRefGoogle Scholar
Williams, R., Bain, D.E. & Trites, A.W. (2002a). Behavioural responses of killer whales (Orcinus orca) to whale-watching boats: Opportunistic observations and experimental approaches. Journal of Zoology (London) 256, 255–270.CrossRefGoogle Scholar
Williams, R., Bain, D.E., Trites, A.W. & Ford, J.K.B. (2002b). Behavioural responses of male killer whales to a ‘leapfrogging’ vessel. Journal of Cetacean Research and Management 4, 305–310.Google Scholar
Williams, R., Lusseau, D. & Hammond, P. (2006). Estimating relative energetic costs of human disturbance to killer whales (Orcinus orca). Biological Conservation 133, 301–311.CrossRefGoogle Scholar
Williams, R., Bain, D.E., Smith, J.C. & Lusseau, D. (2009a). Effects of vessels on behaviour patterns of individual southern resident killer whales Orcinus orca. Endangered Species Research 6, 199–209.CrossRefGoogle Scholar
Williams, R., Lusseau, D. & Hammond, P.S. (2009b). The role of social aggregations and protected areas in killer whale conservation: The mixed blessing of critical habitat. Biological Conservation 142, 709–719.CrossRefGoogle Scholar
Ylitalo, G.M., Matkin, C.O., Buzitis, J., et al. (2001). Influence of life-history parameters on organochlorine concentrations in free-ranging killer whales (Orcinus orca) from Prince William Sound, AK. Science of the Total Environment 281, 183–203.CrossRef

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