Abstract
The feeding ecology of predators can have a profound effect on their life history and behaviour. The killer whale—the apex marine predator—has a cosmopolitan distribution throughout the world’s oceans. Globally, it is a generalist predator with a diverse diet, but regionally, different socially and genetically isolated killer whale populations can have highly specialized foraging strategies involving only a few types of prey. In the eastern North Pacific, the three sympatric killer whale lineages have distinct dietary specializations: one feeds primarily on marine mammals, another on salmon, and the third appears to specialize on sharks. These ecological specializations are associated with distinct patterns of seasonal distribution, group size, social organization, foraging behavior, and acoustic activity. Divergent foraging strategies may have played a major role in the social isolation and genetic divergence of killer whale populations.
Keywords
- Apex predator
- Feeding ecology
- Orcinus orca

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References
Au WWL, Ford JKB, Horne JK, Newman Allman KA (2004) Echolocation signals of free-ranging killer whales (Orcinus orca) and modeling of foraging for Chinook salmon (Oncorhynchus tshawytscha). Acoust Soc Am 115(2):901–909
Baird RW, Abrams PA, Dill LM (1992) Possible indirect interactions between transient and resident killer whales: implications for the evolution of foraging specializations in the genus Orcinus. Oecologia, 89:125–132
Baird RW, Dill LM (1995) Occurrence and behavior of transient killer whales: seasonal and pod-specific variability, foraging, and prey handling. Can J Zool 73:1300–1311
Baird RW, Dill LM (1996) Ecological and social determinants of group size in transient killer whales. Behav Ecol 7:408–416
Baird RW, Hanson MB, Dill LM (2005a) Factors influencing the diving behaviour of fish-eating killer whales: sex differences and diel and interannual variation in diving rates. Can J Zool 83:257–267
Baird RW, McSweeney DJ, Bane C, Barlow J, Salden DR, Antoine LK, LeDuc RG, Webster DL (2005b) Killer whales in Hawaiian waters: information on population identity and feeding habits. Pac Sci 60:523–530
Baird RW, Whitehead H (2000) Social organisation of mammal eating killer whales: group stability and dispersal patterns. Can J Zool 78:2096–2105
Barrett-Lennard LG (2000) Population structure and mating systems of northeastern pacific killer whales. Ph.D. Thesis, University of British Columbia, Vancouver
Barrett-Lennard LG, Ford JKB, Heise KA (1996) The mixed blessing of echolocation: differences in sonar use by fish-eating and mammal-eating killer whales. Anim Behav 51:553–565
Barrett-Lennard LG, Heise K (2006) The natural history and ecology of killer whales. In: Estes JA, DeMaster DP, Doak DF, Williams TM, Brownell RL (eds) Whales, whaling and ocean ecosystems. University of California Press, Berkeley, pp 163–173
Barrett-Lennard LG, Matkin CO, Ellifrit DK, Durban J, Mazzuca L (2005) Black and white versus gray: estimating kill rates, consumption rates, and population-level impacts of transient killer whales feeding on gray whales. Abstract, 16th Biennial Conference on the Biology of Marine Mammals, San Diego, 12–16 December 2005, p 26
Bigg MA (1982) An assessment of killer whale (Orcinus orca) stocks off Vancouver Island, British Columbia. Rep Int Whal Comm 32:655–666
Bigg MA, Ellis GM, Ford JKB, Balcomb KC (1987) Killer whales: a study of their identification, genealogy and natural history in British Columbia and Washington State. Phantom Press, Nanaimo
Bigg MA, Ford JKB, Ellis GM (1985) Two sympatric forms of killer whales off British Columbia and Washington. In: Sixth Biennial Conference on the Biology of Marine Mammals, Vancouver, Canada, November 1985
Bigg MA, MacAskie IB, Ellis G (1976) Abundance and movements of killer whales off eastern and southern Vancouver Island with comments on management. Unpublished report, Arctic Biological Station, Department of Fisheries and Environment, Ste Anne-de-Bellevue
Bigg MA, Olesiuk PF, Ellis GM, Ford JKB, Balcomb KC III (1990) Social organization and genealogy of resident killer whales (Orcinus orca) in the coastal waters of British Columbia and Washington State. Rep Int Whal Comm Spec Issue 12:383–405
Black NA, Schulman-Janiger A, Ternullo RL, Guerrero-Ruiz M (1997) Killer whales of California and western Mexico: a catalog of photo-identified individuals. NOAA Technical Memorandum NOAA-TM-NMFS-SWFSC-247
Candy JR, Quinn TP (1999) Behaviour of adult Chinook salmon (Oncorhynchus tshawytscha) in British Columbia coastal waters determined from ultrasonic telemetry. Can J Zool 77:1161–1169
Dahlheim ME, Schulman-Janiger A, Black N, Ternullo R, Ellifrit D, Balcomb KC (2008) Eastern temperate North Pacific offshore killer whales (Orcinus orca): occurrence, movements, and insights into feeding ecology. Mar Mamm Sci 24:719–729
Deecke VB, Ford JKB, Slater P (2005) The vocal behaviour of mammal-eating killer whales: communicating with costly calls. Anim Behav 69:395–405
Deecke VB, Slater P, Ford JKB (2002) Selective habituation shapes acoustic predator recognition in harbour seals. Nature (Lond) 420:171–173
Dieckmann U, Doebeli M (1999) On the origin of species by sympatric speciation. Nature (Lond) 400:354–357
Ellis GM, Ford JKB, Towers JR (2007) Northern resident killer whales in British Columbia: photo-identification catalogue 2007. Pacific Biological Station, Fisheries and Oceans Canada, Nanaimo. http://www.pac.dfo-mpo.gc.ca/sci/sa/cetacean/default_e.htm
Estes JA, Tinker MT, Williams TM, Doak DF (1998) Killer whale predation on sea otters linking oceanic and nearshore ecosystems. Science 282:473–476
Ford JKB (1984) Call traditions and dialects of killer whales (Orcinus orca) in British Columbia. Ph.D. Dissertation, University of British Columbia, Vancouver
Ford JKB (1989) Acoustic behaviour of resident killer whales (Orcinus orca) off Vancouver Island, British Columbia. Can J Zool 67:727–745
Ford JKB (1991) Vocal traditions among resident killer whales (Orcinus orca) in coastal waters of British Columbia. Can J Zool 69:1454–1483
Ford JKB (2002) Killer whale Orcinus orca. In: Perrin WF, Wursig B, Thewissen JGM (eds) The encyclopedia of marine mammals. Academic, San Diego, pp 669–676
Ford JKB (2006) An assessment of critical habitats of resident killer whales in waters off the Pacific coast of Canada. Fisheries and Oceans Canada, Canadian Science Advisory Secretariat, Ottawa. http://www.dfo-mpo.gc.ca/csas/
Ford JKB (2009) Killer whales Orcinus orca. In: Perrin WF, Würsig B, Thewissen JGM (eds) The encyclopedia of marine mammals, 2nd edn. Academic, San Diego, pp 650–657
Ford JKB, Ellis FM, Nichol LM (1992) Killer whales of the Queen Charlotte Islands: a preliminary study of the abundance, distribution, and population identity of Orcinus orca in the waters of Haida Gwaii. Unpublished report, South Moresby/Haida Gwaii Haanas National Parks Reserve, Canadian Parks Service
Ford JKB, Ellis GM (1999) Transients: mammal-hunting killer whales of British Columbia, Washington, and Southeastern Alaska. UBC Press, Vancouver
Ford JKB, Ellis GM (2006) Selective foraging by fish-eating killer whales Orcinus orca in British Columbia. Mar Ecol Prog Ser 316:185–199
Ford JKB, Ellis GM, Balcomb KC (2000) Killer whales: the natural history and genealogy of Orcinus orca in the waters of British Columbia and Washington. UBC and University of Washington Press, Vancouver
Ford JKB, Ellis GM, Barrett-Lennard LG, Morton AB, Palm RS, Balcomb KC III (1998) Dietary specialization in two sympatric populations of killer whales (Orcinus orca) in coastal British Columbia and adjacent waters. Can J Zool 76:1456–1471
Ford JKB, Ellis GM, Durban JW (2007) An assessment of the potential for recovery of West Coast transient killer whales using coastal waters of British Columbia. Fisheries and Oceans Canada, Canadian Science Advisory Secretariat, Ottawa. http://www.dfo-mpo.gc.ca/csas/
Ford JKB, Ellis GM, Matkin DR, Balcomb KC, Briggs D, Morton AB (2005) Killer whale attacks on minke whales: prey capture and antipredator tactics. Mar Mamm Sci 21:603–618
Ford JKB, Ellis GM, Matkin CO, Wetklo MH, Barrett-Lennard LG, Withler RE (2011) Shark predation and tooth wear in a population of northeastern Pacific killer whales. Aquat Biol 11:213–224
Ford JKB, Reeves RR (2008) Fight or flight: antipredator strategies of baleen whales. Mammal Review, 38:50–86
Forney KA, Wade PR (2006) Worldwide distribution and abundance of killer whales. In: Estes JA, Demaster DP, Doak DF, Williams TM, Brownell RL Jr (eds) Whales, whaling, and ocean ecosystems. University of California Press, Berkeley, pp 145–173
Futuyma DJ, Moreno G (1988) The evolution of ecological specialization. Annu Rev Ecol Syst 19:207–233
Guinet C (1990) Sympatrie des deux categories d’orques dans le detroit de Johnstone, Columbie Britannique. Rev Ecol (Terre Vie) 45:25–34
Guinet C (1992) Hunting behaviour of killer whales (Orcinus orca) around Crozet Islands. Can J Zool 70:1656–1667
Guinet C, Domenici P, de Stephanis R, Barrett-Lennard LG, Ford JKB, Verborgh P (2007) Killer whale predation on bluefin tuna: exploring the hypothesis of the endurance-exhaustion technique. Mar Ecol Prog Ser 347:111–119
Hauser DDW, Logsdon MG, Holmes EE, VanBlaricom GR, Osborne RW (2007) Summer distribution patterns of southern resident killer whales Orcinus orca: core areas and spatial segregation of social groups. Mar Ecol Prog Ser 351:301–310
Heimlich-Boran J (1986) Fishery correlations with the occurrence of killer whales in greater Puget Sound. In: Kirkevold BC, Lockard JS (eds) Behavioral biology of killer whales. Liss, New York, pp 113–131
Heimlich-Boran JR (1988) Behavioural ecology of killer whales (Orcinus orca) in the Pacific Northwest. Can J Zool 66:565–578
Heise K, Barrett-Lennard LG, Saulitis E, Matkin C, Bain D (2003) Examining the evidence for killer whale predation on Steller sea lions in British Columbia and Alaska. Aquat Mamm 29:325–334
Herman DP, Burrows DG, Wade PR, Durban JW, Matkin CO, LeDuc R, Barrett-Lennard LG, Krahn MM (2005) Feeding ecology of eastern North Pacific killer whales Orcinus orca from fatty acid, stable isotope, and organochlorine analyses of blubber biopsies. Mar Ecol Prog Ser 302:275–291
Hoelzel AR (1991) Killer whale predation on marine mammals at Punta Norte, Argentina: food sharing, provisioning and foraging strategy. Behav Ecol Sociobiol 29:197–204
Hoelzel AR, Dahlheim ME, Stern SJ (1998) Low genetic variation among killer whales (Orcinus orca) in the eastern North Pacific, and differentiation between foraging specialists. J Hered 89:121–128
Hoelzel AR, Natoli A, Dahlheim ME, Olavarria C, Baird RW, Black NA (2002) Low worldwide genetic diversity in the killer whale: implications for demographic history. Proc R Soc Lond Ser B 269:1467–1473
Jefferson TA, Stacey PF, Baird RW (1991) A review of killer whale interactions with other marine mammals: predation to co-existence. Mamm Rev 21:151–180
Jones IM (2006) A northeast Pacific offshore killer whale (Orcinus orca) feeding on a Pacific halibut (Hippoglossus stenolepis). Mar Mamm Sci 22:198–200
Krahn MM, Herman DP, Matkin CO, Durban JW, Barrett-Lennard L, Burrows DG, Dahlheim ME, Black N, Leduc RG, Wade PR (2007) Use of chemical tracers in assessing the diet and foraging regions of eastern North Pacific killer whales. Mar Environ Res 63:91–114
Krahn MM, Pitman RL, Burrows DG, Herman DP, Pearce RW (2008) Use of chemical tracers to assess diet and persistent organic pollutants in Antarctic type C killer whales. Mar Mamm Sci 24:643–663
LeDuc RG, Robertson KM, Pitman RL (2008) Mitochondrial sequence divergence among Antarctic killer whale ecotypes is consistent with multiple species. Biol Lett 4:426–429
Lopez JC, Lopez D (1985) Killer whales (Orcinus orca) of Patagonia, and their behavior of intentional stranding while hunting nearshore. J Mammal 66:181–183
Martinez DR, Klinghammer E (1970) The behavior of the whales, Orcinus orca: a review of the literature. Z Tierpsychol 27:828–839
Matkin C, Ellis G, Saulitis E, Barrett-Lennard L, Matkin D (1999) Killer whales of Southern Alaska. North Gulf Oceanic Society, Homer
Matkin CO, Barrett-Lennard LG, Yurk H, Ellifrit D, Trites AW (2007a) Ecotypic variation and predatory behavior among killer whales (Orcinus orca) off the eastern Aleutian Islands. Alaska Fish Bull 105:74–87
Matkin DR, Straley JM, Gabriele CM (2007b) Killer whale feeding ecology and non-predatory interactions with other marine mammals in the Glacier Bay region of Alaska. In: Piatt JF, Gende SM (eds) Proceedings of the Fourth Glacier Bay Science Symposium, pp 155–158. http://pubs.usgs.gov/sir/2007/5047/
McKinnon JS, Mori S, Blackman BK, David U, Kingsley DM, Jamieson L, Chou J, Schluter D (2004) Evidence for ecology’s role in speciation. Nature (Lond) 429:294–298
Miller PJO, Shapiro AD, Tyack PL, Solow AR (2004) Call-type matching in vocal exchanges of free-ranging resident killer whales, Orcinus orca. Anim Behav 67:1099–1107
Miller PJO (2006) Diversity in sound pressure levels and estimated active space of resident killer whale vocalizations. J Comp Physiol A 192:449–459
Mizroch SA, Rice DW (2006) Have North Pacific killer whales switched prey species in response to depletion of the great whale populations? Mar Ecol Prog Ser 310:235–246
Morin PA, Leduc RG, Robertson KM, Hedrick NM, Perrin WF, Etnier M, Wade P, Taylor BL (2006) Genetic analysis of killer whale (Orcinus orca) historical bone and tooth samples to identify western U.S. ecotypes. Mar Mamm Sci 22:897–909
Morin PA, Archer FI, Foote AD, Vilstrup J, Allen EE, Wade P, Durban J, Parsons KL, Pitman RL, Li L, Bouffard P, Abel Nielsen SC, Rasmussen M, Willerslev E, Gilbert MTP, Harkins T (2010) Complete mitochondrial genome phylogeographic analysis of killer whales (Orcinus orca) indicates multiple species. Genome Res 20:908–916
Morton AB (1990) A quantitative comparison of the behavior of resident and transient forms of the killer whale off the central British Columbia coast. Rep Int Whal Comm Spec Issue 12:245–248
Nichol LM, Shackleton DM (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. Can J Zool 74:983–991
Olesiuk PF, Ellis GM, Ford JKB (2005) Life history and population dynamics of northern resident killer whales (Orcinus orca) in British Columbia. Canadian Science Advisory Secretariat, Fisheries and Oceans, Canada. http://www.dfo-mpo.gc.ca/csas/
Osborne RW (1999) A historical ecology of Salish Sea “resident” killer whales (Orcinus orca), with implications for management. Ph.D. Thesis, University of Victoria, British Columbia
Pike GC, MacAskie IB (1969) Marine mammals of British Columbia. Fish Res Board Can Bull 71:1–54
Pitman RL, Ensor P (2003) Three forms of killer whales (Orcinus orca) in Antarctic waters. J Cetacean Res Manag 5:131–139
Pitman RL, Perryman WL, LeRoi D, Eilers E (2007) A dwarf form of killer whale in Antarctica. J Mammal 88:43–48
Reeves RR, Notarbartolo Di Sciara G (2006) The status and distribution of cetaceans in the Black Sea and Mediterranean Sea. IUCN Centre for Mediterranean Cooperation, Malaga, Spain. http://cmsdata.iucn.org/downloads/status_distr_cet_blac_med.pdf
Robinson BW, Wilson DS, Shea GO (1996) Trade-offs of ecological specialization: an intraspecific comparison of pumpkinseed sunfish phenotypes. Ecology 77:170–178
Saulitis E, Matkin C, Barrett-Lennard L, Heise K, Ellis G (2000) Foraging strategies of sympatric killer whale (Orcinus orca) populations in Prince William Sound. Alaska Mar Mamm Sci 16:94–109
Schluter D (2001) Ecology and the origin of species. Trends Ecol Evol 16(7):372–380
Similä T (1997) Behavioral ecology of killer whales in Northern Norway. Norwegian College of Fisheries Science. Ph.D. Thesis, University of Tromso, Tromso
Similä T, Ugarte F (1993) Surface and underwater observations of cooperatively feeding killer whales in northern Norway. Can J Zool 71:1494–1499
Simon M, Ugarte F, Wahlberg M, Miller LA (2005) Icelandic killer whales Orcinus orca use a pulsed call suitable for manipulating the schooling behaviour of herring Clupea harengus. Bioacoustics 16:57–74
Springer AM, Estes JA, van Vliet GB, Williams TM, Doak DF, Danner EM, Forney KA, Phister B (2003) Sequential megafaunal collapse in the North Pacific Ocean: an ongoing legacy of industrial whaling? Proc Natl Acad Sci USA 100:12223–12228
Steiger GH, Calambokidis J, Straley JM, Herman LM, Cerchio S, Salden DR, Urbán J, Jacobsen RJK, von Ziegesar O, Balcomb KC, Gabriele CM, Dahlheim ME, Uchida S, Ford JKB, Ladron de Guevara P, Yamaguchi M, Barlow J (2008) Geographic variation in killer whale attacks on humpback whales in the North Pacific: implications for predation pressure. Endang Species Res 4:247–256
Stephens DW, Krebs JR (1986) Foraging theory. Princeton University Press, Princeton
Stevens TA, Duffiled DA, Asper ED, Hewlett KG, Bolz A, Gage LJ, Bossart GD (1989) Preliminary findings of restriction fragment differences in mitochondrial DNA among killer whales. Can J Zool 67:2592–2595
Strager H (1995) Pod-specific call repertoires and compound calls of killer whales, Orcinus orca Linnaeus, 1758, in the waters of northern Norway. Can J Zool 73:1037–1047
Temte JL, Bigg MA, Wiig O (1991) Clines revisited: the timing of pupping in the harbour seal (Phoca vitulina). J Zool (Lond) 224:617–632
Ternullo R, Black N (2002) Predation behavior of transient killer whales in Monterey Bay, California. In: Fourth International Orca Symposium and Workshop, CEBC-CNRS, France, pp 156–159
Trites AW, Deecke VB, Gregr EJ, Ford JKB, Olesiuk PF (2007) Killer whales, whaling, and sequential megafaunal collapse in the North Pacific: a comparative analysis of the dynamics of marine mammals in Alaska and British Columbia following commercial whaling. Mar Mamm Sci 23:751–765
Via S (2001) Sympatric speciation in animals: the ugly duckling grows up. Trends Ecol Evol 16:381–390
Visser IN, Smith TG, Bullock ID, Green GD, Carlsson OGL, Imberti S (2008) Antarctic peninsula killer whales (Orcinus orca) hunt seals and a penguin on floating ice. Mar Mamm Sci 24:225–234
Wade PR, Barrett-Lennard LG, Black NA, Burkanov VN, Burdin AM, Calambokidis J, Cerchio S, Dahlheim ME, Ford JKB, Friday NA, Fritz LW, Jacobsen JK, Loughlin TR, Matkin CO, Matkin DR, McCluskey SM, Mehta AV, Mizroch SA, Muto MM, Rice DW, Robe P, Clapham P (2007) Killer whales and marine mammal trends in the North Pacific: a re-examination of evidence for sequential megafauna collapse and the prey-switching hypothesis. Mar Mamm Sci 23:766–802
Yurk H, Barrett-Lennard LG, Ford JKB, Matkins C (2002) Cultural transmission within maternal lineages: vocal clans in resident killer whales in southern Alaska. Anim Behav 63:1103–1119
Acknowledgments
We thank R. Baird, C. Chapman, V. Iriarte, C. Matkin, and J. Watson for helpful comments on earlier drafts of this chapter, and M. Malleson and J. Towers for kindly allowing us to use their photographs.
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Ford, J.K.B., Ellis, G.M. (2014). You Are What You Eat: Foraging Specializations and Their Influence on the Social Organization and Behavior of Killer Whales. In: Yamagiwa, J., Karczmarski, L. (eds) Primates and Cetaceans. Primatology Monographs. Springer, Tokyo. https://doi.org/10.1007/978-4-431-54523-1_4
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