Mammalian Biology

, Volume 97, Issue 1, pp 72–79 | Cite as

The summer diet of fish-eating killer whales in the Avacha Gulf of Kamchatka: Are there any preferences?

  • Evgeniia V. VolkovaEmail author
  • Tatiana V. Ivkovich
  • Marina V. Shitova
  • Ekaterina N. Chernyaeva
  • Tatiana V. Malinina
  • Sofia S. Okorokova
  • Alexandr M. Burdin
  • Erich Hoyt
Original investigation


Dietary specialization has led to the origin of several ecotypes of killer whales (Orcinus orca). Comparing the feeding habits of different populations within an ecotype is valuable for understanding the evolution of foraging strategies. In this study we investigated the diet of fish-eating killer whales in the Northwest Pacific Ocean (Avacha Gulf, Kamchatka) and compared it to the published data on Northeast Pacific fish-eating killer whales. We collected prey fragments (N = 141) after hunting events in the summer seasons of 2011–2016 and determined the species of prey. Killer whales fed mainly on various species of Pacific salmon in Avacha Gulf. Coho salmon (Oncorhynchus kisutch) (56% of samples) and chum salmon (O. keta) (31% of samples) predominated in the diet. The results are comparable to the peculiarities of the fish-eating killer whale diet in the Northeast Pacific where killer whales prefer Chinook salmon (O. tshawytscha) most of the summer but shift toward coho and chum in late summer. Avacha Gulf could be an area where killer whales come to hunt coho similar to Prince William Sound in Alaska where killer whales prefer coho to other salmon species. We found significant differences in the diet between sex-age categories. Females (N = 42) and juveniles (N = 15) were more selective and hunted mainly coho while males (N = 53) hunted chum equally to coho. Like Chinook in British Columbia, coho in Kamchatka has a prolonged spawning period. Coho is the most nutritious after Chinook. Coho’s prolonged abundance and nutritional value probably make it preferable to killer whales especially females.


Killer whales Foraging strategy Foraging preferences Prey choice Foraging specialization 


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  1. Agresti, A., 2002. Categorical Data Analysis, 2nd edition. John Wiley & Sons, New York.CrossRefGoogle Scholar
  2. Altschul, S.F., Gish, W., Miller, W., Myers, E.W., Lipman, D.J., 1990. Basic local alignment search tool. J. Mol. Biol., 215, 403–410, Scholar
  3. Antonov, N.P., 2011. Biology, Population Dynamics and Sustainable Use of Fish in Kamchatka and Adjacent Marine Areas [PhD Thesis]. Russian Federal Research Institute of Fisheries and Oceanography, Moscow, Russia [In Russian].Google Scholar
  4. Araújo, M.S., Bolnick, D.I., Layman, CA., 2011. The ecological causes of individual specialisation. Ecol. Lett., 14, 948–958, Scholar
  5. Baird, R.W., Hanson, M.B., Dill, L.M., 2005. Factors influencing the diving behaviour offish-eating killer whales: sex differences and diel and interannual variation in diving rates. Can. J. Zool., 83, 257–267, Scholar
  6. Baird, R.W., McSweeney, D.J., Bane, C., Barlow, J., Salden, D.R., Antoine, L.K., LeDuc, R.G., Webster, D.L., 2006. Killer whales in hawaiian waters: information on population identity and feeding habits. Pacific Sci., 60, 523–530, Scholar
  7. Beck, CA., Iverson, S.J., Bowen, W.D., Blanchard, W., 2007. Sex differences in grey seal diet reflect seasonal variation in foraging behaviour and reproductive expenditure: evidence from quantitative fatty acid signature analysis. J. Anim. Ecol., 76, 490–502, Scholar
  8. Beerman, A., Ashe, E., Preedy, K., Williams, R., 2016. Sexual segregation when foraging in an extremely social killer whale population. Behav. Ecol. Sociobiol., 70, 189–198, Scholar
  9. Belonovich, O., Burkanov, V., 2012. Killer whale (Orcinus orca) depredation on the Greenland halibut (Reinhardtius hippoglossoides) long-line fishery in the Sea of Okhotsk. VII International Conference «Marine Mammals of the Holarctic», 86–90.Google Scholar
  10. Best, P.B., 1999. Food and feeding of sperm whales Physeter macrocephalus off the west coast of South Africa. South African J. Mar. Sci. ISSN, 21, 393–413, Scholar
  11. Bigg, M.A., Olesiuk, P.F., Ellis, G.M., Ford, J.K.B., Balcomb, K.C., 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., 12, 383–405 (special issue).Google Scholar
  12. Bolaños-Jiménez, J., Mignucci-Giannoni, A.A., Blumenthal, J., Bogomolni, A., Casas, J.J., Henríquez, A., Iñíguez Bessega, M., Khan, J., Landrau-Giovannetti, N., Rinaldi, C., Rinaldi, R., Rodríguez-Ferrer, G., Sutty, L., Ward, N., Luksenburg, J.A., 2014. Distribution, feeding habits and morphology of killer whales Orcinus orca in the Caribbean Sea. Mamm. Rev., Scholar
  13. Bolnick, D.I., Svanbäck, R., Fordyce, J.A., Yang, L.H., Davis, J.M., Hulsey, CD., Forister, M.L., 2003. The ecology of individuals: incidence and implications of individual specialization. Am. Nat., 161, 1–28, Scholar
  14. Burdin, a M., Hoyt, E., Filatova, O., Ivkovich, T., Tarasyan, K., Sato, H., 2007. Status of killer whales (Orcinus orca) in Eastern Kamchatka, Russia, based on photo-identification and acoustic studies. Preliminary results. IWC Sci. Comm., 11.Google Scholar
  15. Capella, J.J., Abramson, J.Z., Vilina, Y.A., Gibbons, J., 2014. Observations of killer whales (Orcinus orca) in the fjords of Chilean Patagonia. Polar Biol., 37, 1533–1539, Scholar
  16. Clutton-Brock, T.H., 1977. Some Aspects of Intraspecific Variation in Feeding and Ranging Behaviour in Primates, Primate Ecology: Studies of Feeding and Ranging Behaviour in Lemurs, Monkey and Apes. ACADEMIC PRESS INC. (LONDON) LTD, Scholar
  17. Clutton-Brock, T.H., Iason, G.R., Guinness, F.E., 1987. Sexual segregation and density-related changes in habitat use in male and female red deer (Cervus elaphus). J. Zool., 211, 275–289, Scholar
  18. Emlen, J.M., 1966. The role of time and energy in food preference. Am. Nat., 100, 611–617, Scholar
  19. Filatova, O.A., Fedutin, I.D., Burdin, A.M., Hoyt, E., 2007. The structure of the discrete call repertoire of killer whales Orcinus Orca from Southeast Kamchatka. Bioacoustics, 16, 261–280, Scholar
  20. Filatova, O.A., Borisova, E.A., Shpak, O.V., Meschersky, I.G., Tiunov, A.V., Goncharov, A.A., Fedutin, I.D., Burdin, A.M., 2015. Reproductively isolated ecotypes of killer whales Orcinus orca in the seas of the Russian Far East. Biol. Bull., 42, 674–681, Scholar
  21. Foote, A.D., Newton, J., Piertney, S.B., Willerslev, E., Gilbert, M.T.P., 2009. Ecological, morphological and genetic divergence of sympatric North Atlantic killer whale populations. Mol. Ecol., 18, 5207–5217, Scholar
  22. Ford, J.K.B., Ellis, G.M., 2006. Selective foraging by fish-eating killer whales Orcinus orca in British Columbia. Mar. Ecol. Prog. Ser., 316, 185–199, Scholar
  23. Ford, J.K.B., Ellis, G.M., Barrett-Lennard, L.G., Morton, A.B., Palm, R.S., Balcomb III, K.C., 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, Scholar
  24. Ford, M.J., Hempelmann, J., Hanson, M.B., Ayres, K.L., Baird, R.W., Emmons, CK., Lundin, J.I., Schorr, G.S., Wasser, K., Park, L.K., 2016. Estimation of a killer whale (Orcinus orca) population’s diet using sequencing analysis of DNA from feces. PLoS Biol., 11, 1–14, Scholar
  25. Foster, E.A., Franks, D.W., Morrell, L.J., Balcomb, K.C., Parsons, K.M., van Ginneken, A., Croft, D.P., 2012. Social network correlates of food availability in an endangered population of killer whales, Orcinus orca. Anim. Behav., 83, 731–736, Scholar
  26. Ginnett, T.F., Demment, M.W., 1997. Sex differences in giraffe foraging behavior at two spatial scales. Oecologia, 110, 291–300, Scholar
  27. Groot, C., Margolis, L., 1991. Pacific Salmon Life Histories. UBC press, Vancouver, British Columbia.Google Scholar
  28. Hanson, M.B., Baird, R.W., Ford, J.K.B., Hempelmann-Halos, J., Van Doornik, D.M., Candy, J.R., Emmons, CK., Schorr, G.S., Gisborne, B., Ayres, K.L., Wasser, S.K., Balcomb, K.C., Balcomb-Bartok, K., Sneva, J.G., Ford, M.J., 2010. Species and stock identification of prey consumed by endangered southern resident killer whales in their summer range. Endanger. Species Res., 11, 69–82, Scholar
  29. Hoelzel, A.R., 1993. Foraging behaviour and social group dynamics in Puget Sound killer whales. Anim. Behav. 45 (3), 581–591, Scholar
  30. Ishida, Y., Yano, A., Ban, M., Ogura, M., 2001. Vertical movement of a chum salmon Oncorhynchus keta in the western North Pacific Ocean as determined by a depth-recording archival tag. Fish. Sci., 67, 1030–1035, Scholar
  31. Ivkovich, T., Filatova, O.A., Burdin, A.M., Sato, H., Hoyt, E., 2010. The social organization of resident-type killer whales (Orcinus orca) in Avacha Gulf, Northwest Pacific, as revealed through association patterns and acoustic similarity. Mamm. Biol., 75, 198–210, Scholar
  32. Kamilar, J.M., Pokempner, A.A., 2008. Does body mass dimorphism increase male -female dietary niche separation? A comparative study of primates. Behaviour, 145, 1211–1234, Scholar
  33. Kleiber, M., 1975. Metabolic turnover rate: a physiological meaning of the metabolic rate per unit body weight. J. Theor. Biol., 53, 199–204, Scholar
  34. Kornev, S.I., Belonovich, O., Nikulin, S.V., 2014. Killerwhales (Orcinus orca) and greenland halibut (Reinhardtius hipoglossoides) fishery in the Sea of Okhotsk. Stud. Aquat. Biol. Resour. Kamchatka North-West Pacific, 34, 35–50.Google Scholar
  35. MacArthur, R.H., 1972. The Economics of Consumer Choice, in: Geographical Ecology. Harper & Row, Publishers, New York, pp. 59–70.Google Scholar
  36. MacArthur, R.H., Pianka, E.R., 1966. On optimal use of a patchy environment. Am. Nat., 100, 603–609, Scholar
  37. Miller, P.J.O., Shapiro, A.D., Deecke, V.B., 2010. The diving behaviour of mammal-eating killerwhales (Orcinus orca): variations with ecological not physiological factors. Can. J. Zool., 88, 1103–1112, Scholar
  38. Morin, P.A., Archer, F.I., Foote, A.D., Vilstrup, J., Allen, E.E., Wade, P., Durban, J., Parsons, K., Pitman, R., Li, L., Bouffard, P., Nielsen, S.C.A., Rasmussen, M., Willerslev, E., Gilbert, M.T.P., Harkins, T., 2010. Complete mitochondrial genome phylogeographic analysis of killerwhales (Orcinus orca) indicates multiple species. Genome Res., 20, 908–916, Scholar
  39. Mosher, K.H., 1969. Identification of Pacific Salmon and Steelhead Trout by Scale Characteristics. United States Department of the Interior, Fish and Wildlife Service, Bureau of Commercial Fishes, Washington, D.C.Google Scholar
  40. Nagaylik, M.M., Burdin, A.M., Hoyt, E., 2012. Influence of salmon abundance on the activity offish-eating killerwhales. VII International Conference «Marine Mammals of the Holarctic», 26–29.Google Scholar
  41. Nichol, L.M., Shackleton, D.M., 1996. Seasonal movements and foraging behaviour of resident killerwhales (Orcinus orca) in relation to the inshore distribution of salmon (Oncorhynchus spp.) in British Columbia. Can. J. Zool., 74, 983–991, Scholar
  42. Noren, D.P., 2011. Estimated field metabolic rates and prey requirements of resident killerwhales. Mar. Mammal Sci., 27, 60–77, Scholar
  43. O’Neill, S.M., Ylitalo, G.M., West, J.E., 2014. Energy content of Pacific salmon as prey of northern and southern resident killerwhales. Endanger. Species Res., 25, 265–281, Scholar
  44. Ogura, M., Ishida, Y., 1995. Homing behavior and vertical movements of four species of Pacific salmon (Oncorhynchus spp.) in the central Bering Sea. Can. J. Fish., 5410, 5–7, Scholar
  45. Olesiuk, P.F., Ellis, G.M., Ford, J.K.B., 2005. Life history and population dynamics of northern resident killerwhales (Orcinus orca) in British Columbia. Rep. Int. Whal. Comm. Spec. Issue, 12, 209–243.Google Scholar
  46. Osborne, R., [ Ph.D. Dissertation] 1999. A Historical Ecology of Salish Sea. A Resident KillerWhales (Orcinus orca): With Implications for Management. University of Victoria, Victoria, BC, Canada, pp. 130.Google Scholar
  47. Pianka, E.R., 1974. Resource acquisition and allocation: foraging tactics and feeding efficiency. In: Pianka, E.R. (Ed.), Evolutionary Ecology. Harper & Row, Publishers, New York, pp. 78–83.Google Scholar
  48. Pitman, R.L., Durban, J.W., 2011. Cooperative hunting behavior, prey selectivity and prey handling by pack ice killerwhales (Orcinus orca), type B, in Antarctic Peninsula waters. Mar. Mammal Sci., 28, 16–36, Scholar
  49. Pitman, R., Ensor, P., 2003. Three forms of killerwhales (Orcinus orca) in Antarctic waters. J. Cetacean Res. Manag., 5, 131–139, Scholar
  50. Rasmussen, R.S., Morrissey, M.T., Hebert, P.D., 2009. DNA barcoding of commercially important salmon and trout species (Oncorhynchus and Salmo) from North America. J. Agric. Food Chem. 57 (18), 8379–8385,, Sep 23.CrossRefGoogle Scholar
  51. R-Core-Team., 2015. R: a Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna (Austria).Google Scholar
  52. Reisinger, R.R., Gröcke, D.R., Lübcker, N., McClymont, E.L., Rus Hoelzel, A., Nico De Bruyn, P.J., 2016. Variation in the diet of killer whales Orcinus orca at Marion Island, Southern Ocean. Mar. Ecol. Prog. Ser., 549, 263–274, Scholar
  53. Rendell, L., Whitehead, H., 2001. Culture in whales and dolphins. Behav. Brain Sci. 24 (2), 309–324, Scholar
  54. Riesch, R., Barrett-lennard, L.G., Ellis, G.M., Ford, J.K.B., Deecke, V.B., 2012. Cultural traditions and the evolution of reproductive isolation: ecological speciation in killerwhales? Biol. J. Linn. Soc. 106 (1), 1–17, Scholar
  55. Samarra, F.I.P., Vighi, M., Aguilar, A., Víkingsson, G.A., 2017. Intra-population variation in isotopic niche in herring-eating killerwhales off Iceland. Mar. Ecol. Prog. Ser., 564, 199–210, Scholar
  56. Saulitis, E., Matkin, C., 2000. Foraging Strategies of Sympatric Killer Whale (Orcinus Orca)Populations in Prince William Sound, Alaska. Mar. Mammal Sci., 16, 94–109, Scholar
  57. Schoener, T.W., 1971. Theory of feeding strategies. Annu. Rev. Ecol. Syst., 2, 369–404, Scholar
  58. Shine, R., 1989. Ecological causes forthe evolution of sexual dimorphism: a review of the evidence. Q. Rev. Biol., 64, 419–461, Scholar
  59. Smirnov, A.I., 1975. Biology, Reproduction and Development of Pacific Salmon. Publishing House of Moscow State University, Moscow, Russia [In Russian].Google Scholar
  60. Stephens, D.W., Krebs, J.R., 1986. Foraging Theory. Princeton University Press, Princeton, New JerseyGoogle Scholar
  61. Tarasyan, K.K., Filatova, O.A., Burdin, A.M., Hoyt, E., Sato, H., 2005. Keys forthe status of killer whales in Eastern Kamchatka, Russia: foraging ecology and acoustic behavior. Biosph. Conserv., 6, 73–83, Scholar
  62. Teloni, V., Mark, J.P., Patrick, M.J.O., Peter, M.T., 2008. Shallow food for deep divers: dynamic foraging behavior of male sperm whales in a high latitude habitat. J. Exp. Mar. Bio. Ecol., 354, 119–131, Scholar
  63. Testin, A.I., Pinigin, E.V., Purtov, S.Yu., Mironova, A.M., Burkanov, V.N., 2002. Influence of sea lions and killerwhales on the longline fishery inthe Okhotsk and Bering seas. II International Conference «Marine Mammals of the Holarctic», 2002, 252–254.Google Scholar
  64. Thiemann, G.W., Iverson, S.J., Stirling, I., Obbard, M.E., 2011. Individual patterns of prey selection and dietary specialization in an Arctic marine carnivore. Oikos, 120, 1469–1478, Scholar
  65. Vongraven, D., Bisther, A., 2014. Prey switching by killerwhales inthe north-east Atlantic: observational evidence and experimental insights. J. Mar. Biolog. Assoc. U.K., 94, 1357–1365, Scholar
  66. Ward, R.D., Zemlak, T.S., Innes, B.H., Last, P.R., Hebert, P.D.N., 2005. DNA barcoding Australia’s fish species. Philos. Trans. Biol. Sci., 360, 1847–1857, Scholar
  67. Williams, R., Krkosek, M., Ashe, E., Branch, T.A., Clark, S., Hammond, P.S., Hoyt, E., Noren, D.P., Rosen, D., Winship, A., 2011. Competing conservation objectives for predators and prey: estimating killer whale prey requirements for chinook salmon. PLoS One 6,
  68. Wright, B.M., Ford, J.K.B., Ellis, G.M., Deecke, V.B., Shapiro, A.D., Battaile, B.C., Trites, A.W., 2017. Fine-scale foraging movements by fish-eating killerwhales (Orcinus orca) relate to the vertical distributions and escape responses of salmonid prey (Oncorhynchus spp.). Mov. Ecol. 5, 3, Scholar
  69. Zhivotovsky, L.A., Shaikhaev, E.G., Shitova, M.V., 2013. Identification of salmonid fish using microsatellite markers with identical pcr-primers. Russian J. Mar. Biol. 39 (6), 447–39454, Scholar

Copyright information

© Deutsche Gesellschaft für Säugetierkunde 2019

Authors and Affiliations

  • Evgeniia V. Volkova
    • 1
    Email author
  • Tatiana V. Ivkovich
    • 1
  • Marina V. Shitova
    • 2
  • Ekaterina N. Chernyaeva
    • 3
  • Tatiana V. Malinina
    • 2
  • Sofia S. Okorokova
    • 4
  • Alexandr M. Burdin
    • 5
  • Erich Hoyt
    • 6
  1. 1.Department of Vertebrate Zoology, Faculty of BiologySt. Petersburg State UniversitySt. PetersburgRussia
  2. 2.Vavilov Institute of General Genetics RASMoscowRussia
  3. 3.Theodosius Dobzhansky Center for Genome BioinformaticsSt. Petersburg State UniversitySt. PetersburgRussia
  4. 4.Russian State Agrarian University Moscow Timiryazev Agricultural AcademyMoscowRussia
  5. 5.Kamchatka Branch of Pacific Institute of Geography FEB RASPetropavlovsk-KamchatskyRussia
  6. 6.Whale and Dolphin ConservationPark HouseDorsetUK

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