Polar Biology

, Volume 36, Issue 3, pp 305–320 | Cite as

Harp seal foraging behaviour during summer around Svalbard in the northern Barents Sea: diet composition and the selection of prey

  • Ulf Lindstrøm
  • Kjell T. Nilssen
  • Line M. S. Pettersen
  • Tore Haug
Original Paper


The harp seal Pagophilus groenlandicus is a major high trophic level predator in the Barents Sea, and to better understand their function in the Barents Sea ecosystem, we need to understand their foraging behaviour during their most intensive feeding period. We analysed the diet composition and prey preference of 184 harp seals and 94 faeces samples, sampled in the northern Barents Sea (around Svalbard) during the period May–August in 1996, 1997, and 2004–2006. Concurrent with the sampling of seals, prey availability was assessed in one area in 1996 and 1997 and in two areas in 2006 using standard acoustic methods. Our study showed that harp seal diet composition varied significantly both in time (year) and space, and that their diets appeared to be size dependent. Both subadult (<150 cm) and adult seals were associated with pelagic crustaceans (particularly krill), whereas primarily adult seals were associated with fish (capelin, gadoids and flatfish). Krill was the most important prey group (63 %) followed by polar cod (16 %) and other fish species (10 %). The prey preference of harp seals varied in time and space; polar cod was often preferred by the seals whereas krill was commonly consumed in lower proportion than observed in the survey area. Gadoids and capelin had either been exploited in the same or less proportion as observed in the survey sea. This study emphasises the ecological significance of krill as prime food for harp seals during their intensive feeding period in summer.


Harp seals Diets Prey selection Barents Sea 



The authors thank L. Lindblom, N. E Skavberg and the crew on board M/S Jan Mayen, along with the two reviewers who gave valuable comments to the manuscript. Sophie Smout has kindly proof-read the manuscript.

Supplementary material

300_2012_1260_MOESM1_ESM.doc (57 kb)
Supplementary material 1 (DOC 57 kb)


  1. Abrams PA, Ginzburg LR (2000) The nature of predation: prey dependent, ratio dependent or neither? TREE 15:337–341PubMedGoogle Scholar
  2. Barrett R, Anker-Nilssen T, Gabrielsen GW, Chapdelaine G (2002) Food consumption by seabirds in Norwegian waters. ICES J Mar Sci 59:43–57CrossRefGoogle Scholar
  3. Bodholt H, Nes H, Solli H (1989) A new echo-sounder system. Proc Inst Acoust 11:123–130Google Scholar
  4. Bogstad B, Hauge KH, Ulltang Ø (1997) MULTSPEC—a multispecies model for fish and marine mammals in the Barents Sea. J Northw Atl Fish Sci 22:317–341CrossRefGoogle Scholar
  5. Bogstad B, Haug T, Mehl S (2000) Who eats whom in the Barents Sea. NAMMCO Sci Publ 2:98–119Google Scholar
  6. Bowen WD, Tully D, Boness DJ, Bulheier BM, Marshall GJ (2002) Prey-dependent foraging tactics and prey profitability in a marine mammal. Mar Ecol Prog Ser 244:235–245CrossRefGoogle Scholar
  7. Bradstreet M, Cross W (1982) Trophic relationships at high arctic ice edges. Arctic 35:1–12Google Scholar
  8. Breiby A (1985) Otolitter fra saltvannsfisker i Nord Norge. Tromura Naturvitenskap 53:1–30Google Scholar
  9. Chapskii KK (1961) Nektorye ekologicheskie obosnovanniia sezonnoj dynamiki areala belomorskoj populjatsii grenlandskogo tjulerja (Pagophilus groenlandica) (Some biological factors determining seasonal changes in distribution of the White Sea harp seal population). Trudy Soveschanija, Ichtiologicheskoi Komissii Akademii Nauk. SSSR 12:150–163 (Fisheries Research Board of Canada, Translation Series, 380:1–22)Google Scholar
  10. Charnov E (1976) Optimal foraging: the marginal value theorem. Theor Pop Biol 9:129–136CrossRefGoogle Scholar
  11. Chesson J (1978) Measuring preference in selective predation. Ecology 59:211–215CrossRefGoogle Scholar
  12. Cressie NAC (1993) Statistics for spatial data. Wiley, New York, p 900Google Scholar
  13. Dalpadado P, Skjoldal HR (1996) Abundance, maturity and growth of krill species Thysanoessa inermis and T. longicaudata in the Barents Sea. Mar Ecol Prog Ser 144:175–183CrossRefGoogle Scholar
  14. Dommasnes A, Christensen V, Ellertsen B, Kvamme C, Melle W, Nøttestad L, Pedersen T, Tjelmeland S, Zeller D (2001) An Ecopath model for the Norwegian and Barents Sea. In: Guénette S, Christensen V, Pauly D (eds) Fisheries impacts on North Atlantic ecosystems: models and analyses. Fish Centre Res Rep 9(4):213–240Google Scholar
  15. Efron B, Tibshirani RJ (1993) An introduction to bootstrap. Chapman and Hall, Inc., New York, p 436Google Scholar
  16. Enckell PH (1980) Fältfauna/Kräftdjur. Bokförlaget Sigmun i Lund, pp 685Google Scholar
  17. Eriksen E, Dalpadado P (2011) Long-term changes in Krill biomass and distribution in the Barents Sea: are the changes mainly related to capelin stock size and temperature conditions? Polar Biol 34:1399–1409CrossRefGoogle Scholar
  18. Falk-Petersen S, Hopkins CCE, Sargent JR (1990) Trophic relationships in the pelagic, Arctic food web. In: Barnes M, Gibson RN (eds) Trophic relationship in the marine environment. University Press, Aberdeen, pp 315–333Google Scholar
  19. Falk-Petersen S, Haug T, Hop H, Nilssen KT, Wold A (2009) Transfer of lipids from plankton to blubber of harp and hooded seals off East Greenland. Deep Sea Res II 56:2080–2086CrossRefGoogle Scholar
  20. Finley KJ, Bradstreet MSW, Miller GW (1990) Summer feeding ecology of harp seals (Phoca groenlandica) in relation to Arctic cod (Boreogadus saida) in the Canadian high Arctic. Polar Biol 10:609–618CrossRefGoogle Scholar
  21. Foote KG, Knudsen HP, Korneliussen RJ, Nordbø PE, Røang K (1991) Post processing system for echo sounder data. J Acoust Soc Am 90:37–47CrossRefGoogle Scholar
  22. Frie AK, Potelov VA, Kingsley MCS, Haug T (2003) Trends in age at maturity and growth parameters of female northeast Atlantic harp seals, Pagophilus groenlandicus. ICES J Mar Sci 60:1018–1032CrossRefGoogle Scholar
  23. Fryxell JM, Lundberg P (1994) Diet choice and predator-prey dynamics. Evol Ecol 8:407–421CrossRefGoogle Scholar
  24. Gjøsæter H, Bogstad B (1998) Effects of the presence of herring (Clupea harengus) on the stock–recruitment relationship of Barents Sea capelin (Mallotus villosus). Fish Res 38:57–71CrossRefGoogle Scholar
  25. Gjøsæter H, Bogstad B, Tjelmeland S (2009) Ecosystem effects of the three capelin stock collapses in the Barents Sea. Mar Biol Res 5:40–53CrossRefGoogle Scholar
  26. Grahl-Nielsen O, Haug T, Lindstrøm U, Nilssen KT (2011) Fatty acids in harp seal blubber do not necessarily reflect their diet. Mar Ecol Prog Ser 426:263–276CrossRefGoogle Scholar
  27. Grellier K, Hammond PS (2006) Robust digestion and passage rate estimates for hard parts of grey seal (Halichoerus grypus) prey. Can J Fish Aquat Sci 63:1982–1998CrossRefGoogle Scholar
  28. Hamre J (1994) Biodiversity and exploitation of the main fish stocks in the Norwegian-Barents Sea ecosystem. Biodiv Conserv 3:473–492CrossRefGoogle Scholar
  29. Harbitz A, Lindstrøm U (2001) Stochastic spatial analysis of marine resources with application to minke whale foraging: a synoptic case study from the southern Barents. Sarsia 86:485–5010Google Scholar
  30. Härkönen T (1986) Guide to the otoliths of the bony fishes of the northeast Atlantic. Danbiu ApS, HellerupGoogle Scholar
  31. Hassel MP, May RM (1974) Aggregation of predators and insect parasites and its effect on stability. J Anim Ecol 43:567–594CrossRefGoogle Scholar
  32. Haug T, Nilssen KT, Øien N, Potelov V (1994) Seasonal distribution of harp seals (Phoca groenlandica) in the Barents Sea. Polar Res 13:161–172Google Scholar
  33. Haug T, Lindstrøm U, Nilssen KT (2002) Variations in minke whale (Balaenoptera acutorostrata) diet and body condition in response to ecosystem changes in the Barents Sea. Sarsia 87:409–422CrossRefGoogle Scholar
  34. Haug T, Nilssen KT, Lindblom L (2004) Feeding habits of harp and hooded seals in drift ice waters along the east coast of Greenland in summer and winter. Polar Res 23:35–42CrossRefGoogle Scholar
  35. Haug T, Nilssen KT, Lindblom L, Lindstrøm U (2007) Diets of hooded seals (Cystophora cristata) in coastal waters and drift ice waters along the east coast of Greenland. Mar Biol Res 3:123–133CrossRefGoogle Scholar
  36. Hjermann DØ, Bogstad B, Eikeset AM, Ottersen G, Gjøsæter H, Stenseth NC (2007) Food web dynamics affect Northeast Arctic cod recruitment. Proc R Soc B 274:661–669PubMedCrossRefGoogle Scholar
  37. ICES (2011a) Report of the Arctic Fisheries Working Group, 28 April–4 May 2011, Hamburg, Germany. ICES CM 2011/ACOM:05Google Scholar
  38. ICES (2011b) Report of the joint ICES/NAFO Working Group on Harp and Hooded Seals. St.Andrews, Scotland, UK, 15–19 August 2011. ICES CM 2011/ACOM:22Google Scholar
  39. Johannesen E, Ingvaldsen RB, Bogstad B, Dalpadado P, Eriksen E, Gjøsæter H, Knutsen T, Skern-Mauritzen M, Stiansen JE (2012) Changes in Barents Sea ecosystem state, 1970–2009: climate fluctuations, human impact, and trophic interactions. ICES J Mar Sci 69:880–889CrossRefGoogle Scholar
  40. Kapel FO (2000) Feeding habits of harp and hooded seals in Greenland waters. NAMMCO Sci Publ 2:50–64Google Scholar
  41. Kitandis PK (1997) Introduction to geostatistics: applications to hydrogeology. Cambridge University Press, Cambridge, p 247CrossRefGoogle Scholar
  42. Kotler BP, Brown JS (1988) Environmental heterogeneity and coexistence of desert rodents. Annu Rev Ecol Syst 19:281–307CrossRefGoogle Scholar
  43. Kotler BP, Brown JS (1999) Mechanisms of coexistence of optimal foragers as determinants of the local abundance and distributions of desert granivores. J Mamm 80:361–374CrossRefGoogle Scholar
  44. Lawson JW, Anderson JT, Dalley EL, Stenson GB (1998) Selective foraging by harp seals Phoca groenlandica in nearshore and offshore waters of Newfoundland, 1993 and 1994. Mar Ecol Prog Ser 163:1–10CrossRefGoogle Scholar
  45. Legendre P, Anderson MJ (1999) Distance-based redundancy analysis: testing multispecies responses in multifactorial ecological experiments. Ecol Monogr 69:1–24CrossRefGoogle Scholar
  46. Legendre P, Legendre L (1998) Numerical ecology, 2nd edn. Elsevier Science, Amsterdam, p 853Google Scholar
  47. Lindstrøm U, Harbitz A, Haug T, Nilssen KT (1998) Do harp seals Phoca groenlandica exhibit particular prey preferences? ICES J Mar Sci 55:941–953CrossRefGoogle Scholar
  48. Lindstrøm U, Smouth S, Howell D, Bogstad B (2009) Modelling multi-species interactions in the Barents Sea ecosystem with special emphasis on minke whales and their interactions with cod, herring and capelin. Deep-Sea Res II 56:2068–2079CrossRefGoogle Scholar
  49. MacLennan DN, Simmonds EJ (1992) Fisheries acoustics. Chapman & Hall, London, p 325Google Scholar
  50. Mårtensson PE, Nordøy ES, Messelt EB, Blix AS (1998) Gut length, food transit time and diving habit in phocid seals. Polar Biol 20:213–217CrossRefGoogle Scholar
  51. Matthiopoulos J, Smout S, Winship AJ, Thompson D, Boyd IL, Harwood J (2008) Getting beneath the surface of marine mammal—fisheries competition. Mammal Rev 38:167–188CrossRefGoogle Scholar
  52. Murphy EJ, Morris DJ, Watkins JL, Priddle J (1988) Scales of interaction between Antarctic krill and the environment. In: Sahrhage D (ed) Antarctic Ocean and resources variability. Springer, Berlin, pp 120–130CrossRefGoogle Scholar
  53. Nilssen KT (1995) Seasonal distribution, condition and feeding habits of Barents Sea harp seals (Phoca groenlandica). In: Blix AS, Walløe L, Ulltang Ø (eds) Whales, seals, fish and man. Elsevier Science BV, Amsterdam, pp 241–254Google Scholar
  54. Nilssen KT, Haug T, Potelov V, Stasenkov VA, Timoshenko YK (1995a) Food habits of harp seals (Phoca groenlandica) during lactation and moult in March–May in the southern Barents Sea and White Sea. ICES J Mar Sci 52:33–41CrossRefGoogle Scholar
  55. Nilssen KT, Haug T, Potelov V, Timoshenko YK (1995b) Food habits and food availability of harp seals (Phoca groenlandica) during early summer and autumn in the northern Barents Sea. Polar Biol 15:485–493CrossRefGoogle Scholar
  56. Nilssen KT, Haug T, Grotnes PE, Potelov V (1997) Seasonal variation in body condition of adult Barents Seaharp seals (Phoca groenlandica). J Northw Atl Fish Sci 22:17–25CrossRefGoogle Scholar
  57. Nilssen KT, Pedersen OP, Folkow LP, Haug T (2000) Food consumption estimates of Barents Sea harp seals. NAMMCO Sci Publ 2:9–27Google Scholar
  58. Nordøy ES, Folkow LP, Potelov V, Prischemikhin V, Blix AS (2008) Seasonal distribution and dive behaviour of harp seals (Pagophilus groenlandicus) of the White Sea-Barents Sea stock. Polar Biol 31:1119–1135CrossRefGoogle Scholar
  59. Øigård TA, Lindstrøm U, Nilssen KT, Haug T (in revision) Functional relationship between harp seal body condition, environment and prey in the Barents Sea. Mar Ecol Prog SerGoogle Scholar
  60. Oksanen J (2011) Multivariate analyses of ecological communities in R: vegan tutorial. http://cc.oulu.fi/~jarioksa/opetus/metodi/vegantutor.pdf
  61. Pethon P (1985) Ascheougs store fiskebog. Aschehoug H and company (Nygaard W) A/S, NorwayGoogle Scholar
  62. Piatt JF, Methven DA (1992) Threshold foraging behaviour of baleen whales. Mar Ecol Prog Ser 84:205–210CrossRefGoogle Scholar
  63. Potelov V, Nilssen KT, Svetochev V, Haug T (2000) Feeding habits of harp (Phoca groenlandica) and hooded seals (Cystophora cristata) during late winter, spring and early summer in the Greenland Sea. NAMMCO Sci Publ 2:40–49Google Scholar
  64. Rosenzweig ML, MacArthur RH (1963) Graphical representation and stability conditions of predator-prey interactions. Am Nat 97:209–223CrossRefGoogle Scholar
  65. Ryg M, Lydersen C, Markusen NH, Smith TG, Øritsland NA (1990) Estimating the blubber content of phocid seals. Can J Fish Aquat Sci 47:1223–1227CrossRefGoogle Scholar
  66. Sakshaug E, Bjørge A, Gulliksen B, Loeng H, Mehlum F (1992) Økosystem Barentshavet. Universitetsforlaget, Oslo, p 304Google Scholar
  67. Sergeant D (1973) Feeding, growth, and productivity of northwest Atlantic harp seals (Pagophilus groenlandicus). J Fish Res Board Can 30:17–29CrossRefGoogle Scholar
  68. Sih A, Englund G, Wooster D (1998) Emergent impacts of multiple predators on prey. TREE 13:350–355PubMedGoogle Scholar
  69. Skjoldal HR, Hassel A, Rey F (1987) Spring phytoplankton development and zooplankton reproduction in the central Barents Sea in the period 1979–1984. In: Loeng H (ed) The effect of oceanographic conditions on distribution and population dynamics of commercial fish stocks in the Barents Sea. Proceedings of the 3rd Soviet–Norwegian Symposium, IMR/PINRO, Bergen, pp 59–89Google Scholar
  70. ter Braak CJF, Verdonschot PMF (1995) Canonical correspondence analysis and related multivariate methods in aquatic ecology. Aquat Sci 57:255–289CrossRefGoogle Scholar
  71. Vincent TLS, Scheel D, Brown JS, Vincent TL (1996) Trade-offs and coexistence in consumer-resource models: it all depends on what and where you eat. Am Nat 148:1038–1058CrossRefGoogle Scholar
  72. Wassmann P, Reigstad M, Haug T, Rudels B, Carroll ML, Hop H, Gabrielsen GW, Falk-Petersen S, Denisenko SG, Arashkevich E, Slagstad D, Pavlova O (2006) Food webs and carbon flux in the Barents Sea. Progr Oceanogr 71:232–287CrossRefGoogle Scholar
  73. Windsland K, Lindstrøm U, Nilssen KT, Haug T (2007) Relative abundance and size composition of prey in the common minke whale diet in selected areas of the northeast Atlantic during 2000–04. J Cetacean Res Manage 9:167–178Google Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2012

Authors and Affiliations

  • Ulf Lindstrøm
    • 1
  • Kjell T. Nilssen
    • 1
  • Line M. S. Pettersen
    • 2
  • Tore Haug
    • 1
  1. 1.Institute of Marine ResearchTromsøNorway
  2. 2.University of TromsøTromsøNorway

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