Polar Biology

, Volume 37, Issue 7, pp 911–939 | Cite as

A trophic mass balance model of the eastern Chukchi Sea with comparisons to other high-latitude systems

  • G. Andy Whitehouse
  • Kerim Aydin
  • Timothy E. Essington
  • George L. HuntJr.
Original Paper


The Chukchi Sea is a seasonally ice-covered, marginal Arctic-shelf sea that possesses both large petroleum reserves and abundant biological communities, including migratory mammals and seabirds. We developed a mass balance food web model for the eastern Chukchi Sea to evaluate the trophic structure of this ecosystem and to compare food web properties of the Chukchi Sea to those of other high-latitude marine ecosystems. We compiled data on biomass levels, diet composition, demographic rates (production, consumption), and fishery removals, and used these data to construct an Ecopath trophic mass balance model. The majority of biomass was concentrated in benthic invertebrates and most of the mass flow above trophic level 2.0 was through these groups. We found that density estimates of most fish groups derived from trawl survey data using area-swept methods were insufficient to match the consumptive demands of predators, and that densities needed to be several-fold greater to meet modeled demand. We also used a set of system metrics derived from a common modeling framework to highlight differences in ecosystem structure between the eastern Chukchi Sea and other high-latitude systems. The extent of benthic dominance observed in the eastern Chukchi Sea was unique among the systems examined, both in terms of food web structure and associated mass flows between benthic and pelagic components. In relation to total biomass density, the eastern Chukchi Sea had low production when compared with the other systems, and this lower turnover rate suggests that recovery from disturbance might be slow.


Arctic cod Boreogadus saida Chukchi Sea Ecopath Comparative approach Trophic structure 



Funding for this project was provided by the Alaska Fisheries Science Center’s Resource Ecology and Ecosystem Modeling (REEM) Program. This work was improved by the insightful comments and suggestions of P. Boveng, J. Clarke, L. Eisner, M. Ferguson, N. Friday, S. Kotwicki, K. Laidre, M. Martin, B. Norcross, S. Moore, K. Rand, E. Regehr, R. Suydam, and K. Weinberg. Thank you to B. Lauth and the REEM Food Habits Laboratory for supplying data. Thanks also to A. Grieg, G. Lang, G. Mundell and N. Roberson for additional database support. S. Gaichas and I. Ortiz provided invaluable assistance with modeling and graphics. We are grateful to M. Ferguson, L. Logerwell, and two anonymous reviewers who provided valuable comments and criticisms on earlier versions of this paper. The findings and conclusions in the paper are those of the authors and do not necessarily represent the views of the National Marine Fisheries Service. This publication is [partially] funded by the Joint Institute for the Study of the Atmosphere and Ocean (JISAO) under NOAA Cooperative Agreement No. NA10OAR4320148, Contribution No. 2101.

Supplementary material

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Supplementary material 1 (PDF 258 kb)
300_2014_1490_MOESM2_ESM.pdf (511 kb)
Supplementary material 2 (PDF 510 kb)
300_2014_1490_MOESM3_ESM.pdf (392 kb)
Supplementary material 3 (PDF 392 kb)


  1. Aars J, Lunn NJ, Derocher AE (eds) (2006) Polar bears: Proceedings of the 14th working meeting of the IUCN/SSC polar bear specialist group, 20–24 June 2005, Seattle, Washington, USA. IUCN, Gland, Switzerland and Cambridge, UKGoogle Scholar
  2. Abbott DP (1966) The Ascidians. In: Wilimovsky NJ, Wolfe JN (eds) Environment of the Cape Thompson Region, Alaska. U.S. Atomic Energy Commission, Oak Ridge, TN, pp 839–841Google Scholar
  3. ACIA (2004) Impacts of a warming Arctic: Arctic climate impact assessment. Cambridge University Press, Cambridge, UK; New York, NYGoogle Scholar
  4. Acuña JL, Deibel D, Saunders PA, Booth B, Hatfield E, Klein B, Mei ZP, Rivkin R (2002) Phytoplankton ingestion by appendicularians in the North Water. Deep-Sea Res Part II-Top Stud Oceanogr 49:5101–5115. doi: 10.1016/s0967-0645(02)00180-7 Google Scholar
  5. Allen KR (1971) Relation between production and biomass. J Fish Res Bd Can 28:1573–1581Google Scholar
  6. Allen BM, Angliss RP (2010) Alaska marine mammal stock assessments, 2009. U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-206, 276 ppGoogle Scholar
  7. Allen BM, Angliss RP (2012) Alaska marine mammal stock assessments, 2011. U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-234, 288 ppGoogle Scholar
  8. Allen BM, Angliss RP (2013) Alaska marine mammal stock assessments, 2012. U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-245, 282 ppGoogle Scholar
  9. Alverson DL, Wilimovsky NJ (1966) Fishery investigations of the Southeastern Chukchi Sea. In: Wilimovsky NJ, Wolfe JN (eds) Environment of the Cape Thompson Region, Alaska. U.S. Atomic Energy Commission, Oak Ridge, TN, pp 843–860Google Scholar
  10. Ambrose WG, Clough LM, Tilney PR, Beer L (2001) Role of echinoderms in benthic remineralization in the Chukchi Sea. Mar Biol 139:937–949Google Scholar
  11. Ashjian CJ, Campbell RG, Welch HE, Butler M, Van Keuren D (2003) Annual cycle in abundance, distribution, and size in relation to hydrography of important copepod species in the western Arctic Ocean. Deep-Sea Res Part I-Oceanogr Res Pap 50:1235–1261. doi: 10.1016/s0967-0637(03)00129-8 Google Scholar
  12. Atkinson EG, Percy JA (1992) Diet comparison among demersal marine fish from the Canadian Arctic. Polar Biol 11:567–573Google Scholar
  13. Aydin KY (2004) Age structure or functional response? Reconciling the energetics of surplus production between single-species models and ECOSIM. Afr J Mar Sci 26:289–301. doi: 10.2989/18142320409504062 Google Scholar
  14. Aydin K, Gaichas S, Ortiz I, Kinzey D, Friday N (2007) A comparison of the Bering Sea, Gulf of Alaska, and Aleutian Islands large marine ecosystems through food web modeling. U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-178, 298 ppGoogle Scholar
  15. Båmstedt U, Karlson K (1998) Euphausiid predation on copepods in coastal waters of the Northeast Atlantic. Mar Ecol Prog Ser 172:149–168. doi: 10.3354/meps172149 Google Scholar
  16. Banse K, Mosher S (1980) Adult body mass and annual production biomass relationships of field populations. Ecol Monogr 50:355–379. doi: 10.2307/2937256 Google Scholar
  17. Barber WE, Smith RL, Weingartner TJ (1994) Fisheries oceanography of the northeast Chukchi Sea final report. U.S. Dep Int, OCS Study MMS-93-0051Google Scholar
  18. Barber WE, Smith RL, Vallarino M, Meyer RM (1997) Demersal fish assemblages of the northeastern Chukchi Sea, Alaska. Fish Bull US 95:195–208Google Scholar
  19. Barlow J, Boveng P (1991) Modeling age-specific mortality for marine mammal populations. Mar Mamm Sci 7:50–65. doi: 10.1111/j.1748-7692.1991.tb00550.x Google Scholar
  20. Belikov SE (1995) Status of polar bear populations in the Russian Arctic 1993. In: Wiig Ø, Born EW, Garner GW (eds) Polar Bears: Proceedings of the eleventh working meeting of the IUCN/SSC Polar Bear Specialist Group, 25–27 Jan 1993, Copenhagen, Denmark. IUCN, Gland, Switzerland and Cambridge, UK, p 192Google Scholar
  21. Bengtson JL, Hiruki-Raring LM, Simpkins MA, Boveng PL (2005) Ringed and bearded seal densities in the eastern Chukchi Sea, 1999–2000. Polar Biol 28:833–845. doi: 10.1007/s00300-005-0009-1 Google Scholar
  22. Benoit D, Simard Y, Gagne J, Geoffroy M, Fortier L (2010) From polar night to midnight sun: photoperiod, seal predation, and the diel vertical migrations of polar cod (Boreogadus saida) under landfast ice in the Arctic Ocean. Polar Biol 33:1505–1520. doi: 10.1007/s00300-010-0840-x Google Scholar
  23. Bentzen TW, Follmann EH, Amstrup SC, York GS, Wooller MJ, O’Hara TM (2007) Variation in winter diet of southern Beaufort Sea polar bears inferred from stable isotope analysis. Can J Zool 85:596–608. doi: 10.1139/z07-036 Google Scholar
  24. Berchok CL, Clapham PJ, Crance J, Moore SM, Napp J, Overland J, Stabeno P (2012) Passive acoustic detection and monitoring of endangered whales in the Arctic (Beaufort, Chukchi) and ecosystem observations in the Chukchi Sea: Biophysical Moorings and Climate Modeling Annual Report. Submitted to the Bureau of Ocean Energy Management., 15 pp
  25. Blanchard AL, Feder HM (in press) Interactions of habitat complexity and environmental characteristics with macrobenthic community structure at multiple spatial scales. Deep-Sea Res Part II-Top Stud Oceanogr. doi: 10.1016/j.dsr2.2013.09.022
  26. Blanchard JL, Pinnegar JK, Mackinson S (2002) Exploring marine mammal-fishery interactions using ‘Ecopath with Ecosim’: Modelling the Barents Sea ecosystem. Sci Ser Tech Rep No 117. CEFAS Lowestoft, 52 ppGoogle Scholar
  27. Blanchard AL, Parris CL, Knowlton AL, Wade NR (2013) Benthic ecology of the northeastern Chukchi Sea. Part I. Environmental characteristics and macrofaunal community structure, 2008–2010. Cont Shelf Res 67:52–66. doi: 10.1016/j.csr.2013.04.021 Google Scholar
  28. Bluhm BA, Gradinger R (2008) Regional variability in food availability for arctic marine mammals. Ecol Appl 18:S77–S96. doi: 10.1890/06-0562.1 PubMedGoogle Scholar
  29. Bluhm BA, Coyle KO, Konar B, Highsmith R (2007) High gray whale relative abundances associated with an oceanographic front in the south-central Chukchi Sea. Deep-Sea Res Part II-Top Stud Oceanogr 54:2919–2933. doi: 10.1016/j.dsr2.2007.08.015 Google Scholar
  30. Bluhm BA, Iken K, Hardy SM, Sirenko BI, Holladay BA (2009) Community structure of epibenthic megafauna in the Chukchi Sea. Aquat Biol 7:269–293. doi: 10.3354/ab00198 Google Scholar
  31. Bluhm BA, Iken K, Hopcroft RR (2010) Observations and exploration of the Arctic’s Canada Basin and the Chukchi Sea: the Hidden Ocean and RUSALCA expeditions. Deep-Sea Res Part II-Top Stud Oceanogr 57:1–4. doi: 10.1016/j.dsr2.2009.08.001 Google Scholar
  32. Boé JL, Hall A, Qu X (2009) September sea-ice cover in the Arctic Ocean projected to vanish by 2100. Nat Geosci 2:341–343. doi: 10.1038/ngeo467 Google Scholar
  33. Bogoslovskaya LS, Votorogov LM, Semenova TN (1982) Distribution and feeding of gray whales off Chukotka in the summer and autumn of 1980. Rep Int Whal Commn 32:385–389Google Scholar
  34. Boveng PL, Bengtson JL, Buckley TW, Cameron MF, Dahle SP, Kelly BP, Megrey BA, Overland JE, Williamson NJ (2009) Status review of the spotted seal (Phoca largha). U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-200, 153 ppGoogle Scholar
  35. Bradstreet MSW, Cross WE (1982) Trophic relationships at High Arctic ice edges. Arctic 35:1–12Google Scholar
  36. Bradstreet MSW, Finley KJ, Sekerak AD, Griffiths WB, Evans CR, Fabijan MF, Stallard HE (1986) Aspects of the biology of Arctic cod (Boreogadus saida) and its importance in Arctic marine food chains. Can Tech Rep Fish Aquat Sci 1491, 193 ppGoogle Scholar
  37. Brey T (2012) A multi-parameter artificial neural network model to estimate macrobenthic invertebrate productivity and production. Limnol Oceanogr Meth 10:581–589. doi: 10.4319/lom.2012.10.581 Google Scholar
  38. Brey T, Gerdes D (1998) High Antarctic macrobenthic community production. J Exp Mar Biol Ecol 231:191–200. doi: 10.1016/s0022-0981(98)00060-4 Google Scholar
  39. Brodeur RD, Terazaki M (1999) Springtime abundance of chaetognaths in the shelf region of the northern Gulf of Alaska, with observations on the vertical distribution and feeding of Sagitta elegans. Fish Oceanogr 8:93–103. doi: 10.1046/j.1365-2419.1999.00099.x Google Scholar
  40. Bukhtiyarov YA, Frost KJ, Lowry LF (1984) New information on foods of the spotted seal, Phoca largha, in the Bering Sea in spring. In: Fay FH, Fedoseev GA (eds) Soviet-American cooperative research on marine mammals. Volume 1: Pinnipeds. U.S. Dep. Commer., NOAA Tech. Rep. NMFS-12, 55–59 ppGoogle Scholar
  41. Burns JJ, Frost KJ (1983) Natural history and ecology of the bearded seal, Erignathus barbatus. US Dep Commer, NOAA, OCSEAP Final Rep 11:311–392Google Scholar
  42. Byers T, Prach RW (1988) Diet of the kelp snailfish, Liparis tunicatus, in Jones Sound, Canadian High Arctic. Can Field-Nat 102:242–245Google Scholar
  43. Calvert W, Stirling I (1990) Interactions between polar bears and overwintering Walruses in the Central Canadian High Arctic. Int Conf Bear Res and Manage 8:351–356Google Scholar
  44. Cameron MF, Bengtson JL, Boveng PL, Jansen JK, Kelly BP, Dahle SP, Logerwell EA, Overland JE, Sabine CL, Waring GT, Wilder JM (2010) Status review of the bearded seal (Erignathus barbatus). U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-211, 246 ppGoogle Scholar
  45. Campbell RG, Sherr EB, Ashjian CJ, Plourde S, Sherr BF, Hill V, Stockwell DA (2009) Mesozooplankton prey preference and grazing impact in the western Arctic Ocean. Deep-Sea Res Part II-Top Stud Oceanogr 56:1274–1289. doi: 10.1016/j.dsr2.2008.10.027 Google Scholar
  46. Christensen V, Pauly D (1992) Ecopath II-a software for balancing steady-state ecosystem models and calculating network characteristics. Ecol Model 61:169–185. doi: 10.1016/0304-3800(92)90016-8 Google Scholar
  47. Christensen V, Walters CJ (2004a) Ecopath with Ecosim: methods, capabilities and limitations. Ecol Model 172:109–139. doi: 10.1016/j.ecolmodel.2003.09.003 Google Scholar
  48. Christensen V, Walters CJ (2004b) Trade-offs in ecosystem-scale optimization of fisheries management policies. Bull Mar Sci 74:549–562Google Scholar
  49. Christensen V, Walters CJ, Pauly D (2005) Ecopath with Ecosim: A user’s guide. Fisheries Centre, University of British Columbia, Vancouver, Canada,, 154 pp
  50. Christensen V, Walters CJ, Pauly D, Forrest R (2008) Ecopath with Ecosim version 6 User Guide., 235 pp
  51. Christiansen JS, Hop H, Nilssen EM, Joensen J (2012) Trophic ecology of sympatric Arctic gadoids, Arctogadus glacialis (Peters, 1872) and Boreogadus saida (Lepechin, 1774), in NE Greenland. Polar Biol 35:1247–1257. doi: 10.1007/s00300-012-1170-y Google Scholar
  52. Coachman LK, Aagaard K, Tripp RB (1975) Bering Strait: the regional physical oceanography. University of Washington Press, Seattle 172 pGoogle Scholar
  53. Comiso JC, Parkinson CL, Gersten R, Stock L (2008) Accelerated decline in the Arctic Sea ice cover. Geophys Res Lett 35. doi: 10.1029/2007gl031972
  54. Conover RJ, Herman AW, Prinsenberg SJ, Harris LR (1986) Distribution of and feeding by the copepod Pseudocalanus under fast ice during the Arctic spring. Science 232:1245–1247. doi: 10.1126/science.232.4755.1245 PubMedGoogle Scholar
  55. Cooney RT, Coyle KO (1982) Trophic implications of cross-shelf copepod distributions in the southeastern Bering Sea. Mar Biol 70:187–196. doi: 10.1007/bf00397684 Google Scholar
  56. Cooper LW, Lalande C, Pirtle-Levy R, Larsen IL, Grebmeier JM (2009) Seasonal and decadal shifts in particulate organic matter processing and sedimentation in the Bering Strait Shelf region. Deep-Sea Res Part II-Top Stud Oceanogr 56:1316–1325. doi: 10.1016/j.dsr2.2008.10.025 Google Scholar
  57. Coyle KO, Cooney RT (1988) Estimating carbon flux to pelagic grazers in the ice-edge zone of the eastern Bering Sea. Mar Biol 98:299–306. doi: 10.1007/bf00391208 Google Scholar
  58. Coyle KO, Gillispie JA, Smith RL, Barber WE (1997) Food habits of four demersal Chukchi Sea fishes. In: Reynolds JB (ed) Fish ecology in Arctic North America, American Fisheries Society symposium 19. American Fisheries Society, Bethesda, MD, pp 310–318Google Scholar
  59. Craig PC (1987) Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986. U.S. Dep Int, MMS, Alaska Socioeconomic Studies Program, Tech Rep No 129, 67 ppGoogle Scholar
  60. Craig PC, Griffiths WB, Haldorson L, McElderry H (1982) Ecological studies of Arctic cod (Boreogadus saida) in Beaufort Sea coastal waters, Alaska. Can J Fish Aquat Sci 39:395–406Google Scholar
  61. Cui XH, Grebmeier JM, Cooper LW (2012) Feeding ecology of dominant groundfish in the northern Bering Sea. Polar Biol 35:1407–1419. doi: 10.1007/s00300-012-1180-9 Google Scholar
  62. Cusson M, Bourget E (2005) Global patterns of macroinvertebrate production in marine benthic habitats. Mar Ecol Prog Ser 297:1–14. doi: 10.3354/meps297001 Google Scholar
  63. Dalby JE (1992) Prey of the sea anemone Stomphia didemon (Anthozoa: Actiniaria) on the west coast of Canada. Can Field-Nat 106:403–404Google Scholar
  64. Dalpadado P, Yamaguchi A, Ellertsen B, Johannessen S (2008) Trophic interactions of macro-zooplankton (krill and amphipods) in the Marginal Ice Zone of the Barents Sea. Deep-Sea Res Part II-Top Stud Oceanogr 55:2266–2274. doi: 10.1016/j.dsr2.2008.05.016 Google Scholar
  65. Davis ND, Volkov AV, Efimkin AY, Kuznetsova NA, Armstrong JL, Sakai O (2009) Review of BASIS salmon food habits studies. N Pac Anadr Fish Comm Bull 5:197–208Google Scholar
  66. DeBruyn TD, Evans TJ, Miller S, Perham C, Regehr E, Karyn R, Wilder J, Lierheimer LJ (2010) Polar bear conservation in the United States. In: Obbard ME, Thiemann GW, Peacock E, DeBruyn TD (eds) Polar Bears: Proceedings of the 15th working meeting of the IUCN/SSC polar bear specialist group, Copenhagen, Denmark, 29 June–3 July 2009. IUCN, Gland, Switzerland and Cambridge, UK, pp 2005–2009Google Scholar
  67. Dehn LA, Sheffield GG, Follmann EH, Duffy LK, Thomas DL, O’Hara TM (2007) Feeding ecology of phocid seals and some walrus in the Alaskan and Canadian Arctic as determined by stomach contents and stable isotope analysis. Polar Biol 30:167–181. doi: 10.1007/s00300-006-0171-0 Google Scholar
  68. DeRiddler C, Lawrence JM (1982) Food and feeding mechanisms: Echinoidea. In: Jangoux M, Lawrence JM (eds) Echinoderm nutrition. A.A Balkema, Rotterdam, pp 57–115Google Scholar
  69. Derocher AE, Wiig Ø, Andersen M (2002) Diet composition of polar bears in Svalbard and the western Barents Sea. Polar Biol 25:448–452. doi: 10.1007/s00300-002-0364-0 Google Scholar
  70. Douglas DC (2010) Arctic sea ice decline projected changes in timing and extent of sea ice in the Bering and Chukchi Seas. U.S. Geological Survey Open-File Report 2010-1176, 32 ppGoogle Scholar
  71. Dunton KH, Goodall JL, Schonberg SV, Grebmeier JM, Maidment DR (2005) Multi-decadal synthesis of benthic-pelagic coupling in the western arctic: role of cross-shelf advective processes. Deep-Sea Res Part II-Top Stud Oceanogr 52:3462–3477. doi: 10.1016/j.dsr2.2005.09.007 Google Scholar
  72. Dunton KH, Grebmeier JM, Trefry JH, Cooper LW (2012) The COMIDA-CAB Project: an overview of the biological and chemical characteristics of the northern Chukchi Sea benthos. In: Chukchi Sea Offshore Monitoring in Drilling Area (COMIDA): Chemical and Benthos (CAB) Final Report. Prepared For: Bureau of Ocean Energy Management, US Department of the Interior, 3801 Centerpoint Drive, Suite 500, Anchorage, AK 99503, USA,, 6–19 pp
  73. Dyck MG, Kebreab E (2009) Estimating the energetic contribution of polar bear (Ursus maritimus) summer diets to the total energy budget. J Mammal 90:585–593. doi: 10.1644/08-mamm-a-103r2.1 Google Scholar
  74. Emson RH, Mladenov PV, Barrow K (1991) The feeding mechanism of the basket star Gorgonocephalus arcticus. Can J Zool 69:449–455. doi: 10.1139/z91-070 Google Scholar
  75. Essington TE (2004) Getting the right answer from the wrong model: evaluating the sensitivity of multispecies fisheries advice to uncertain species interactions. Bull Mar Sci 74:563–581Google Scholar
  76. Essington TE (2007) Evaluating the sensitivity of a trophic mass-balance model (Ecopath) to imprecise data inputs. Can J Fish Aquat Sci 64:628–637. doi: 10.1139/f07-042 Google Scholar
  77. Essington TE, Kitchell JF, Walters CJ (2001) The von Bertalanffy growth function, bioenergetics, and the consumption rates of fish. Can J Fish Aquat Sci 58:2129–2138. doi: 10.1139/cjfas-58-11-2129 Google Scholar
  78. Fadely BS, Piatt JF, Hatch SA, Roseneau DG (1989) Populations, productivity, and feeding habits of seabirds at Cape Thompson, Alaska : final report. OCS Study MMS 89-0014. U.S. Fish and Wildlife Service, Alaska Fish and Wildlife Research Center, Anchorage, Alaska, 429 ppGoogle Scholar
  79. Fall JA, Brown C, Braem N, Simon JJ, Simeone WE, Holen DL, Naves L, Hutchinson-Scarborough L, Lemons T, Krieg TM (2011) Alaska subsistence salmon fisheries 2008 annual report. Alaska Department of Fish and Game Division of Subsistence, Technical Paper No. 359. Anchorage, AKGoogle Scholar
  80. Fay FH (1982) Ecology and biology of the Pacific walrus, Odobenus rosmarus divergens Illiger. North American Fauna 74:1–279Google Scholar
  81. Fay FH, Feder HM, Stoker SW (1977) An estimation of the impact of the Pacific walrus population on its food resources in the Bering Sea. U.S. Marine Mammal Commission, Report Nos. MMC-75/06 and MMC-74/03. Washington, D.C. 38 ppGoogle Scholar
  82. Fay FH, Kelly BP, Gehnrich PH, Sease JL, Hoover AA (1986) Modern populations, migrations, demography, trophics, and historical status of the Pacific walrus. US Dep Commer, NOAA, OCSEAP Final Rep 37:231–376Google Scholar
  83. Feder HM, Jewett SC (1978) Survey of the epifaunal invertebrates of Norton Sound, southeastern Chukchi Sea, and Kotzebue Sound. IMS Report R78-1. Institute of Marine Science, University of Alaska, Fairbanks, AK, 124 ppGoogle Scholar
  84. Feder HM, Jewett SC (1981) Feeding interactions in the eastern Bering Sea with emphasis on the benthos. In: Hood DW, Calder JA (eds) The Eastern Bering Sea Shelf: oceanography and resources, vol 2. Dep Commer, NOAA, Office of Marine Pollution Assessment, University of Washington Press, Seattle, WA, pp 1229–1261Google Scholar
  85. Feder HM, Paul AJ, Hoberg M, Jewett S, Matheke G, McCumby K, McDonald J, Rice R, Shoemaker P (1981) Distribution, abundance, community structure and trophic relationships of the nearshore benthos of Cook Inlet. US Dep Commer, NOAA, OCSEAP Final Rep 14:45–676Google Scholar
  86. Feder HM, Foster NR, Jewett SC, Weingartner TJ, Baxter R (1994) Mollusks in the northeastern Chukchi Sea. Arctic 47:145–163Google Scholar
  87. Feder HM, Jewett SC, Blanchard A (2005) Southeastern Chukchi Sea (Alaska) epibenthos. Polar Biol 28:402–421. doi: 10.1007/s00300-004-0683-4 Google Scholar
  88. Feder HM, Jewett SC, Blanchard AL (2007) Southeastern Chukchi Sea (Alaska) macrobenthos. Polar Biol 30:261–275. doi: 10.1007/s00300-006-0180-z Google Scholar
  89. Feder HM, Iken K, Blanchard AL, Jewett SC, Schonberg S (2011) Benthic food web structure in the southeastern Chukchi Sea: an assessment using δ13C and δ15N analyses. Polar Biol 34:521–532. doi: 10.1007/s00300-010-0906-9 Google Scholar
  90. Frank PG, Bleakney JS (1978) Asexual reproduction, diet, and anomalies of the anemone Nematostella vectensis in Nova Scotia. Can Field-Nat 92:259–263Google Scholar
  91. Freeman MMR (1973) Polar bear predation on beluga in the Canadian Arctic. Arctic 26:162–163Google Scholar
  92. Frost KJ, Lowry LF (1981) Trophic importance of some marine gadids in northern Alaska and their body-otolith size relationships. Fish Bull US 79:187–192Google Scholar
  93. Frost KJ, Lowry LF (1983) Demersal Fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort seas, 1976-77. U.S. Dep Commer, NOAA Tech Rep NMFS SSRF-764Google Scholar
  94. Frost KJ, Lowry LF (1984) Trophic relationships of vertebrate consumers in the Alaskan Beaufort Sea. In: Barnes PW, Schell DM, Reimnitz E (eds) The Alaskan Beaufort Sea: ecosystems and environments. Academic Press, Orlando, FL, pp 381–401Google Scholar
  95. Fukuchi M, Sasaki H, Hattori H, Matsuda O, Tanimura A, Handa N, McRoy CP (1993) Temporal variability of particulate flux in the northern Bering Sea. Cont Shelf Res 13:693–704. doi: 10.1016/0278-4343(93)90100-c Google Scholar
  96. Furness R, Monaghan P (1987) Seabird ecology. Tertiary level biology. Chapman & Hall, New York 164 pGoogle Scholar
  97. Gaichas S, Skaret G, Falk-Petersen J, Link JS, Overholtz W, Megrey BA, Gjøsæter H, Stockhausen WT, Dommasnes A, Friedland KD, Aydin K (2009) A comparison of community and trophic structure in five marine ecosystems based on energy budgets and system metrics. Prog Oceanogr 81:47–62. doi: 10.1016/j.pocean.2009.04.005 Google Scholar
  98. Gaichas SK, Odell G, Aydin KY, Francis RC (2012) Beyond the defaults: functional response parameter space and ecosystem-level fishing thresholds in dynamic food web model simulations. Can J Fish Aquat Sci 69:2077–2094. doi: 10.1139/f2012-099 Google Scholar
  99. Gall AE, Day RH, Weingartner TJ (2013) Structure and variability of the marine-bird community in the northeastern Chukchi Sea. Cont Shelf Res 67:96–115. doi: 10.1016/j.csr.2012.11.004 Google Scholar
  100. Garner GW, Knick ST, Douglas DC (1990) Seasonal movements of adult female polar bears in the Bering and Chukchi Seas. Int Conf Bear Res and Manage 8:219–226Google Scholar
  101. Garner GW, Belikov SE, Stishov MS, Barnes VG, Arthur SM (1994) Dispersal patterns of maternal polar bears from the denning concentration on Wrangel Island. Int Conf Bear Res and Manage 9:401–410Google Scholar
  102. Gautier DL, Bird KJ, Charpentier RR, Grantz A, Houseknecht DW, Klett TR, Moore TE, Pitman JK, Schenk CJ, Schuenemeyer JH, Sorensen K, Tennyson ME, Valin ZC, Wandrey CJ (2009) Assessment of undiscovered oil and gas in the Arctic. Science 324:1175–1179. doi: 10.1126/science.1169467 PubMedGoogle Scholar
  103. Geoffroy M, Robert D, Darnis G, Fortier L (2011) The aggregation of polar cod (Boreogadus saida) in the deep Atlantic layer of ice-covered Amundsen Gulf (Beaufort Sea) in winter. Polar Biol 34:1959–1971. doi: 10.1007/s00300-011-1019-9 Google Scholar
  104. George JCC, Zeh J, Suydam R, Clark C (2004) Abundance and population trend (1978–2001) of western Arctic bowhead whales surveyed near Barrow, Alaska. Mar Mamm Sci 20:755–773. doi: 10.1111/j.1748-7692.2004.tb01191.x Google Scholar
  105. Gilbert JR, Fedoseev GA, Seagars D, Razlivalov E, LaChugin A (1992) Aerial census of Pacific walrus, 1990. U.S. Fish and Wildlife Service Admin. Rep. R7/MMM Tech. Rep. 92-1. U.S. Dep. Int., Anchorage, AK, 33 ppGoogle Scholar
  106. Gillispie JA, Smith RL, Barbour E, Barber WE (1997) Distribution, abundance, and growth of Arctic cod in the northeastern Chukchi Sea. In: Reynolds JB (ed) Fish Ecology in Arctic North America, American Fisheries Society Symposium 19. American Fisheries Society, Bethesda, MD, pp 81–89Google Scholar
  107. Goddard P, Lauth B, Britt L, Armistead C (2013) Results of the 2012 Arctic EIS Chukchi Sea bottom trawl survey of groundfish and invertebrate resources. Arctic Eis Annual Report for October 1, 2012–September 30, 2013. NOAA/Alaska Fisheries Science Center Submission to the Bureau of Ocean Energy Management October 2013, Retrieved from:, 80 pp
  108. Gradinger RR, Bluhm BA (2004) In-situ observations on the distribution and behavior of amphipods and Arctic cod (Boreogadus saida) under the sea ice of the High Arctic Canada Basin. Polar Biol 27:595–603. doi: 10.1007/s00300-004-0630-4 Google Scholar
  109. Grebmeier JM (1993) Studies of pelagic-benthic coupling extended onto the Soviet continental shelf in the northern Bering and Chukchi seas. Cont Shelf Res 13:653–668. doi: 10.1016/0278-4343(93)90098-i Google Scholar
  110. Grebmeier JM (2012) Shifting patterns of life in the Pacific Arctic and Sub-Arctic Seas. Annu Rev Mar Sci 4:63–78. doi: 10.1146/annurev-marine-120710-100926 Google Scholar
  111. Grebmeier JM, McRoy CP (1989) Pelagic-benthic coupling on the shelf of the northern Bering and Chukchi Seas. III. Benthic food supply and carbon cycling. Mar Ecol Prog Ser 53:79–91. doi: 10.3354/meps053079 Google Scholar
  112. Grebmeier JM, McRoy CP, Feder HM (1988) Pelagic-benthic coupling on the shelf of the northern Bering and Chukchi seas. I. Food supply source and benthic biomass. Mar Ecol Prog Ser 48:57–67. doi: 10.3354/meps048057 Google Scholar
  113. Grebmeier JM, Feder HM, McRoy CP (1989) Pelagic-benthic coupling on the shelf of the Northern Bering and Chukchi seas. II. Benthic community structure. Mar Ecol Prog Ser 51:253–268. doi: 10.3354/meps051253 Google Scholar
  114. Grebmeier JM, Cooper LW, Feder HM, Sirenko BI (2006a) Ecosystem dynamics of the Pacific-influenced Northern Bering and Chukchi Seas in the Amerasian Arctic. Prog Oceanogr 71:331–361. doi: 10.1016/j.pocean.2006.10.001 Google Scholar
  115. Grebmeier JM, Overland JE, Moore SE, Farley EV, Carmack EC, Cooper LW, Frey KE, Helle JH, McLaughlin FA, McNutt SL (2006b) A major ecosystem shift in the northern Bering Sea. Science 311:1461–1464. doi: 10.1126/science.1121365 PubMedGoogle Scholar
  116. Grebmeier JM, Moore SE, Overland JE, Frey KE, Gradinger R (2010) Biological response to recent pacific arctic sea ice retreats. Eos 91:161–162Google Scholar
  117. Hansell DA, Goering JJ (1990) Pelagic nitrogen flux in the northern Bering Sea. Cont Shelf Res 10:501–519. doi: 10.1016/0278-4343(90)90079-2 Google Scholar
  118. Hansell DA, Goering JJ, Walsh JJ, McRoy CP, Coachman LK, Whitledge TE (1989) Summer phytoplankton production and transport along the shelf break in the Bering Sea. Cont Shelf Res 9:1085–1104. doi: 10.1016/0278-4343(89)90060-5 Google Scholar
  119. Harris JL, MacIsaac K, Gilkinson KD, Kenchington EL (2009) Feeding biology of Ophiura sarsii Lütken, 1855 on Banquereau bank and the effects of fishing. Mar Biol 156:1891–1902. doi: 10.1007/s00227-009-1222-1 Google Scholar
  120. Hazard KW, Lowry LF (1984) Benthic prey in a bowhead whale from the northern Bering Sea. Arctic 37:166–168Google Scholar
  121. Hewitt DA, Hoenig JM (2005) Comparison of two approaches for estimating natural mortality based on longevity. Fish Bull US 103:433–437Google Scholar
  122. Highsmith RC, Coyle KO (1990) High productivity of northern Bering Sea benthic amphipods. Nature 344:862–864. doi: 10.1038/344862a0 Google Scholar
  123. Highsmith RC, Coyle KO (1992) Productivity of arctic amphipods relative to gray whale energy requirements. Mar Ecol Prog Ser 83:141–150. doi: 10.3354/meps083141 Google Scholar
  124. Highsmith RC, Coyle KO, Bluhm BA, Konar B (2006) Gray whales in the Bering and Chukchi seas. In: Estes JA, DeMaster DP, Doak DF, Williams TM, Brownell RL Jr (eds) Whales, whaling, and ocean ecosystems. University of California Press, Berkeley, CA, pp 303–313Google Scholar
  125. Hollowed AB, Aydin KY, Essington TE, Ianelli JN, Megrey BA, Punt AE, Smith ADM (2011) Experience with quantitative ecosystem assessment tools in the northeast Pacific. Fish Fish 12:189–208. doi: 10.1111/j.1467-2979.2011.00413.x Google Scholar
  126. Hop H, Welch HE, Crawford RE (1997) Population structure and feeding ecology of Arctic cod schools in the Canadian High Arctic. In: Reynolds JB (ed) Fish Ecology in Arctic North America, American Fisheries Society Symposium 19. American Fisheries Society, Bethesda, MD, pp 68–80Google Scholar
  127. Hopcroft RR, Day RH (2013) Introduction to the special issue on the ecology of the northeastern Chukchi Sea. Cont Shelf Res 67:1–4. doi: 10.1016/j.csr.2013.06.017 Google Scholar
  128. Hopcroft RR, Kosobokova KN, Pinchuk AI (2010) Zooplankton community patterns in the Chukchi Sea during summer 2004. Deep-Sea Res Part II-Top Stud Oceanogr 57:27–39. doi: 10.1016/j.dsr2.2009.08.003 Google Scholar
  129. Hovelsrud GK, McKenna M, Huntington HP (2008) Marine mammal harvests and other interactions with humans. Ecol Appl 18:S135–S147. doi: 10.1890/06-0843.1 PubMedGoogle Scholar
  130. Hunt GL Jr, Megrey BA (2005) Comparison of the biophysical and trophic characteristics of the Bering and Barents Seas. ICES J Mar Sci 62:1245–1255. doi: 10.1016/j.icejms.2005.04.008 Google Scholar
  131. Hunt GL Jr, Burgeson B, Sanger GA (1981) Feeding ecology of seabirds of the eastern Bering Sea. In: Hood DW, Calder JA (eds) The Eastern Bering Sea Shelf: oceanography and resources, vol 2. Dep Commer, NOAA, Office of Marine Pollution Assessment, University of Washington Press, Seattle, WA, pp 629–647Google Scholar
  132. Hunt GL, Jr., Kato H, McKinnell SM (eds) (2000) Predation by marine birds and mammals in the Subarctic North Pacific Ocean. PICES Sci. Rep. No. 14. North Pacific Marine Science Organization, Sidney, B.C., CanadaGoogle Scholar
  133. Hunt GL, Jr., Drinkwater KF (eds) (2005) Ecosystem studies of sub-arctic seas (ESSAS) science plan. GLOBEC Report No. 19Google Scholar
  134. Hunt GL, Blanchard AL, Boveng P, Dalpadado P, Drinkwater KF, Eisner L, Hopcroft RR, Kovacs KM, Norcross BL, Renaud P, Reigstad M, Renner M, Skjoldal HR, Whitehouse A, Woodgate RA (2013) The Barents and Chukchi Seas: comparison of two Arctic shelf ecosystems. J Mar Syst 109–110:43–68Google Scholar
  135. Iken K, Bluhm B, Dunton K (2010) Benthic food-web structure under differing water mass properties in the southern Chukchi Sea. Deep-Sea Res Part II-Top Stud Oceanogr 57:71–85. doi: 10.1016/j.dsr2.2009.08.007 Google Scholar
  136. IPCC (2007) Summary for policymakers. In: Solomon S, Qin D, Manning M et al (eds) Climate change 2007: The physical basis. Contribution of working group I to the fourth assessment report of the intergovernmental panel on climate change. Cambridge University Press, Cambridge, UK and New York, NYGoogle Scholar
  137. Iverson SJ, Stirling I, Lang SLC (2006) Spatial and temporal variation in the diets of polar bears across the Canadian Arctic: indicators of changes in prey populations and environment. In: Boyd I, Wanless S, Camphuysen CJ (eds) Top Predators in Marine Ecosystems: their role in monitoring and management. Cambridge University Press, Cambridge, UK; New York, USA, 98–117 ppGoogle Scholar
  138. Jakobsson M (2002) Hypsometry and volume of the Arctic Ocean and its constituent seas. Geochem Geophys Geosyst 3. doi: 10.1029/2001gc000302
  139. Jewett SC, Feder HM (1980) Autumn food of adult starry flounders, Platichthys stellatus, from the northeastern Bering Sea and the southeastern Chukchi Sea. J Conseil 39:7–14Google Scholar
  140. Johnson ML, Fiscus CH, Ostenson BT, Barbour ML (1966) Marine mammals. In: Wilimovsky NJ, Wolfe JN (eds) Environment of the Cape Thompson Region, Alaska. U.S. Atomic Energy Commission, Oak Ridge, TN, pp 877–924Google Scholar
  141. Kleiber M (1961) The fire of life; an introduction to animal energetics. Wiley, New York 454 pGoogle Scholar
  142. Kochnev AA (2006) Research on polar bear autumn aggregations on Chukotka, 1989–2004. In: Aars J, Lunn NJ, Derocher AE (eds) Polar Bears: proceedings of the 14th working meeting of the IUCN/SSC polar bear specialist group, 20–24 June 2005, Seattle, Washington, USA. IUCN, Gland, Switzerland and Cambridge, UK, pp 157–165Google Scholar
  143. Kruger LM, Griffiths CL (1998) Sea anemones as secondary consumers on rocky shores in the south-western Cape, South Africa. J Nat Hist 32:629–644. doi: 10.1080/00222939800770331 Google Scholar
  144. Laake JL, Punt A, Hobbs R, Ferguson M, Rugh D, Breiwick J (2009) Re-analysis of gray whale southbound migration surveys 1967–2006. U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-203, 55 ppGoogle Scholar
  145. Lane PVZ, Llinás L, Smith SL, Pilz D (2008) Zooplankton distribution in the western Arctic during summer 2002: hydrographic habitats and implications for food chain dynamics. J Mar Syst 70:97–133. doi: 10.1016/j.jmarsys.2007.04.001 Google Scholar
  146. Lang GM, Dodd KA, Livingston PA (2005) Groundfish food habits and predation on commercially important prey species in the eastern Bering Sea from 1997 through 2001. U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-158, 230 ppGoogle Scholar
  147. Levinsen H, Turner JT, Nielsen TG, Hansen BW (2000) On the trophic coupling between protists and copepods in arctic marine ecosystems. Mar Ecol Prog Ser 204:65–77. doi: 10.3354/meps204065 Google Scholar
  148. Lønne OJ, Gabrielsen GW (1992) Summer diet of seabirds feeding in sea-ice-covered waters near Svalbard. Polar Biol 12:685–692Google Scholar
  149. Lønne OJ, Gulliksen B (1989) Size, age and diet of polar cod, Boreogadus saida (Lepechin 1773), in ice covered waters. Polar Biol 9:187–191. doi: 10.1007/bf00297174 Google Scholar
  150. Lovvorn JR, Cooper LW, Brooks ML, De Ruyck CC, Bump JK, Grebmeier JM (2005) Organic matter pathways to zooplankton and benthos under pack ice in late winter and open water in late summer in the north-central Bering Sea. Mar Ecol Prog Ser 291:135–150. doi: 10.3354/meps291135 Google Scholar
  151. Lowry LF (1993) Foods and feeding ecology. In: Burns JJ, Montague JJ, Cowles CJ (eds) The Bowhead Whale. Society for Marine Mammalogy, Lawrence, pp 201–238Google Scholar
  152. Lowry LF, Burns JJ (1980) Foods utilized by bowhead whales near Barter Island, Alaska, autumn 1979. Marine Fisheries Review 42:88–91Google Scholar
  153. Lowry LF, Fay FH (1984) Seal eating by walruses in the Bering and Chukchi seas. Polar Biol 3:11–18. doi: 10.1007/bf00265562 Google Scholar
  154. Lowry LF, Frost KJ (1981a) Distribution, growth, and foods of Arctic cod (Boreogadus saida) in the Bering, Chukchi, and Beaufort seas. Can Field-Nat 95:186–191Google Scholar
  155. Lowry LF, Frost KJ (1981b) Feeding and trophic relationships of Phocid seals and walruses in the eastern Bering Sea. In: Hood DW, Calder JA (eds) The Eastern Bering Sea Shelf: oceanography and resources, vol 2. Dep Commer, NOAA, Office of Marine Pollution Assessment, University of Washington Press, Seattle, WA, pp 813–824Google Scholar
  156. Lowry LF, Frost KJ, Burns JJ (1978) Food of ringed seals and bowhead whales near Point Barrow, Alaska. Can Field-Nat 92:67–70Google Scholar
  157. Lowry LF, Frost KJ, Burns JJ (1980a) Feeding of bearded seals in the Bering and Chukchi seas and trophic interactions with Pacific walruses. Arctic 33:330–342Google Scholar
  158. Lowry LF, Frost KJ, Burns JJ (1980b) Variability in the diet of ringed seals, Phoca hispida, in Alaska. Can J Fish Aquat Sci 37:2254–2261Google Scholar
  159. Lowry LF, Frost KJ, Calkins DG, Swartzman GL, Hills S (1982) Feeding habits, food requirements, and status of Bering Sea marine mammals. Council Documents Nos. 19 and 19A. North Pacific Fisheries Management Council, Anchorage, AK, 574 ppGoogle Scholar
  160. Lowry LF, Frost KJ, Burns JJ (1983) Trophic relationships among ice-inhabiting phocid seals and functionally related marine mammals in the Chukchi Sea. US Dep Commer, NOAA, OCSEAP Final Rep 19:179–229Google Scholar
  161. Lowry LF, Burns JJ, Nelson RR (1987) Polar bear, Ursus maritimus, predation on belugas, Delphinapterus leucas, in the Bering and Chukchi seas. Can Field-Nat 101:141–146Google Scholar
  162. Lunn NJ, Stirling I (1985) The significance of supplemental food to polar bears during the ice-free period of Hudson Bay. Can J Zool 63:2291–2297Google Scholar
  163. Lunn NJ, Schliebe S, Born EW (eds) (2002) Polar Bears: Proceedings of the 13th Working Meeting of the IUCN/SSC Polar Bear Specialist Group, Nuuk, Greenland. IUCN, Gland, Switzerland and Cambridge, UKGoogle Scholar
  164. Massin C (1982) Food and feeding mechanisms: Holothuroidea. In: Jangoux M, Lawrence JM (eds) Echinoderm nutrition. A.A Balkema, Rotterdam, pp 43–55Google Scholar
  165. Matsuno K, Yamaguchi A, Hirawake T, Imai I (2011) Year-to-year changes of the mesozooplankton community in the Chukchi Sea during summers of 1991, 1992 and 2007, 2008. Polar Biol 34:1349–1360. doi: 10.1007/s00300-011-0988-z Google Scholar
  166. McTigue ND, Dunton KH (in press) Trophodynamics and organic matter assimilation pathways in the northeast Chukchi Sea, Alaska. Deep-Sea Res Part II-Top Stud Oceanogr. doi: 10.1016/j.dsr2.2013.07.016
  167. Mecklenburg CW, Stein DL, Sheiko BA, Chernova NV, Mecklenburg TA, Holladay BA (2007) Russian-American long-term census of the Arctic: benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004. Northwest Nat 88:168–187Google Scholar
  168. Megrey BA, Aydin KY (2009) A macrodescriptor perspective of ecological attributes for the Bering and Barents Seas. Deep-Sea Res Part II-Top Stud Oceanogr 56:2132–2140. doi: 10.1016/j.dsr2.2008.11.024 Google Scholar
  169. Megrey BA, Link JS, Hunt GL Jr, Moksness E (2009) Comparative marine ecosystem analysis: applications, opportunities, and lessons learned. Prog Oceanogr 81:2–9. doi: 10.1016/j.pocean.2009.04.002 Google Scholar
  170. Mincks SL, Smith CR, DeMaster DJ (2005) Persistence of labile organic matter and microbial biomass in Antarctic shelf sediments: evidence of a sediment ‘food bank’. Mar Ecol Prog Ser 300:3–19. doi: 10.3354/meps300003 Google Scholar
  171. Moore IA, Moore JW (1974) Food of shorthorn sculpin, Myoxocephalus scorpius, in Cumberland Sound area of Baffin Island. J Fish Res Bd Can 31:355–359Google Scholar
  172. Moore SE, George JC, Sheffield G, Bacon J, Ashjian CJ (2010) Bowhead Whale Distribution and Feeding near Barrow, Alaska, in Late Summer 2005–06. Arctic 63:195–205Google Scholar
  173. Morissette L, Pedersen T, Nilsen M (2009) Comparing pristine and depleted ecosystems: the Sorfjord, Norway versus the Gulf of St. Lawrence, Canada. Effects of intense fisheries on marine ecosystems. Prog Oceanogr 81:174–187. doi: 10.1016/j.pocean.2009.04.013 Google Scholar
  174. Moss JH, Murphy JM, Farley EV, Eisner LB, Andrews AG (2009) Juvenile pink and chum salmon distribution, diet, and growth in the northern Bering and Chukchi seas. N Pac Anadr Fish Comm Bull 5:191–196Google Scholar
  175. Mueter FJ, Litzow MA (2008) Sea ice retreat alters the biogeography of the Bering Sea continental shelf. Ecol Appl 18:309–320. doi: 10.1890/07-0564.1 PubMedGoogle Scholar
  176. Nerini M (1984) A review of gray whale feeding ecology. In: Jones ML, Swartz SL, Leatherwood S (eds) The Gray Whale, Eschrichtius robustus. Academic Press, Orlando, FL, pp 423–450Google Scholar
  177. Norcross BL, Holladay BA, Busby MS, Mier KL (2010) Demersal and larval fish assemblages in the Chukchi Sea. Deep-Sea Res Part II-Top Stud Oceanogr 57:57–70. doi: 10.1016/j.dsr2.2009.08.006 Google Scholar
  178. NPFMC (2009) Fishery management plan for fish resources of the Arctic management area. North Pacific Fishery Management Council, 605 W 4th Avenue, Suite 306, Anchorage AK, 99501,
  179. Ovsyanikov N (2010) Polar bear research on Wrangel Island and in the Central Arctic Basin. In: Obbard ME, Thiemann GW, Peacock E, DeBruyn TD (eds) Polar Bears: Proceedings of the 15th working meeting of the IUCN/SSC polar bear specialist group, Copenhagen, Denmark, 29 June–3 July 2009. IUCN, Gland, Switzerland and Cambridge, UK, pp 171–178Google Scholar
  180. Parker-Stetter SL, Horne JK, Weingartner TJ (2011) Distribution of polar cod and age-0 fish in the US Beaufort Sea. Polar Biol 34:1543–1557. doi: 10.1007/s00300-011-1014-1 Google Scholar
  181. Patent DH (1970) Life history of the basket star, Gorgonocephalus eucnemis (Müller and Troschel) (Echinodermata; Ophiuroidea). Ophelia 8:145–160Google Scholar
  182. Pauly D, Soriano-Bartz M, Palomares ML (1993) Improved construction, parametrization, and interpretation of steady-state ecosystem models. In: Christensen V, Pauly D (eds) Trophic models of aquatic ecosystems, ICLARM Conf Proc No. 26. International Center for Living Aquatic Resources Management; International Council for the Exploration of the Sea: Danish International Development Agency, Manila, Philippines, Copenhagen, Denmark, 1–13 ppGoogle Scholar
  183. Perez MA (1990) Review of marine mammal population and prey information for Bering Sea ecosystem studies. U.S. Dep Commer, NOAA Tech Memo NMFS F/NWC-186, 86 ppGoogle Scholar
  184. Perez MA, McAlister WB, Mooney EE (1990) Estimated feeding rate relationship for marine mammals based on captive animal data. U.S. Dep Commer, NOAA Tech Memo NMFS F/NWC-184, 36 ppGoogle Scholar
  185. Piatt JF, Springer AM (2003) Advection, pelagic food webs and the biogeography of seabirds in Beringia. Mar Ornithol 31:141–154Google Scholar
  186. Piepenburg D (2005) Recent research on Arctic benthos: common notions need to be revised. Polar Biol 28:733–755. doi: 10.1007/s00300-005-0013-5 Google Scholar
  187. Pirtle-Levy R, Grebmeier JM, Cooper LW, Larsen IL (2009) Chlorophyll a in Arctic sediments implies long persistence of algal pigments. Deep-Sea Res Part II-Top Stud Oceanogr 56:1326–1338. doi: 10.1016/j.dsr2.2008.10.022 Google Scholar
  188. Plagányi ÉE (2007) Models for an ecosystem approach to fisheries. FAO Fisheries Technical Paper No. 477. Rome, 108 ppGoogle Scholar
  189. Plagányi ÉE, Butterworth DS (2004) A critical look at the potential of Ecopath with ECOSIM to assist in practical fisheries management. Afr J Mar Sci 26:261–287. doi: 10.2989/18142320409504061 Google Scholar
  190. Polovina JJ (1984) Model of a coral reef ecosystem I. The ECOPATH model and its application to French Frigate Shoals. Coral Reefs 3:1–11. doi: 10.1007/bf00306135 Google Scholar
  191. Polovina JJ, Ow MD (1985) An approach to estimating an ecosystem box model. Fish Bull US 83:457–460Google Scholar
  192. Questel JM, Clarke C, Hopcroft RR (2013) Seasonal and interannual variation in the planktonic communities of the northeastern Chukchi Sea during the summer and early fall. Cont Shelf Res 67:23–41. doi: 10.1016/j.csr.2012.11.003 Google Scholar
  193. Ramsay MA, Hobson KA (1991) Polar bears make little use of terrestrial food webs: evidence from stable-carbon isotope analysis. Oecologia 86:598–600. doi: 10.1007/bf00318328 Google Scholar
  194. Ravelo AM, Konar B, Trefry JH, Grebmeier JM (in press) Epibenthic community variability in the northeastern Chukchi Sea. Deep-Sea Research Part II-Topical Studies in Oceanography. doi: 10.1016/j.dsr2.2013.07.017
  195. Regehr EV, Amstrup SC, Stirling I (2006) Polar bear population status in the southern Beaufort Sea. U.S. Geological Survey Open-File Report 2006-1337, 20 ppGoogle Scholar
  196. Renaud PE, Berge J, Varpe O, Lonne OJ, Nahrgang J, Ottesen C, Hallanger I (2012) Is the poleward expansion by Atlantic cod and haddock threatening native polar cod, Boreogadus saida? Polar Biol 35:401–412. doi: 10.1007/s00300-011-1085-z Google Scholar
  197. Rice RL (1981) Feeding habits of Crangonid shrimps and some aspects of sediment-detrital food systems in lower Cook Inlet, Alaska. US Dep Commer, NOAA, OCSEAP Final Rep 14:611–676Google Scholar
  198. Rice RL, McCumby KI, Feder HM (1980) Food of Pandalus borealis, Pandalus hypsinotus, and Pandalus goniurus (Pandalidae, Decapoda) from Lower Cook Inlet, Alaska. Proceedings of the National Shellfisheries Association 70:47–54Google Scholar
  199. Rooney N, McCann K, Gellner G, Moore JC (2006) Structural asymmetry and the stability of diverse food webs. Nature 442:265–269. doi: 10.1038/nature04887 PubMedGoogle Scholar
  200. Rugh DJ, Shelden KEW (1993) Polar bears, Ursus maritimus, feeding on beluga whales, Delphinapterus leucas. Can Field-Nat 107:235–237Google Scholar
  201. Runge JA, Ingram RG (1988) Underice grazing by planktonic, calanoid copepods in relation to a bloom of ice microalgae in southeastern Hudson Bay. Limnol Oceanogr 33:280–286Google Scholar
  202. Runge JA, Ingram RG (1991) Under-ice feeding and diel migration by the planktonic copepods Calanus glacialis and Pseudocalanus minutus in relation to the ice algal production cycle in southeastern Hudson Bay, Canada. Mar Biol 108:217–225. doi: 10.1007/bf01344336 Google Scholar
  203. Ruppert EE, Barnes RD (eds) (1994) Invertebrate zoology, 6th edn. Saunders College Pub, Fort WorthGoogle Scholar
  204. Sakshaug E (2004) Primary and secondary production in the Arctic Seas. In: Stein R, Macdonald RM (eds) The organic carbon cycle in the Arctic Ocean. Springer, Berlin, pp 57–81Google Scholar
  205. Sambrotto RN, Goering JJ, McRoy CP (1984) Large yearly production of phytoplankton in the western Bering Strait. Science 225:1147–1150. doi: 10.1126/science.225.4667.1147 PubMedGoogle Scholar
  206. Schliebe SL, Evans TJ (1995) Polar Bear Management in Alaska 1988-92. In: Wiig Ø, Born E, Garner GW (eds) Polar Bears: proceedings of the eleventh working meeting of the IUCN/SSC polar bear specialist group. IUCN, Gland, Switzerland and Cambridge, UK, pp 139–144Google Scholar
  207. Schonberg SV, Clarke JT, Dunton KH (in press) Distribution, abundance, biomass and diversity of benthic infauna in the northeast Chukchi Sea, Alaska: Relation to environmental variables and marine mammals. Deep-Sea Research Part II-Topical Studies in Oceanography. doi: 10.1016/j.dsr2.2013.11.004
  208. Schreiber EA, Burger J (eds) (2001) Biology of marine birds. Marine biology. CRC Press, Boca RatonGoogle Scholar
  209. Seaman GA, Lowry LF, Frost KJ (1982) Foods of belukha whales (Delphinapterus leucas) in western Alaska. Cetology 44:1–19Google Scholar
  210. Sheffield G, Grebmeier JM (2009) Pacific walrus (Odobenus rosmarus divergens): differential prey digestion and diet. Mar Mamm Sci 25:761–777. doi: 10.1111/j.1748-7692.2009.00316.x Google Scholar
  211. Sherr EB, Sherr BF, Hartz AJ (2009) Microzooplankton grazing impact in the Western Arctic Ocean. Deep-Sea Res Part II-Top Stud Oceanogr 56:1264–1273. doi: 10.1016/j.dsr2.2008.10.036 Google Scholar
  212. Shuert PG, Walsh JJ (1993) A coupled physical-biological model of the Bering-Chukchi seas. Cont Shelf Res 13:543–573. doi: 10.1016/0278-4343(93)90094-e Google Scholar
  213. Sigler MF, Renner M, Danielson SL, Eisner LB, Lauth RR, Kuletz KJ, Logerwell EA, Hunt GL Jr (2011) Fluxes, fins, and feathers relationships among the Bering, Chukchi, and Beaufort Seas in a time of climate change. Oceanography 24:250–265Google Scholar
  214. Sirenko BI, Gagaev SY (2007) Unusual abundance of macrobenthos and biological invasions in the Chukchi Sea. Russ J Mar Biol 33:355–364. doi: 10.1134/s1063074007060016 Google Scholar
  215. Smith TG (1980) Polar bear predation of ringed and bearded seals in the land-fast sea ice habitat. Can J Zool 58:2201–2209Google Scholar
  216. Smith TG (1985) Polar bears, Ursus maritimus, as predators of belugas, Delphinapterus leucas. Can Field-Nat 99:71–75Google Scholar
  217. Smith TG, Sjare B (1990) Predation of belugas and narwhals by polar bears in nearshore areas of the Canadian High Arctic. Arctic 43:99–102Google Scholar
  218. Smith RL, Barber WE, Vallarino M, Barbour E, Fitzpatrick E (1997a) Population biology of the Bering flounder, Hippoglossoides robustus, from the northeastern Chukchi Sea. In: Reynolds JB (ed) Fish ecology in Arctic North America, American Fisheries Society Symposium 19. American Fisheries Society, Bethesda, MD, pp 127–132Google Scholar
  219. Smith RL, Barber WE, Vallarino M, Gillispie JA, Ritchie A (1997b) Population biology of the Arctic staghorn sculpin, from the northeastern Chukchi Sea. In: Reynolds JB (ed) Fish Ecology in Arctic North America, American Fisheries Society Symposium 19. American Fisheries Society, Bethesda, MD, pp 133–139Google Scholar
  220. Sparks AK, Pereyra WT (1966) Benthic Invertebrates of the Southeastern Chukchi Sea. In: Wilimovsky NJ, Wolfe JN (eds) Environment of the Cape Thompson Region, Alaska. U.S. Atomic Energy Commission, Oak Ridge, TN, pp 817–838Google Scholar
  221. Springer AM, McRoy CP (1993) The paradox of pelagic food webs in the northern Bering Sea-III. Patterns of primary production. Cont Shelf Res 13:575–599. doi: 10.1016/0278-4343(93)90095-f Google Scholar
  222. Springer AM, Roseneau DG, Murphy EC, Springer MI (1984) Environmental controls of marine food webs: food habits of seabirds in the eastern Chukchi Sea. Can J Fish Aquat Sci 41:1202–1215. doi: 10.1139/f84-142 Google Scholar
  223. Springer AM, McRoy CP, Turco KR (1989) The paradox of pelagic food webs in the northern Bering Sea-II. Zooplankton communities. Cont Shelf Res 9:359–386. doi: 10.1016/0278-4343(89)90039-3 Google Scholar
  224. Springer AM, McRoy CP, Flint MV (1996) The Bering Sea Green Belt: shelf-edge processes and ecosystem production. Fish Oceanogr 5:205–223. doi: 10.1111/j.1365-2419.1996.tb00118.x Google Scholar
  225. Stabeno PJ, Farley EV, Kachel NB, Moore S, Mordy CW, Napp JM, Overland JE, Pinchuk AI, Sigler MF (2012) A comparison of the physics of the northern and southern shelves of the eastern Bering Sea and some implications for the ecosystem. Deep-Sea Res Part II-Top Stud Oceanogr 65–70:14–30. doi: 10.1016/j.dsr2.2012.02.019 Google Scholar
  226. Stevenson DE, Lauth RR (2012) Latitudinal trends and temporal shifts in the catch composition of bottom trawls conducted on the eastern Bering Sea shelf. Deep-Sea Res Part II-Top Stud Oceanogr 65–70:251–259. doi: 10.1016/j.dsr2.2012.02.021 Google Scholar
  227. Stirling I, Archibald WR (1977) Aspects of predation of seals by polar bears. J Fish Res Bd Can 34:1126–1129Google Scholar
  228. Stirling I, McEwan EH (1975) The caloric value of whole ringed seals (Phoca hispida) in relation to polar bear (Ursus maritimus) ecology and hunting behavior. Can J Zool 53:1021–1027. doi: 10.1139/z75-117 PubMedGoogle Scholar
  229. Stoker SW (1981) Benthic invertebrate macrofauna of the eastern Bering/Chukchi continental shelf. In: Hood DW, Calder JA (eds) The Eastern Bering Sea Shelf: oceanography and resources, vol 2. Dep Commer, NOAA, Office of Marine Pollution Assessment, University of Washington Press, Seattle, WA, pp 1069–1090Google Scholar
  230. Stroeve J, Holland MM, Meier W, Scambos T, Serreze M (2007) Arctic sea ice decline: faster than forecast. Geophys Res Lett 34. doi: 10.1029/2007gl029703
  231. Tarverdieva MI (1981) Feeding habits of snow crabs Chionoecetes opilio and C. bairdi in the Bering Sea. Zool Zhurnal 60:991–997Google Scholar
  232. Thiemann GW, Iverson SJ, Stirling I (2008) Polar bear diets and arctic marine food webs: insights from fatty acid analysis. Ecol Monogr 78:591–613. doi: 10.1890/07-1050.1 Google Scholar
  233. Thomson DH (1986) Distribution, production, and ecology of gray whale prey species. US Dep Commer, NOAA, OCSEAP Final Rep 43:227–292Google Scholar
  234. Trites AW, Livingston PA, Mackinson S, Vasconcellos MC, Springer AM, Pauly D (1999) Ecosystem change and the decline of marine mammals the eastern Bering Sea : testing the ecosystem shift and commercial whaling hypotheses. Univ. British Columbia Fish. Centr. Res. Rep. 1999, vol 7, 106 ppGoogle Scholar
  235. U.S. Dep. Int. (2010) Secretary Salazar announces comprehensive strategy for offshore oil and gas development and exploration: press release March 31, 2010.
  236. U.S. Dep. Int. (2011) Chukchi Sea OCS oil & gas lease sale 193 record of decision. U.S. Dep. Interior, Bureau of Ocean Energy Management, Washington D.C.,
  237. USFWS (2003) Beringian Seabird Colony Catalog-database. U.S. Fish and Wildlife Service, Migratory Bird Management, Anchorage, AK,
  238. USFWS (2010a) Stock assessment report: Polar bear (Ursus maritimus): Chukchi/Bering seas stock. U.S. Fish and Wildlife Service, Anchorage, AK,
  239. USFWS (2010b) Stock assessment report: Polar bear (Ursus maritimus): Southern Beaufort Sea stock. U.S. Fish and Wildlife Service, Anchorage, AK,
  240. Ver Hoef JM, Cameron MF, Boveng PL, London JM, Moreland EE (2013) A spatial hierarchical model for abundance of three ice-associated seal species in the eastern Bering Sea. Statistical Methodology. doi: 10.1016/j.stamet.2013.03.001 Google Scholar
  241. Vincent TLS, Scheel D, Hough KR (1998) Some aspects of diet and foraging behavior of Octopus dofleini (Wülker, 1910) in its northernmost range. Mar Ecol-Pubbl Stn Zool Napoli 19:13–29Google Scholar
  242. Wakabayashi K, Bakkala RG, Alton MS (1985) Methods of the U.S.-Japan demersal trawl surveys. In: Bakkala RG, Wakabayashi K (eds) Results of the cooperative U.S.-Japan groundfish investigations in the Bering Sea during May-August 1979. Int N Pac Fish Comm Bull, vol 44, 7–29 ppGoogle Scholar
  243. Walkusz W, Majewski A, Reist JD (2012) Distribution and diet of the bottom dwelling Arctic cod in the Canadian Beaufort Sea. J Mar Syst. doi: 10.1016/j.jmarsys.2012.04.004 Google Scholar
  244. Walsh JJ, McRoy CP, Coachman LK, Goering JJ, Nihoul JJ, Whitledge TE, Blackburn TH, Parker PL, Wirick CD, Shuert PG, Grebmeier JM, Springer AM, Tripp RD, Hansell DA, Djenidi S, Deleersnijder E, Henriksen K, Lund BA, Andersen P, Müller-Karger FE, Dean K (1989) Carbon and nitrogen cycling within the Bering/Chukchi seas: source regions for organic matter effecting AOU demands of the Arctic Ocean. Prog Oceanogr 22:277–359. doi: 10.1016/0079-6611(89)90006-2 Google Scholar
  245. Walsh JJ, Dieterle DA, Maslowski W, Grebmeier JM, Whitledge TE, Flint M, Sukhanova IN, Bates N, Cota GF, Stockwell D, Moran SB, Hansell DA, McRoy CP (2005) A numerical model of seasonal primary production within the Chukchi/Beaufort Seas. Deep-Sea Res Part II-Top Stud Oceanogr 52:3541–3576. doi: 10.1016/j.dsr2.2005.09.009 Google Scholar
  246. Wang MY, Overland JE (2009) A sea ice free summer Arctic within 30 years? Geophys Res Lett 36. doi: 10.1029/2009gl037820
  247. Warner GF (1982) Food and feeding mechanisms: Ophiuroidea. In: Jangoux M, Lawrence JM (eds) Echinoderm nutrition. A.A Balkema, Rotterdam, pp 161–181Google Scholar
  248. Wassmann P, Reigstad M (2011) Future Arctic Ocean Seasonal Ice Zones and Implications for Pelagic-Benthic Coupling. Oceanography 24:220–231Google Scholar
  249. Weingartner T, Aagaard K, Woodgate R, Danielson S, Sasaki Y, Cavalieri D (2005) Circulation on the north central Chukchi Sea shelf. Deep-Sea Res Part II-Top Stud Oceanogr 52:3150–3174. doi: 10.1016/j.dsr2.2005.10.015 Google Scholar
  250. Welch HE, Bergmann MA, Siferd TD, Martin KA, Curtis MF, Crawford RE, Conover RJ, Hop H (1992) Energy flow through the marine ecosystem of the Lancaster Sound Region, Arctic Canada. Arctic 45:343–357Google Scholar
  251. Whitehouse GA (2013) A preliminary mass-balance food web model of the eastern Chukchi Sea. U.S. Dep Commer, NOAA Tech Memo NMFS-AFSC-262, 162 ppGoogle Scholar
  252. Wolotira RJ (1985) Saffron cod (Eleginus gracilis) in western Alaska: the resource and its potential. U.S. Dep Commer, NOAA Tech Memo NMFS F/NWC-79. National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Northwest and Alaska Fisheries Center, Seattle, WA, 119 ppGoogle Scholar
  253. Wolotira RJ, Sample TM, Morin M (1977) Demersal fish and shellfish resources of Norton Sound, the Southeastern Chukchi Sea, and adjacent waters in the baseline year 1976. U.S. Dep Commer, NWAFC Processed Report, 292 p. Northwest and Alaska Fish. Cent., Natl. Mar. Fish. Serv., NOAA, 7600 Sand Point Way NE, Seattle WA 98115, 292 ppGoogle Scholar
  254. Woodgate RA, Aagaard K (2005) Revising the Bering Strait freshwater flux into the Arctic Ocean. Geophys Res Lett 32. doi: 10.1029/2004gl021747
  255. Woodgate RA, Aagaard K, Weingartner TJ (2005) Monthly temperature, salinity, and transport variability of the Bering Strait through flow. Geophys Res Lett 32. doi: 10.1029/2004gl021880
  256. Wyllie-Echeverria T, Wooster WS (1998) Year to-year variations in Bering Sea ice cover and some consequences for fish distributions. Fish Oceanogr 7:159–170. doi: 10.1046/j.1365-2419.1998.00058.x Google Scholar
  257. Zador S, Gaichas S (eds) (2010) Ecosystem considerations for 2011, Appendix C. Stock Assessment and fishery evaluation report for the groundfish resources of the Bering Sea/Aleutian Islands Regions. North Pacific Fishery Management Council, Anchorage, Alaska,

Copyright information

© Springer-Verlag Berlin Heidelberg 2014

Authors and Affiliations

  • G. Andy Whitehouse
    • 1
    • 2
    • 3
  • Kerim Aydin
    • 2
  • Timothy E. Essington
    • 1
  • George L. HuntJr.
    • 1
  1. 1.School of Aquatic and Fishery SciencesUniversity of WashingtonSeattleUSA
  2. 2.NOAA Alaska Fisheries Science CenterSeattleUSA
  3. 3.Joint Institute for the Study of the Atmosphere and OceanUniversity of WashingtonSeattleUSA

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