Oceanography The Official Magazine of
The Oceanography Society
Volume 24 Issue 03

View Issue TOC
Volume 24, No. 3
Pages 232 - 248


Arctic Marine Biodiversity: An Update of Species Richness and Examples of Biodiversity Change

By Bodil A. Bluhm , Andrey V. Gebruk, Rolf Gradinger, Russell R. Hopcroft , Falk Huettmann, Ksenia N. Kosobokova, Boris I. Sirenko, and Jan Marcin Weslawski  
Jump to
Article Abstract Citation References Copyright & Usage
Article Abstract

The societal need for—and urgency of obtaining—basic information on the distribution of Arctic marine species and biological communities has dramatically increased in recent decades as facets of the human footprint alter Arctic marine biodiversity. The primary goals of this article are to present updated species inventories based on focused biodiversity research over the last decade, to give examples of emerging recent changes in diversity as indicators of environmental change, and to recommend future diversity-related research areas. Species inventories across all eukaryotic taxonomic levels now total close to 8,000 species, with several thousand additional benthic species predicted to be recorded or discovered in the future. The currently known Arctic species richness estimate includes close to 2,000 phytoplankton taxa, over 1,000 ice-associated protists, greater than 50 ice-associated metazoans, ~ 350 multicellular zooplankton species, over 4,500 benthic protozoans and invertebrates, at least 160 macro-algae, 243 fishes, 64 seabirds, and 16 marine mammals. Endemic and abundant species are present in all three environmental realms (sea ice, water column, and seafloor) and across phyla. The few published time series on Arctic marine biodiversity have detected interannual and interdecadal variability or change both in pelagic and benthic habitats, and at virtually all trophic levels. We identify knowledge gaps and stress the urgency to fill them. We recommend regular, strategic, and sustained monitoring of Arctic marine biodiversity in a public, open-access fashion in order to provide comprehensive data to inform management, conservation, and other decisions.


Bluhm, B.A., A.V. Gebruk, R. Gradinger, R.R. Hopcroft, F. Huettmann, K.N. Kosobokova, B.I. Sirenko, and J.M. Weslawski. 2011. Arctic marine biodiversity: An update of species richness and examples of biodiversity change. Oceanography 24(3):232–248, https://doi.org/10.5670/oceanog.2011.75.


Aars, J., N.F. Lunn, and A.E. Derocher, eds. 2006. Polar Bears: Proceedings of the 14th Working Meeting of the IUCN/SSC Polar Bears Specialist Group, June 20–24, 2005, Seattle,Washington, USA. International Union for Conservation of Nature, Cambridge, MA.

ACIA. 2005. Arctic Climate Impact Assessment. Cambridge University Press, Cambridge, 1,042 pp.

Agatha, S., M. Spindler, and N. Wilbert. 1993. Ciliated Protozoa (Ciliophora) from Arctic sea ice. Acta Protozoologica 32:261–268.

Ambrose, W.G. Jr., C. von Quillfeldt, L.M. Clough, P.V.R. Tilney, and T. Tucker. 2005. The sub-ice algal community in the Chukchi Sea: Large- and small-scale patterns of abundance based on images from a remotely operated vehicle. Polar Biology 28:784–795, https://doi.org/10.1007/s00300-005-0002-8.

Andriashev, A.P., and N.V. Chernova. 1995. Annotated list of fishlike vertebrates and fish of the Arctic seas and adjacent waters. Journal of Ichthyology 35:81–123.

Andronov, V.N., and K.N. Kosobokova. 2011. New species of small, bathypelagic calanoid copepods from the Arctic Ocean: Brodskius arcticus sp. nov. (Tharybidae) and three new species of Pertsoviusgen nov. (Discoidae). Zootaxa 2,809:33–46.

Archambault, P., P.V.R. Snelgrove, J.A.D. Fisher, J.-M. Gagnon, D.J. Garbary, M. Harvey, E.L. Kenchington, V. Lesage, M. Levesque, C. Lovejoy, and others. 2010. From sea to sea: Canada’s three oceans of biodiversity. PLoS ONE 5(8):e12182, https://doi.org/10.1371/journal.pone.0012182

Arndt, C.E., G. Fernandez-Leborans, L. Seuthe, J. Berge, and B. Gulliksen. 2005. Ciliated epibionts on the Arctic sympagic amphipod Gammarus wilkitzkii as indicators for sympago-benthic coupling. Marine Biology 147:643–652, https://doi.org/10.1007/s00227-005-1599-4.

Berge, J., G. Johnsen, F. Nilsen, B. Gulliksen, and D. Slagstad. 2005. Ocean temperature oscillations enable reappearance of blue mussel Mytilus edulis in Svalbard after 1000 years of absence. Marine Ecology Progress Series 303:167–175, https://doi.org/10.3354/meps303167.

Beuchel, F., B. Gulliksen, and M.L. Carroll. 2006. Long-term patterns of rocky bottom macrobenthic community structure in an Arctic fjord (Kongsfjorden, Svalbard) in relation to climate variability (1980–2003). Journal of Marine Systems 63:35–48, https://doi.org/10.1016/j.jmarsys.2006.05.002.

Blacker, R. 1957. Benthic animals as indicators of hydrographic conditions and climatic change in Svalbard waters. Fishery Investigations, series 2, 20(10):1–59.

Błachowiak-Samołyk, K. 2008. Contrasting zooplankton communities (Arctic vs. Atlantic) in the European Arctic Marginal Ice Zone. Oceanologica 50:363–389.

Bluhm, B.A., A.G. Ambrose Jr., M. Bergmann, L.M. Clough, A.V. Gebruk, C. Hasemann, K. Iken, M. Klages, I.R. MacDonald, P.E. Renaud, and others. 2011. Diversity of the Arctic deep-sea benthos. Marine Biodiversity 41:87–107, https://doi.org/10.1007/s12526-010-0078-4.

Bluhm, B., R. Gradinger, and S. Schnack-Schiel. 2010a. Sea ice meio- and macrofauna. Pp. 357–393 in Sea Ice. D. Thomas and G. Dieckmann, eds, Wiley-Blackwell, New York.

Bluhm, B.A., K. Iken, S.L. Mincks, B.I. Sirenko, and B.A. Holladay. 2009. Community structure of epibenthic megafauna in the Chukchi Sea. Aquatic Biology 7:269–293, https://doi.org/10.3354/ab00198.

Bluhm, B.A., D. Watts, and F. Huettmann. 2010b. Free database availability, metadata and the Internet: An example of two high latitude components of the Census of Marine Life. Pp. 233–244 in Spatial Complexity, Informatics and Wildlife Conservation. F. Huettmann and S.A. Cushman, eds, Springer, Tokyo.

Brinkmeyer, R., K. Knittel, J. Juergens, H. Weyland, R. Amann, and E. Helmke. 2003. Diversity and structure of bacterial communities in Arctic versus Antarctic pack ice. Applied Environmental Microbiology 69:6,610–6,619, https://doi.org/10.1128/AEM.69.11.6610-6619.2003.

Born, E.W. 2005. An assessment of the effects of hunting and climate on walruses in Greenland. PhD thesis, University of Oslo, Oslo.

Brodeur, R.D., M.B. Decker, L. Ciannelli, J.E. Purcell, N.A. Bond, P.J. Stabeno, E. Acuna, and G.L. Hunt Jr. 2008. Rise and fall of jellyfish in the eastern Bering Sea in relation to climate regime shifts. Progress in Oceanography 77:103–111, https://doi.org/10.1016/j.pocean.2008.03.017.

Bubnova, E.N. 2010. Fungal diversity in bottom sediments of the Kara Sea. Botanica Marina 53:595–600, https://doi.org/10.1515/BOT.2010.063.

Bucklin, A., R.R. Hopcroft, K.N. Kosobokova, L.M. Nigro, B.D. Ortman, R.M. Jennings, and C.J. Sweetman. 2010. DNA barcoding of Arctic Ocean holozooplankton for species identification and recognition. Deep-Sea Research Part II 57:40–48, https://doi.org/10.1016/j.dsr2.2009.08.005.

Buzhinskaja, G., ed. 2011. Illustrated Keys to Free-Living Invertebrates of Eurasian Arctic Seas and Adjacent Deep Waters, vol. 2. Nemertea, Cephalorhyncha, Oligochaeta, Hirudinida, Pogonophora, Echiura, Sipuncula, Phoronida, and Brachiopoda. Alaska Sea Grant, University of Alaska Fairbanks, Fairbanks, AK.

CAFF (Conservation of Arctic Flora & Fauna). 2001. Arctic Flora and Fauna: Status and Conservation. Edita Helsinki, 278 pp.

CAFF. 2010. Arctic Biodiversity Trends 2010: Selected Indicators of Change. CAFF International Secretariat, Akureyri, Iceland, 121 pp.

Carlson, D. 2011. A lesson in sharing. Nature 469:293, https://doi.org/10.1038/469293a.

Carr, C.M. 2010. The Polychaeta of Canada: Exploring diversity and distribution patterns using DNA barcodes. MSc Thesis, University of Guelph, Guelph, Ontario.

Carr, C.M. 2011. Polychaete diversity and distribution patterns in Canadian marine waters. Marine Biodiversity, https://doi.org/10.1007/s12526-011-0095-y.

Carroll, M.L., and J. Carroll. 2003. The Arctic Seas. Pp. 127–156 in Biogeochemistry of Marine Systems. K.D. Black and G.B. Shimmield, eds, CRC Press, Boca Raton, FL.

Carstens, J., D. Hebbeln, and G. Wefer. 1997. Distribution of planktic foraminifera at the ice margin in the Arctic (Fram Strait). Marine Micropaleontology 29:257–269, https://doi.org/10.1016/S0377-8398(96)00014-X.

Carstens, J., and G. Wefer. 1992. Recent distribution of planktonic foraminifera in the Nansen Basin, Arctic Ocean. Deep-Sea Research Part A 39:S507–S524, https://doi.org/10.1016/S0198-0149(06)80018-X.

Chaban, E.M. 2008. Opisthobranchiate mollusca of the orders Cephalaspidea, Thecosomata and Gymnosomata (Mollusca, Opisthobranchia) of the Chukchi Sea and the Bering Strait. Pp. 149–162 in Fauna and Zoogeography of Benthos of the Chukchi Sea. B.I. Sirenko and S.V. Vassilenko, eds, Explorations of the Fauna of the Seas 61(69).

Collins, R.E., G. Rocap, and J.W. Deming. 2010. Persistence of bacterial and archaeal communities in sea ice through an Arctic winter. Environmental Microbiology 12:1,828–1,841.

Corstorphine, E.A. 2010. DNA barcoding of echinoderms: Species diversity and patterns of molecular evolution. MSc Thesis, University of Guelph, Guelph, Ontario.

Coyle, K.O., B.A. Bluhm, B. Konar, A. Blanchard, and R.C. Highsmith. 2007. Amphipod prey of gray whales in the northern Bering Sea: Comparison of biomass and distribution between the 1980s and 2002–2003. Deep-Sea Research Part II 54:2,906–2,918, https://doi.org/10.1016/j.dsr2.2007.08.026.

Daugbjerg, N., and Ø. Moestrup. 1993. Four new species of Pyramimonas (Prasinophyceae) from arctic Canada including a light and electron microscopic description of Pyramimonas quadrifolia sp. nov. European Journal of Phycology 28:3–16, https://doi.org/10.1080/09670269300650021.

DeBroyer, C., and B. Danis. 2011. How many species in the Southern Ocean? Towards a dynamic inventory of the Antarctic marine species. Deep-Sea Research Part II 58:5–17, https://doi.org/10.1016/j.dsr2.2010.10.007.

Divoky, G.J. 1982. The occurrence and behavior of nonbreeding Horned Puffins at Black Guillemot colonies in northern Alaska. Wilson Bulletin 94:356–358.

Dunton, K.H., J.L. Goodall, S.V. Schonberg, J.M. Grebmeier, and D.R. Maidment. 2005. Multi-decadal synthesis of benthic-pelagic coupling in the western Arctic: Role of cross-shelf advective processes. Deep-Sea Research Part II 52:3,462–3,477, https://doi.org/10.1016/j.dsr2.2005.09.007.

Dunton, K.H., E.K. Reimnitz, and S. Schonberg. 1982. An Arctic kelp community in the Alaskan Beaufort Sea. Arctic 35:465–484.

Ebert, D.A., W.T. White, K.J. Goldman, L.J.V. Compagno, T.S. Daly Engel, and R.D. Ward. 2010. Resurrection and redescription of Squalus suckleyi (Girard, 1854) from the North Pacific, with comments on the Squalus acanthias subgroup (Squaliformes: Squalidae). Zootaxa 2,612:2,240.

Feder, H.M., S.C. Jewett, and A. Blanchard. 2005. Southeastern Chukchi Sea (Alaska) epibenthos. Polar Biology 28:402–421, https://doi.org/10.1007/s00300-004-0683-4.

Ferguson, S.H. 2009. Killer whales on the rise in the Canadian Arctic. The Circle 4:20–23.

Gagaev, S.Y. 2009. Terebellides irinae sp. n.: A new species of the genus Terebellides (Polychaeta, Terebellidae) from the Canada Basin of the Arctic Ocean. Russian Journal of Marine Biology 35:73–75, https://doi.org/10.1134/S1063074009060042.

Gaston, A.J. 2004. Seabirds: A Natural History. Yale University Press, London, 224 pp.

Gaston, K.J. 2010. Biodiversity. Pp. 27–44 in Conservation Biology for All. J.S. Sodhi and P.R. Ehrlich, eds, Oxford University Press.

George, J.C., J. Zeh, R. Suydam, and C. Clark. 2004. Abundance and population trend (1978–2001) of western arctic bowhead whales surveyed near Barrow, Alaska. Marine Mammal Science 20:755–773, https://doi.org/10.1111/j.1748-7692.2004.tb01191.x.

Gilchrist, H.G., and M.L. Mallory. 2005. Declines in abundance and distribution of the ivory gull (Pagophila eburnea) in Arctic Canada. Biological Conservation 121:303–309, https://doi.org/10.1016/j.biocon.2004.04.021.

Gill, M.J., K. Crane, R. Hindrum, P. Arneberg, I. Bysveen, N.V. Denisenko, V. Gofman, A. Grant-Friedman, G. Gudmundsson, R.R. Hopcroft, and others. 2011. Arctic Marine Biodiversity Monitoring Plan (CBMP-Marine Plan). Final Draft, Circumpolar Biodiversity Monitoring Program, Environment Canada, 174 pp.

Glud, R.N., M. Kühl, F. Wenzhöfer, and S. Rysgaard. 2002. Benthic diatoms of a high Arctic fjord (Young Sound, NE Greenland): Importance for ecosystem primary production. Marine Ecology Progress Series 238:15–29, https://doi.org/10.3354/meps238015.

Gosselin, M., M. Levasseur, P.A. Wheeler, R.A. Horner, and B.C. Booth. 1997. New measurements of phytoplankton and ice algal production in the Arctic Ocean. Deep-Sea Research Part II 44:1,623–1,644, https://doi.org/10.1016/S0967-0645(97)00054-4.

Gradinger, R. 2009. Sea-ice algae: Major contributors to primary production and algal biomass in the Chukchi and Beaufort Seas during May/June 2002. Deep-Sea Research Part II 56:1,201–1,212, https://doi.org/10.1016/j.dsr2.2008.10.016.

Gradinger, R., B.A. Bluhm, R.R. Hopcroft, A. Gebruk, K.N. Kosobokova, B. Sirenko, and J.M. Węsławski. 2010a. Chapter 10. Marine life in the Arctic. Pp. 183–202 in Life in the World’s Ocean: Diversity, Distribution, Abundance. A. McIntyre, ed., Wiley-Blackwell.

Gradinger, R., B.A. Bluhm, and K. Iken. 2010b. Arctic sea ice ridges: Safe havens for sea ice fauna during periods of extreme ice melt? Deep-Sea Research Part II 57:86–95, https://doi.org/10.1016/j.dsr2.2009.08.008.

Gradinger, R., M.R. Kaufman, and B.A. Bluhm. 2009. The pivotal role of sea ice sediments for the seasonal development of near-shore Arctic fast ice biota. Marine Ecology Progress Series 394:49–63, https://doi.org/10.3354/meps08320.

Grebmeier, J.M., L.W. Cooper, H.M. Feder, and B. Sirenko. 2006a. Ecosystem dynamics of the Pacific-influenced Northern Bering and Chukchi Seas in the Amerasian Arctic. Progress in Oceanography 71:331–361, https://doi.org/10.1016/j.pocean.2006.10.001.

Grebmeier, J.M., J.E. Overland, S.E. Moore, E.V. Farley, E.C. Carmack, L.W. Cooper, K.E. Frey, J.H. Helle, F.A. McLaughlin, and S.L. McNutt. 2006b. A major ecosystem shift in the northern Bering Sea. Science 311:1,461–1,464, https://doi.org/10.1126/science.1121365.

Grebmeier, J.M., S.E. Moore, J.E. Overland, K.E. Frey, and R.R. Gradinger. 2010. Biological response to recent Pacific Arctic sea ice retreats. Eos, Transactions, American Geophysical Union 91:161–162, https://doi.org/10.1029/2010EO180001.

Hamazaki, T., L. Fair, L. Watson, and E. Brennan. 2005. Analyses of Bering Sea bottom-trawl surveys in Norton Sound: Absence of regime shift effect on epifauna and demersal fish. ICES Journal of Marine Science 62:1,597–1,602, https://doi.org/10.1016/j.icesjms.2005.06.003.

Hardy, S.M., C.M. Carr, M. Hardman, D. Steinke, E. Corstorphine, and C. Mah. 2011. Molecular diversity of Arctic fauna: Genetic tools provide new insights in taxonomy and biogeography of marine organisms. Marine Biodiversity 41:195–210, https://doi.org/10.1007/s12526-010-0056-x.

Hegseth, E.N., and A. Sundfjord. 2008. Intrusion and blooming of Atlantic phytoplankton species in the high Arctic. Journal of Marine Systems 74:108–119, https://doi.org/10.1016/j.jmarsys.2007.11.011.

Hop, H., and O. Pavlova. 2008. Distribution and biomass transport of ice amphipods in drifting sea ice around Svalbard. Deep-Sea Research Part I 55:2,292–2,307, https://doi.org/10.1016/j.dsr2.2008.05.023.

Hopcroft, R.R., K.N. Kosobokova, and A.I. Pinchuk. 2010. Zooplankton community patterns in the Chukchi Sea during summer 2004. Deep-Sea Research Part II 57:27–39, https://doi.org/10.1016/j.dsr2.2009.08.003.

Horner, R., and G.C. Schrader. 1982. Relative contributions of ice algae, phytoplankton, and benthic microalgae to primary production in nearshore regions of the Beaufort Sea. Arctic 35:485–503.

Huettmann, F., and S. Hazlett. 2010. Changing the Arctic: Adding immediate protection to the equation. Alaska Park Science 8:95–96.

Huettmann, F., Y. Artukhin, O. Gilg, and G. Humphries. 2011. Predictions of 27 pelagic seabird distributions using public environmental variables, assessed with compiled colony data: A first digital IPY and GBIF Open Access synthesis platform. Marine Biodiversity 41:141–179, https://doi.org/10.1007/s12526-011-0083-2.

Huntingon, H.P., and S.E. Moore. 2008. Assessing the impacts of climate change on Arctic marine mammals. Ecological Applications 18:S1–S2, https://doi.org/10.1890/06-0282.1.

Itaki, T., M. Ito, H. Narita, N. Ahagon, and H. Sakai. 2003. Depth distribution of radiolarians from the Chukchi and Beaufort Seas, western Arctic. Deep-Sea Research Part I 50:1,507–1,522, https://doi.org/10.1016/j.dsr.2003.09.003.

Jakobsson, M., A. Grantz, Y. Kristoffersen, and R. Macnab. 2004. Physiography and bathymetry of the Arctic Ocean. Pp. 1–6 in The Organic Carbon Cycle in the Arctic Ocean. R. Stein and R. MacDonald, eds, Springer, Berlin.

Johnsen, K.I., B. Alfthan, L. Hislop, and J.F. Skaalvik, eds. 2010. Protecting Arctic Biodiversity. United Nations Environment Programme, GRID-Arendal, Norway. Available online at: http://www.grida.no (accessed July 20, 2011).

Junge, K., H. Eicken, and J.W. Deming. 2004. Bacterial activity at –2 to –20°C in Arctic wintertime sea ice. Applied Environmental Microbiology 70:550–555, https://doi.org/10.1128/AEM.70.1.550-557.2004.

Kędra, M., M. Włodarska-Kowalczuk, and J.M. Węsławski. 2010. Decadal change in macrobenthic soft-bottom community structure in a high Arctic fjord (Kongsfjorden, Svalbard). Polar Biology 33:1–11, https://doi.org/10.1007/s00300-009-0679-1.

Knudsen, S.W., and P.R. Møller. 2008. Careproctus kidoi, a new Arctic species of snailfish (Teleostei: Liparidae) from Baffin Bay. Ichthyological Research 55:175–182, https://doi.org/10.1007/s10228-007-0034-x.

Kosobokova, K.N., H-J. Hirche, and R.R. Hopcroft. 2011. Patterns of zooplankton diversity through the depths of the Arctic’s central basins. Marine Biodiversity 41:29–50, https://doi.org/10.1007/s12526-010-0057-9.

Kovacs, K.M., S. Moore, C. Lydersen, and J.E. Overland. 2011. Impacts of changing sea ice conditions on Arctic marine mammals. Marine Biodiversity 41:181–194, https://doi.org/10.1007/s12526-010-0061-0.

Krasnov, Y.V., and R.T. Barrett. 1997. The first record of North Atlantic Gannets Morus bassanus breeding in Russia. Seabird 19:54–57.

Kuklinski, P., and P.D. Taylor. 2008. Arctic species of the cheilostome bryozoan Microporealla, with a redescription of the type species. Journal of Natural History 42:2,893–2,906, https://doi.org/10.1080/00222930802126904.

Laakmann, S., M. Kochzius, and H. Auel. 2009. Ecological niches of Arctic deep-sea copepods: Vertical partitioning, dietary preferences and different trophic levels minimize inter-specific competition. Deep-Sea Research Part I 56:741–756, https://doi.org/10.1016/j.dsr.2008.12.017.

Laidre, K.L., I. Stirling, L. Lowry, Ø. Wiig, M.P. Heide-Jørgensen, and S. Ferguson. 2008. Quantifying the sensitivity of arctic marine mammals to climate-induced habitat change. Ecological Applications 18:S97–S125, https://doi.org/10.1890/06-0546.1.

Li, H., Y. Yu, W. Luo, Y. Zeng, and B. Chen. 2009. Bacterial diversity in surface sediments from the Pacific Arctic Ocean. Extremophiles 13:233–246, https://doi.org/10.1007/s00792-009-0225-7.

Lønne, O.J., and G.W. Gabrielsen. 1992. Summer diet of seabirds feeding in sea-ice-covered waters near Svalbard. Polar Biology 12:685–692.

Lovejoy, C., E. Pierre, P.E. Galand, and D.L. Kirchman. 2011. Picoplankton diversity in the Arctic Ocean and surrounding seas. Marine Biodiversity 41:5–12, https://doi.org/10.1007/s12526-010-0062-z.

Mace, G.M., W. Cramer, S. Diaz, D.P. Faith, A. Larigauderie, P. Le Prestre, M. Palmer, C. Perrings, R.J. Scholes, M. Walpole, and others. 2010. Biodiversity targets after 2010. Current Opinion in Environmental Sustainability 2:1–6, https://doi.org/10.1016/j.cosust.2010.03.003.

Magurran, E.M. 2004. Measuring Biological Diversity. Blackwell Publishing, Malden, MA, 256 pp.

Mathieson, A.C., G.E. Moore, and F.T. Short. 2010. A floristic comparison of seaweeds from James Bay and three contiguous northeastern Canadian Arctic sites. Rhodora 112:396–433, https://doi.org/10.3119/09-12.1.

Mecklenburg, C.W., P.R. Møller, and D. Steinke. 2011. Biodiversity of Arctic marine fishes: Taxonomy and zoogeography. Marine Biodiversity 41:109–140, https://doi.org/10.1007/s12526-010-0070-z.

Mecklenburg, C.W., D.L. Stein, B.A. Sheiko, N.V. Chernova, T.A. Mecklenburg, and B.A. Holladay. 2007. Russian-American long-term census of the Arctic: Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004. Northwestern Naturalist 88:168–187, https://doi.org/10.1898/1051-1733(2007)88[168:RLCOTA]2.0.CO;2.

Moline, M.A., N.J. Karnovsky, Z. Brown, G.J. Divoky, T.K. Frazer, C.A. Jacoby, J.J. Torres, and W.R. Fragser. 2008. High latitude changes in ice dynamics and their impact on polar marine ecosystems. Annals of the New York Academy of Sciences 1,134:267–319, https://doi.org/10.1196/annals.1439.010.

Møller, P.R. 2001. Redescription of the Lycodes pallidus species complex (Pisces, Zoarcidae), with a new species from the Arctic/North Atlantic Ocean. Copeia 2001:972–996, https://doi.org/10.1643/0045-8511(2001)001[0972:ROTLPS]2.0.CO;2.

Moore, S.E. 2008. Marine mammals as ecosystem sentinels. Journal of Mammology 89:534–540, https://doi.org/10.1644/07-MAMM-S-312R1.1.

Moore, S.E., D.P. DeMaster, and P.K. Dayton. 2000. Cetacean habitat selection in the Alaskan Arctic during summer and autumn. Arctic 53:432–447.

Moore, S.E., J.M. Grebmeier, and J.R. Davies. 2003. Gray whale distribution relative to forage habitat in the northern Bering Sea: Current conditions and retrospective summary. Journal of Zoology 81:734–742, https://doi.org/10.1139/z03-043.

Mueter, F.J., and M.A. Litzow. 2008. Sea ice retreat alters the biogeography of the Bering Sea continental shelf. Ecological Applications 18:309–320, https://doi.org/10.1890/07-0564.1.

Naidu, A.S. 1988. Marine surficial sediments. Section 1.4 in Bering, Chukchi and Beaufort Seas: Coastal and Ocean Zones Strategic Assessment Data Atlas. NOAA Strategic Assessment Branch, Ocean Assessment Division, Washington, DC.

Norcross, B.L., B.A. Holladay, M.S. Busby, and K.L. Mier. 2010. Demersal and larval fish assemblages in the Chukchi Sea. Deep-Sea Research Part II 57:57–70, https://doi.org/10.1016/j.dsr2.2009.08.006.

Nozais, C., M. Gosselin, C. Michel, and G. Tita. 2001. Abundance, biomass, composition and grazing impact of the sea-ice meiofauna in the North Water, northern Baffin Bay. Marine Ecology Progress Series 217:235–250, https://doi.org/10.3354/meps217235.

O’Corre-Crowe, G.M. 2008. Climate change and the molecular ecology of Arctic marine mammals. Ecological Applications 18:S56–S76.

Oppel, S., and F. Huettmann, 2010. Using a Random Forest model and public data to predict the distribution of prey for marine wildlife management. Pp. 151–163 in Spatial Complexity, Informatics, and Wildlife Management. S.A. Cushman and F. Huettmann, eds, Springer, https://doi.org/10.1007/978-4-431-87771-4_8.

Orensanz, J., B. Ernst, D.A. Armstrong, P. Stabeno, and P. Livingston. 2004. Contraction of the geographic range of distribution of snow crab (Chionoecetes opilio) in the Eastern Bering Sea: An environmental ratchet? CalCOFI Report 45:65–79.

Pang, K.-L., R.K.K. Chow, C.-W. Chan, and L.L.P. Vrijmoed. 2011. Diversity and physiology of marine lignicolous fungi in Arctic waters: A preliminary account. Polar Research 30, https://doi.org/10.3402/polar.v30i0.5859.

Pasternak, A., E. Arashkevich, K. Tande, and T. Falkenhaug. 2001. Seasonal changes in feeding, gonad development and lipid stores in Calanus finmarchicus and C. hyperboreus from Malangen, northern Norway. Marine Biology 138:1,141–1,152, https://doi.org/10.1007/s002270100553.

Perovich, D.K., S.V. Nghiem, T. Markus, and A. Schweiger. 2007. Seasonal evolution and interannual variability of the local solar energy absorbed by the Arctic sea-ice–ocean system. Journal of Geophysical Research 112, C03005, https://doi.org/10.1029/2006JC003558.

Pertsova, N.M., and K.N. Kosobokova. 2010. Interannual and seasonal variation of the population structure, abundance, and biomass of the Arctic copepod Calanus glacialis in the White Sea. Oceanology 50:531–541, https://doi.org/10.1134/S0001437010040090.

Petersen, A., D. Irons, T. Anker-Nilssen, Y. Artukhin, R. Barrett, D. Boertmann, C. Egevang, M.G. Gavrilo, G. Gilchrist, M. Hario, and others. 2008. Framework for a Circumpolar Arctic Seabird Monitoring Network. CAFF CBMP Report No. 15. CAFF International Secretariat, Akureyri, Iceland, 67 pp.

Petersen, M.R., J.B. Grand, and C.P. Dau. 2000. Spectacled eider: Somateria fischeri. A. Poole and F. Gill, eds, The Birds of North America Inc., no. 57, Philadelphia, PA, 24 pp.

Piatt, J.F., and A.S. Kitaysky. 2002. Tufted Puffin: Fratercula cirrata. The Birds of North America Online, A. Poole, ed., Cornell Laboratory of Ornithology, Ithaca, NY. Available online at: http://bna.birds.cornell.edu/bna/species/708 (accessed June 21, 2011).

Piepenburg, D., P. Archambault, W.G. Ambrose Jr., A. Blanchard, B.A. Bluhm, M.L. Carroll, K. Conlan, M. Cusson, H.M. Feder, J.M. Grebmeier, and others. 2011. Towards a pan-Arctic inventory of the species diversity of the macro- and megabenthic fauna of the Arctic shelf seas. Marine Biodiversity 41:51–70, https://doi.org/10.1007/s12526-010-0059-7.

Piraino, S., B.A. Bluhm, R. Gradinger, and F. Boero. 2008. Sympagohydra tuuli gen. nov. and sp. nov. (Cnidaria: Hydrozoa) a cool hydroid from the Arctic sea ice. Journal of the Marine Biological Association of the United Kingdom 88:1,637–1,641, https://doi.org/10.1017/S0025315408002166.

Poulin, M., N. Daugbjerg, R. Gradinger, L. Ilyash, T. Ratkova, and C. von Quillfeldt. 2011. The pan-Arctic biodiversity of marine phytoplankton and sea-ice unicellular eukaryotes: A first-attempt assessment. Marine Biodiversity 41:13–28, https://doi.org/10.1007/s12526-010-0058-8.

Raskoff, K.A. 2010. Bathykorus bouilloni: A new genus and species of deep-sea jellyfish from the Arctic Ocean (Hydrozoa, Narcomedusae, Aeginidae). Zootaxa 2361:57–67.

Reid, P.C., D.G. Johns, M. Edwards, M. Starr, M. Poulin, and P. Snoeijs. 2007. A biological consequence of reducing Arctic ice cover: Arrival of the Pacific diatom Neodenticula seminae in the North Atlantic for the first time in 800,000 years. Global Change Biology 13:1,910–1,921, https://doi.org/10.1111/j.1365-2486.2007.01413.x.

Renaud, P.E., M. Włodarska-Kowalczuk, H. Trannum, B. Holte, J.M. Weslawski, S. Cochrane, S. Dahle, and B. Gulliksen. 2007. Multidecadal stability of benthic community structure in a high-Arctic glacial fjord (van Mijenfjord, Spitsbergen). Polar Biology 30:295–305, https://doi.org/10.1007/s00300-006-0183-9.

Rogacheva, A.V. 2007. Revision of the Arctic group of species of the family Elpidiidae (Elasipodida, Holothuroidea). Marine Biology Research 3:376–396, https://doi.org/10.1080/17451000701781880.

Rogacheva, A.V. 2011. Taxonomy and distribution of the genus Kolga Danielssen and Koren, 1879 (Elpidiidae, Holothuroidea, Echinodermata). Journal of the Marine Biological Association of the United Kingdom, https://doi.org/10.1017/S0025315411000427.

Ronowicz, M., and P. Schuchert. 2007. Halecium arcticum (Cnidaria: Hydrozoa), a new species of hydroid from Spitsbergen. Zootaxa 1,549:55–62.

Sakshaug, E. 2004. Primary and secondary production in the Arctic Seas. Pp. 57–82 in The Organic Carbon Cycle in the Arctic Ocean. R. Stein and R. MacDonald, eds, Springer, Berlin.

Sirenko, B.I. 2001. List of species of free-living invertebrates of Eurasian Arctic seas and adjacent deep waters. Explorations of the Fauna of the Seas 51:1–129.

Sirenko B.I. 2009. The prosobranchs of the gastropods (Mollusca, Gastropoda, Prosobranchia) of the Chukchi Sea and Bering Strait, their species composition and distribution. Explorations of the Fauna of the Seas 64(72):104–153.

Sirenko, B.I., and S.Y. Gagaev. 2007. Unusual abundance of macrobenthos and biological invasions in the Chukchi Sea. Russian Journal of Marine Biology 33:355–364, https://doi.org/10.1134/S1063074007060016.

Sirenko, B.I., C. Clarke, R.R. Hopcroft, F. Huettmann, B.A. Bluhm, and R. Gradinger, eds. 2010. The Arctic Register of Marine Species (ARMS) Compiled by the Arctic Ocean Diversity (ArcOD) Project. Available online at: http://www.marinespecies.org/arms (accessed February 12, 2011).

Soltwedel, T., E. Bauerfeind, M. Bergmann, N. Budaeva, E. Hoste, N. Jaeckisch, K. von Juterzenka, J. Matthiessen, V. Mokievsky, E.-M. Nöthig, and others. 2005. HAUSGARTEN, multidisciplinary investigations at a deep-sea, long-term observatory in the Arctic Ocean. Oceanography 18(3):46–61. [pdf]

Stockwell, D.A., T.E. Whitledge, S.I. Zeeman, K.O. Coyle, J.M. Napp, R.D. Brodeur, A.I. Pinchuk, and G.L. Hunt. 2001. Anomalous conditions in the south-eastern Bering Sea, 1997: Nutrients, phytoplankton and zooplankton. Fisheries Oceanography 10:99–116, https://doi.org/10.1046/j.1365-2419.2001.00158.x.

Sun, M.-Y., L.M. Clough, M.L. Carroll, J. Dai, W.G. Ambrose Jr., and G.R. Lopez. 2009. Different responses of two common Arctic macrobenthic species (Macoma balthica and Monoporeia affinis) to phytoplankton and ice algae: Will climate change impacts be species specific? Journal of Experimental Marine Biology and Ecology 376:110–121, https://doi.org/10.1016/j.jembe.2009.06.018.

Suydam, R.S., D.L. Dickson, J.B. Fadely, and L.T. Quakenbush. 2000. Population declines of King and Common eiders of the Beaufort Sea. Condor 102:219–222, https://doi.org/10.1650/0010-5422(2000)102[0219:PDOKAC]2.0.CO;2.

Tchesunov, A.V., and F. Riemann. 1995. Arctic sea ice nematodes (Monhysteroidea) with descriptions of Cryonemacrassum gen. n. sp. n. and C. tenue. Nematologica 41:35–50, https://doi.org/10.1163/003925995X00035.

Tett, P., and E.D. Barton. 1995. Why are there about 5000 species of phytoplankton in the sea? Journal of Plankton Research 17:1,693–1,704, https://doi.org/10.1093/plankt/17.8.1693.

Thomas, D., and G. Dieckmann, eds. 2009. Sea Ice: An Introduction to its Physics Chemistry Biology and Geology. Wiley-Blackwell, New York, NY, 416 pp.

Vassilenko, S.V., and V.V. Petryashov, eds. 2009. Illustrated Keys to Free-Living Invertebrates of Eurasian Arctic Seas and Adjacent Deep Waters, vol. 1. Rotifera, Pycnogonida, Cirripedia, Leptostraca, Mysidacea, Hyperiidea, Caprellidea, Euphausiacea, Dendrobranchiata, Pleocyemata, Anomura, and Brachyura. Alaska Sea Grant, University of Alaska Fairbanks, 192 pp.

Vermeij, G.J., and P.D. Roopnarine. 2008. The coming Arctic invasion. Science 321:780–781, https://doi.org/10.1126/science.1160852.

Vinogradova, N.G. 1997. Zoogeography of the abyssal and hadal zones. Advances in Marine Biology 32:326–387, https://doi.org/10.1016/S0065-2881(08)60019-X.

Wassmann, P., C.M. Duarte, S. Agusti, and M.K. Sejr. 2011. Footprints of climate change in the Arctic marine ecosystem. Global Change Biology 17:1,235–1,249, https://doi.org/10.1111/j.1365-2486.2010.02311.x.

Wassmann, P., M. Reigstad, T. Haug, B. Rudels, M.L. Carroll, H. Hop, G.W. Gabrielsen, S. Falk-Petersen, S.G. Denisenko, E. Arashkevich, and others. 2006. Food webs and carbon flux in the Barents Sea. Progress in Oceanography 71:232–287, https://doi.org/10.1016/j.pocean.2006.10.003.

Wei, C.-K., G.T. Rowe, E. Escobar-Briones, A. Boetius, T. Soltwedel, M.J. Caley, Y. Soliman, F. Huettmann, F. Qu, Z. Yu, and others. 2010. Global patterns and predictions of seafloor biomass using random forests. PLoS ONE 5(12):e15323, https://doi.org/10.1371/journal.pone.0015323.

Wesławski, J.M., M.A. Kendall, M. Włodarska-Kowalczuk, K. Iken, M. Kędra, J. Legezynska, and M.K. Sejr. 2011. Arctic fjord and coastal macrobenthic biodiversity. Marine Biodiversity 41:71–86.

Węsławski, J.M., S. Kwaśniewski, and L. Stempniewicz. 2008. Warming in the Arctic may result in the negative effects of increased biodiversity. Polarforschung 78:105–108.

Wesławski, J.M., J. Wiktor Jr., and L. Kotwicki. 2010. Increase in biodiversity in the arctic rocky littoral, Sorkappland, Svalbard, after 20 years of climate warming. Marine Biodiversity 40:123–130.

Wesławski, J.M., J. Wiktor, M. Zajączkowski, and S. Swerpel. 1993. Intertidal zone of Svalbard. 1. Macroorganism distribution and biomass. Polar Biology 13:73–108.

Wiencke, C., M.N. Clayton, I. Gomez, K. Iken, U.H. Lüder, C.D. Amsler, U. Karsten, D. Hanelt, K. Bischof, and K. Dunton. 2007. Life strategy, ecophysiology and ecology of seaweeds in polar waters. Review in Environmental Science and Biotechnology, https://doi.org/10.1007/s11157-006-9106-z.

Wilce, R.T. 1990. Role of the Arctic Ocean as a bridge between the Atlantic and Pacific Oceans: Fact and hypothesis. Pp. 323–348 in Evolutionary Biogeography of the Marine Algae of the North Atlantic. D.J. Garbary and G.R. South, eds, Springer-Verlag.

Wilce, R.T. 1994. The Arctic subtidal as habitat for macrophytes. Pp. 89–91 in Seaweed Ecology and Physiology. C.S. Lobban and P.J. Harrison, eds, Cambridge University Press, Cambridge.

Wilce, R.T., P.M. Pedersen, and S. Sekida. 2009. Chukchi pedicellata gen. et sp. nov. and C. endophytica nov. comb., Arctic endemic brown algae (Phaeophyceae). Journal of Phycology 45:272–286, https://doi.org/10.1111/j.1529-8817.2008.00631.x.

Wulff, A., K. Iken, M.L. Quartino, A. Al-Handal, C. Wiencke, and M.N. Clayton. 2009. Biodiversity, biogeography and zonation of marine benthic micro- and macroalgae in the Arctic and Antarctic. Botanica Marina 52:491–507, https://doi.org/10.1515/BOT.2009.072.

Wyllie-Echeverria, T., and W.S. Wooster. 1998. Year to year variations in Bering Sea ice cover and some consequences for fish distributions. Fisheries Oceanography 7:159–170, https://doi.org/10.1046/j.1365-2419.1998.00058.x.

Zoeckler, C. 2011. In press. Chapter 9: Status, threat and protection of Arctic waterbirds. In Protection of the Three Poles. F. Huettmann, ed., Springer Japan.

Copyright & Usage

This is an open access article made available under the terms of the Creative Commons Attribution 4.0 International License (https://creativecommons.org/licenses/by/4.0/), which permits use, sharing, adaptation, distribution, and reproduction in any medium or format as long as users cite the materials appropriately, provide a link to the Creative Commons license, and indicate the changes that were made to the original content. Images, animations, videos, or other third-party material used in articles are included in the Creative Commons license unless indicated otherwise in a credit line to the material. If the material is not included in the article’s Creative Commons license, users will need to obtain permission directly from the license holder to reproduce the material.