The polar bear (Ursus maritimus) is the apex predator of the Arctic but its distribution throughout the Pleistocene and Holocene has not previously been reported. Although natural death specimens of this species (‘fossils’) are rare, archaeological remains are much more common. This historical compilation presents the record of known ancient polar bear remains from fossil and archaeological contexts before AD 1910. Most remains date within the Holocene and derive from human habitation sites within the modern range of the species, with extralimital specimens documented in the north Atlantic during the late Pleistocene and in the southern Bering Sea during the middle Holocene reflecting natural expansions of sea ice during known cold periods. The single largest polar bear assemblage was recovered from an archaeological site on Zhokhov Island, Russia, occupied ca. 8,250–7,800 a BP during the warmer-than-today Holocene Climatic Optimum: 5,915 polar bear bones were recovered, representing 28% of all remains identified. Polar bear fossils and archaeological remains across the Arctic are most often found in proximity to areas where polynyas (recurring areas of thin ice or open water) are known today and which likely occurred in the past, including for the oldest known fossil from Svalbard (ca. 130–115 k a BP) and the oldest known archaeological specimens from Zhokhov Island (ca. 8,000 a BP). This pattern indicates that as they do today, polar bears may have been most commonly found near polynyas throughout their known historical past because of their need for ice-edge habitats at which to hunt seals.
The polar bear (Ursus maritimus) personifies the Arctic: it is the quintessential species of the northern sea ice habitat. It is usually classified as a marine mammal because individuals can (and often do) spend their entire lives on sea ice. However, females that make maternity dens on the coast may spend up to 8 months at a time on land and many bears in some regions spend at least a few months on land during the ice-free season (Amstrup 2003; Andersen et al. 2012; Castro de la Guardia et al. 2017; Ramsay & Stirling 1988; Rode et al. 2015; Stirling 1997). The species is currently well distributed across the shallow peripheral seas of the Arctic (Chukchi, Beaufort, Barents, Kara and Laptev) but also occurs in sub-Arctic regions with seasonal sea ice in winter and spring (including Hudson Bay, Labrador Sea, Davis Strait, Denmark Strait, and the Bering Sea) (Figure 1).
The most carnivorous and predatory of all bears, the polar bear occupies the top of the Arctic food chain, subsisting primarily on ringed seals (Phoca hispida) and to a lesser degree on bearded seals (Erignathus barbatus), which have a similar circumpolar distribution (Amstrup 2003). However, polar bears also occasionally hunt other Arctic seal species, walrus (Odobenus rosmarus), and small Arctic whales (Heide-Jørgensen et al. 2002; Kochnev 2002; Pereverzev & Kochnev 2012; Thiemann et al. 2007) and will readily scavenge the natural-death or human-hunted carcasses of walrus and large whales (Kavry et al. 2006; Laidre et al. 2018).
Apex predators like polar bears have virtually no natural enemies aside from humans. As a consequence, polar bears are either killed by humans or die a natural death. By far the most common cause of death for polar bears is starvation, which is a natural consequence of injury, illness, old age, lack of hunting experience, and intra-species competition (Amstrup 2003; Calvert et al. 1986; DeMaster, Kingsley & Stirling 1980; Derocher & Stirling 1992; Derocher & Stirling 1995; Ramsay & Stirling 1988; Stirling 1974; Stirling 2002; Stirling & Lunn 1997). These deaths usually occur during the winter when bears are on the sea ice, which means skeletal remains eventually sink to the bottom, never to be found. Very rarely, a polar bear may die of starvation or be killed by another bear on land during the summer/fall ice-free season or a pregnant or post-partum female may die on land in her maternity snow den over the winter, but scavenger activity ensures few skeletal remains survive. For these reasons, skeletal remains of polar bears that have died a natural death are rarely found as paleontological specimens unless they are quickly buried. In this regard, the polar bear stands in marked contrast to its ancestral species, the terrestrial-dwelling brown bear (Ursus arctos) which has a rich fossil record (Barnes et al. 2002; Davison et al. 2011; Edwards et al. 2011; Edwards et al. 2014; Kurtén 1968; Kurtén 1988).
However, as polar bears were hunted by humans across the entire Arctic during the Holocene, archaeological remains of polar bears are much more plentiful and provide the primary historical perspective on the distribution and range of the species since the end of the Last Glacial Maximum (LGM, ca. 11,700 a BP) (Table 1). A number of archaeologists have pointed out that proximity to polynyas may explain the location of many human settlements in the Eastern Arctic (Andreasen 1997; Gotfredsen 2010; Gotfredsen, Appelt & Hastrup 2018; Grønnow 2016; Grønnow et al. 2011; Hastrup, Mosbech & Grønnow 2018; Henshaw 2003; Jeppesen et al. 2018; Kroon, Jakobsen & Pedersen 2010; Schledermann 1980; Sørensen & Gulløv 2012; Woollett, Henshaw & Wake 2000). Polynyas are recurring areas of thin ice or open water within the pack ice caused by strong prevailing winds or currents that allow concentrations of marine mammals and birds to feed over the winter and/or spring; these include the wide offshore cracks in the ice called ‘flaw’ polynyas that develop between the edge of shorefast ice and offshore pack ice (Henderson et al. 2021; Stirling 1997; Stirling & Cleator 1981; Stringer & Groves 1991). The major polynyas mentioned in regard to ancient human habitation are the North Water between Ellesmere Island and northwest Greenland, and the Northeast and Sirius Waters off northeast Greenland, although others may have been just as significant in providing human hunters with access to the abundant wildlife they needed to survive (Figure 2). Biologists have also noted the importance of both large and small polynyas to polar bear health and survival in the Canadian Arctic and Greenland (Heide-Jørgensen et al. 2016; Henderson et al. 2021; Stirling 1980; Stirling, Cleator & Smith 1981; Vibe 1950; Vibe 1967). Therefore, this analysis explores the historical distribution of ancient polar bear remains across the entire Arctic in relation to expansions of sea ice extent during known cold periods and as it overlaps areas where polynya conditions currently prevail (or may have in the past), as has been suggested for natural-death bowhead whale remains (Balaena mysticetus) in the Canadian Arctic during the middle to late Holocene (Dyke & England 2003; Dyke, Hooper & Savelle 1996).
|MAP #||SPECIMEN LOCATION †||AGE (A BP) ‡||TYPE OF DATE||RELATIVE AGE§||SPECIMEN TYPE||REFERENCE|
|1.||NE Point St. Paul Island Pribilofs USA||ca. 55||[shot 1895]||Late Holocene (LIA)||1 skull (old M)||Ray 1971|
|2.||St. Lawrence Is. (Kawarin grave) USA||ca. 40||ethnographic||Late Holocene (LIA)||89 skulls (ritual feature)||NPS 2013a|
|3.||Pottery House St. Matthew Is. USA||ca. 430–350||on deposit||Late Holocene (LIA)||9 assorted elements||Frink et al. 2001|
|4a.||Walakpa Site (late) USA||ca. 550–0||on deposit||Late Holocene (LIA)||13 assorted elements||Stanford 1976|
|4b.||Walakpa Site (middle) USA||ca. 1,050–550||on deposit||Late Holocene (MWP)||6 assorted elements||Stanford 1976|
|4c.||Walakpa Site (early) USA||ca. 1,450–1,150||on deposit||Late Holocene (DAC)||15 assorted elements||Stanford 1976|
|5.||St. Lawrence Is. (3 sites) USA||ca. 2,000–0||on deposit||Late Holocene (LIA-RWP)||present (not quantified)||Dumond 1998; Collins 1937; Murray 2008|
|6.||St. Lawrence Is. (Kukulik) USA||ca. 2,000–0||on deposit||Late Holocene (LIA-RWP)||287 skulls (in human burials)||NPS 2013b|
|7.||St. Lawrence Is. (Hillside site) USA||ca. 1,800–1,550||on deposit||Late Holocene (RWP)||present (not quantified)||Collins 1937; Dumond 1998; Arnold 2000|
|8.||Cape Espenberg Seward Peninsula USA||ca 2,500||on deposit||Late Holocene (NEO)||1 bone||Saleeby 1994|
|9a.||Qagnax Cave St. Paul Island USA †||4,830 ± 40||Beta-182978||Middle Holocene (NEO)||1 radius (distal) juvenile||Veltre et al. 2008|
|9b.||Qagnax Cave USA||4,410 ± 60||SPC-03–76||Middle Holocene (NEO)||1 phalanx adult||Veltre et al. 2008|
|9c.||Qagnax Cave USA||not dated||n/a||Middle Holocene (NEO)?||248 bones from 6 adult bears (2 M/4 F)||Veltre et al. 2008|
|10.||Bogoslov Cave St. Paul Island USA †||not dated||on deposit||Middle Holocene (NEO)?||2 adults 1 juvenile (15 bones/fragments total)||Ray 1971|
|11.||Margaret Bay (UNL-48) Unalaska Is. USA||ca. 4,700–4,100||on deposit||Middle Holocene (NEO)||102 (4 individuals)||Davis 2001; Murray 2008|
|12.||Washount (NjVi-2, H3) Herschel Is. CAN||ca. 400–260||on deposit||Late Holocene (LIA)||7 assorted elements||Friesen & Hunston 1994|
|13.||Agvik (OkRn1) Banks Is. CAN||ca. 500–300||on deposit||Late Holocene (LIA)||28 assorted elements||Kotar 2016|
|14.||Nelson River site Banks Is. CAN||ca. 650–50||on deposit||Late Holocene (LIA)||70 individuals||Arnold 1986; Moody & Hodgetts 2013|
|15.||Co-Op (OdPp-2, H1, H5) Victoria Is. CAN||ca. 500–50||on deposit||Late Holocene (LIA)||193 assorted elements||Lamy & Spitery 1991; Moody & Hodgetts 2013|
|16.||Lady Franklin Pt. (NdPd-2) Victoria Is. CAN||ca. 650–50||on deposit||Late Holocene (LIA)||4 assorted elements||Taylor 1972; Desjardins 2018|
|17.||Pingiqqalik (NgHd-1) Foxe Basin CAN||ca. 600–400||on deposit||Late Holocene (LIA)||55 assorted elements||Desjardins 2018|
|18.||Naujan (MdHs-1) Foxe Basin CAN||ca. 650–50||on deposit||Late Holocene (LIA)||1 bone||Mathiassen 1927; Desjardins 2018|
|19.||Sadlermiut (KkHh-1) Southampton Is. CAN||ca. 650–50||on deposit||Late Holocene (LIA)||38 assorted elements||Collins 1956; Collins 1981; Desjardins 2018|
|20.||Qijurittuq (IbGk-3, H1) Hudson Bay CAN||ca. 200||on deposit||Late Holocene (LIA)||2 assorted elements||Desrosiers et al. 2010|
|21.||Staffe Is. Labrador CAN||ca. 650–50||on deposit||Late Holocene (LIA)||present (not quantified)||Kaplan & Woollett 2016|
|22.||Nachvak Fjord group (IgCx-3; IgCv-7) Labrador CAN||ca. 650–50||on deposit||Late Holocene (LIA)||17 assorted elements||Swinarton 2008; Desjardins 2018|
|23.||Oakes Bay (HeCg-8) Labrador CAN||ca 270–170||on deposit||Late Holocene (LIA)||5 assorted elements||Woollett 2010|
|24.||Iglosiatik Is. Labrador CAN||ca. 650–50||on deposit||Late Holocene (LIA)||present (not quantified)||Kaplan & Woollett 2016|
|25.||JfEl-10 Quebec (Hudson Strait) CAN||ca. 650–50||on deposit||Late Holocene (LIA)||31 assorted elements||Lofthouse 2003; Desjardins 2018|
|26.||Talaguak Baffin Is. on Hudson Strait CAN||ca. 650–50||on deposit||Late Holocene (LIA)||13 assorted elements||Sabo 1981; Desjardins 2018|
|27.||Outer Frobisher Bay sites (KfDe-5; KfDf-2; KeDe-7) Baffin Is.CAN||ca. 650–50||on deposit||Late Holocene (LIA)||17 assorted elements||Henshaw 1995; Desjardins 2018|
|28.||Cumberland Sound (LlDj-1) Baffin Is. CAN||ca. 600–100||on deposit||Late Holocene (LIA)||3 assorted elements||Schledermann 1975; Dejardins 2018|
|29.||Hazard Inlet group (PaJs-3; PaJs-4; PaJs-13) Somerset Is. CAN||ca. 650–50||on deposit||Late Holocene (LIA)||24 assorted elements||Whitridge 1992; Dejardins 2018|
|30.||Learmonth (PeJr-1) Somerset Is. CAN||ca. 650–50||on deposit||Late Holocene (LIA)||146 assorted elements||Taylor & McGhee 1979; Rick 1980; Dejardins 2018|
|31.||Porden Pt. group (RbJr-1; RbJr-4; RbJr-5) Devon Is. CAN||ca. 650–50||on deposit||Late Holocene (LIA)||132 assorted elements||Park 1989; Dejardins 2018|
|32.||Porden Pt. (RbJq-6) Devon Is. CAN||ca. 700–600||on deposit||Late Holocene (LIA/MWP)||3 assorted elements||Howse 2019|
|33a.||Peale Pt. (KkDo-1) Baffin Is. CAN||ca. 650–100||on deposit||Late Holocene (LIA)||16 assorted elements||Stenton 1987|
|33b.||Peale Pt. (KkDo-1) Baffin Is. CAN||ca. 850–750||on deposit||Late Holocene (MWP)||3 assorted elements||Stenton 1987|
|34.||Sanirajak (NeHd-1) Foxe Basin CAN||ca. 750–450||on deposit||Late Holocene (LIA/MWP)||2 assorted elements||Desjardins 2013|
|35.||Kuukpak (NiTs-1, H1) Mackenzie R. CAN||ca. 750–350||on deposit||Late Holocene (LIA/MWP)||1 bone||Betts & Friesen 2006|
|36a.||Amundsen Gulf (Tiktalik NkRi-3, H5) CAN||ca. 750–650||on deposit||Late Holocene (MWP)||28 assorted elements||Moody & Hodgetts 2013|
|36b.||Amundsen Gulf (Pearce Point, Vaughn, Jackson sites)||ca. 650–50||on deposit||Late Holocene (LIA)||at least 4 elements||Morrison 2000; Taylor 1972; Moody & Hodgetts 2013|
|37.||Bell site (NiNg-2) Victoria Is. CAN||ca. 850–650||on deposit||Late Holocene (MWP)||4 assorted elements||Howse 2019|
|38.||Port Refuge (Snowdrift) Devon Is. CAN||ca. 1,000||on deposit||Late Holocene (MWP)||present (not quantified)||McGhee 1979; McGhee 1981|
|39.||Hornby Head (RbJq-1, H2, H3) Devon Is. CAN||ca. 1,100–650||on deposit||Late Holocene (MWP)||17 assorted elements||Howse 2019|
|40.||Brooman Point Bathurst Is. CAN||ca. 900||on deposit||Late Holocene (MWP)||present (not quantified)||McGhee 1984; Murray 2008|
|41a.||Skraeling Is. (SfFk-4, H2–12, 17–23) Ellesmere CAN||ca. 850–650||on deposit||Late Holocene (MWP)||235 assorted elements||McCullough 1989|
|41b.||Eskimobyen (SgFm-4, H25, H26) Ellesmere CAN||ca. 850–650||on deposit||Late Holocene (MWP)||53 assorted elements||McCullough 1989|
|41c.||Sverdrup Skraeling Is. (SfFk-5, H6) Ellesmere CAN||ca. 850–650||on deposit||Late Holocene (MWP)||13 assorted elements||McCullough 1989|
|42.||Skraeling Is. (SfFk-4, H 14–16) NE Ellesmere CAN||ca. 850–650||on deposit||Late Holocene (MWP)||66 assorted elements||Howse 2019; McCullough 1989|
|43.||Cape Garry (PcJq-5) Somerset Is. CAN||ca. 950–750||on deposit||Late Holocene (MWP)||21 assorted elements||Rick 1980; Dejardins 2018|
|44a.||Co-Op (OdPp-2, H1, H5) Victoria Is. CAN||1,350 ± 40||Gif-8434||Late Holocene (DAC)||1 bone||Harington 2003|
|44b.||Co-Op (OdPp-2, H1, H5) Victoria Is. CAN||1,310 ± 40||Gif-8178||Late Holocene (DAC)||1 bone||Harington 2003|
|45a.||Co-Op (OdPp-2, H2) Victoria Is. CAN||1,560 ± 65||Gif-7512||Late Holocene (RWP/DAC)||1 bone||Harington 2003|
|45b.||Lady Franklin Pt. (NdPd-2) Victoria Is.||1,510 ± 30||CAMS-66368||Late Holocene (RWP/DAC)||1 humerus||Savelle et al. 2012; Ingolfsson & Wiig 2009|
|46.||Cape Richard Collinson CAN||2,135 ± 120||Beta-18129||Late Holocene (RWP)||canine tooth||Harington 2003|
|47.||Seahorse Gully (IeKn 6) CAN||ca. 2,600–2,400||on deposit||Late Holocene (NEO)||present (not quantified)||Nash 1976|
|48.||Port Refuge (upper beach) Devon Is. CAN||ca. 4,000||on deposit||Late Holocene (NEO)||2 assorted elements||McGhee 1979; McGhee 1981|
|49.||Port Refuge (Gull Cliff) Devon Is. CAN||ca. 4,000–3,000||on deposit||Late Holocene (NEO)||3 assorted elements||McGhee 1979; McGhee 1981|
|50.||Port Refuge (Lower Beach) Devon Is. CAN||ca. 2,500||on deposit||Late Holocene (NEO)||2 assorted elements||McGhee 1979; McGhee 1981|
|51a.||Gulf of Boothia central CAN||3,265 ± 15||UCI-42204||Late Holocene (NEO)||1 bone||Dyke et al. 2011|
|51b.||Gulf of Boothia central CAN||3,515 ± 15||UCI-42211||Late Holocene (NEO)||1 bone||Dyke et al. 2011|
|51c.||Gulf of Boothia central CAN||3,290 ± 15||UCI-42210||Late Holocene (NEO)||1 bone||Dyke et al. 2011|
|51d.||Gulf of Boothia central CAN||3,765 ± 15||UCI-2207||Late Holocene (NEO)||1 bone||Dyke et al. 2011|
|52.||Baillie Island CAN †||not dated||on deposit||Pleistocene||1 bone||Harington 2003; Vincent 1989|
|53.||Scoresby Sound (House of Beads) GRE||ca. 150–50||on deposit||Late Holocene (LIA)||2 assorted elements||Sandell & Sandell 1991; Sørensen & Gulløv 2012|
|54.||Scoresby Sound (Skærgårdshalvøen 1) GRE||ca. 150–50||on deposit||Late Holocene (LIA)||a few elements||Degerbøl 1936|
|55.||Nugarsuk GRE||ca. 300–100||on deposit||Late Holocene (LIA)||5 assorted elements||Møhl 1979|
|56.||Walrus Is. (caches/shelters) GRE||ca. 550–100||on deposit||Late Holocene (LIA)||16 assorted elements||Gotfredson 2010; Grønnow et al. 2011|
|57.||Clavering Is. (sites 69, 78, 96, 105) GRE||ca. 550–100||on deposits||Late Holocene (LIA)||25 assorted elements||Gotfredson 2010|
|58.||Fladstrand (site 41) GRE||ca. 550–100||on deposit||Late Holocene (LIA)||91 assorted elements||Gotfredson 2010|
|59.||Dødemandsbugten (sites 45–47) GRE||ca. 550–100||on deposit||Late Holocene (LIA)||66 assorted elements||Sørensen et al. 2009; Gotfredson 2010|
|60.||Sephus Müller Næs (NEWland) GRE||460 ± 60||AAR-1776||Late Holocene (LIA)||1 bone||Andreasen 1997|
|61.||Qeqertaaraq (H1 + midden) GRE||ca. 850–750||on deposit||Late Holocene (MWP)||19 assorted elements||Howse 2019; Dejardins 2018|
|62.||Washington Land GRE||960 ± 60||AAR-5775||Late Holocene (MWP)||1 bone||Bennike 2002|
|63.||Washington Land GRE||1,415 ± 60||AAR-5774||Late Holocene (DAC)||1 bone||Bennike 2002|
|64.||Kolnæs Peary Land GRE||1,440 ± 45||K-352||Late Holocene (DAC)||R. mandible||Bennike 1991; Harington 2003|
|65.||Vandfeldsnaes Brønlund Fjord GRE||1,520 ± 110||AAR-1357||Late Holocene (RWP)||1 ulna||Bennike 1997|
|66.||Saqqaq Disko Bay GRE||ca. 2,900||on deposit||Late Holocene (NEO)||present (not quantified)||Gotfredsen 1992; Bennike 1997|
|67.||Solbakken (Hall Land) GRE||ca. 4,000–3,500||on deposit||Late Holocene (NEO)||9 assorted elements (mostly one individual)||Darwent 2003; Murray 2008|
|68.||Adam C. Knuth (Peary Land) GRE||ca. 4,000–3,500||on deposit||Late Holocene (NEO)||3 assorted elements||Darwent 2003; Murray 2008|
|69.||Pearylandville (Peary Land) GRE||ca. 4,000–3,500||on deposit||Late Holocene (NEO)||2 assorted elements||Darwent 2003; Murray 2008|
|70a.||Sønderland GRE||3,320 ± 85||K-5928||Late Holocene (NEO)||1 bone||Rasmussen 1996|
|70b.||Disko Bay GRE||3,470 ± 85||K-5930||Late Holocene (NEO)||1 bone||Rasmussen 1996|
|71.||Norde Eskimonœsset NEWland GRE||4,076± 90||AAR-1773||Late Holocene (NEO)||1 bone||Andreasen 1997|
|72.||Nuulliit (Thule) GRE||5,060 ± 95 uncal||K-2560||Middle Holocene (NEO)||1 bone||Knuth 1978; Bennike 1997; Grønnow & Jensen 2003|
|73.||Cape Schmidt RUS||ca. 100||ethnographic||Late Holocene (LIA)||+50 skulls (2 ritual features)||Kochneva 2007; Vdovin 1977|
|74.||Yamal Peninsula RUS||ca.250–50||ethnographic||Late Holocene (LIA)||‘many skulls’ (ritual feature)||Kochneva 2007; Kishchinskiy 1976|
|75.||Vaygach Island RUS||ca. 250–50||ethnographic||Late Holocene (LIA)||‘many’ skulls (ritual feature)||Kochneva 2007; Nordenscheldt 1881|
|76a.||Tiutei-Sale 1 (late) RUS||ca. 850–650||on deposit||Late Holocene (MWP)||89 assorted elements (5 individuals)||Fedorova et al. 1998; Nomokonova et al. 2018|
|76b.||Tiutei-Sale 1 (early) RUS||ca.1,350–1,150||on deposit||Late Holocene (DAC)||42 assorted elements (6 individuals)||Fedorova et al. 1998; Nomokonova et al. 2018|
|76c.||Tiutei-Sale 1 (early/late) RUS||ca. 1,350–650||on deposit||Late Holocene DAC/MWP)||164 assorted elements (10 individuals)||Fedorova et al. 1998; Nomokonova et al. 2018|
|77.||Dezhnevo Bering St. RUS||ca. 1,500–900||on deposit||Late Holocene (DAC)||33 assorted elements||Gusev et al. 1999; Savinetsky et al. 2004|
|78.||Cape Schmidt RUS||ca. 1,250–1,150||on deposit||Late Holocene (DAC)||skulls from human burials||Dikov 1988|
|79.||Cape Schmidt RUS||ca. 1,950–1,350||on deposit||Late Holocene (RWP)||‘many’ skulls (ritual feature)||Dikov 1988|
|80.||Cape Baranov Kolyma R. mouth RUS||ca. 1,855–1,525||on deposit||Late Holocene (RWP)||16 assorted elements||Bland 2008; Vereshchagin 1969|
|81.||Mainland south of Laptev Strait RUS||not dated||on deposit||Late Holocene?||present (not quantified)||Vereshchagin 1969|
|82.||Tikai (Laptev Sea) RUS||not dated||on deposit||Late Holocene?||present (not quantified)||Vereshchagin 1969|
|83.||Vaygach Island RUS †||1,971 ± 25||OxA-23631||Late Holocene (RWP)||R. ulna||Boeskorov et al. 2018|
|84.||Ekven Bering St. RUS||<2,700 BP uncal||on deposit||Late Holocene (NEO)||10 assorted elements||Savinetsky et al. 2004|
|85.||Devil’s Gorge Wrangel Is. RUS||ca. 3,620–2,950||on deposit||Late Holocene (NEO)||1 skull fragment; 1 claw||Dikov 1988; Tein 1977; Tein 1978|
|86.||Zhokhov Island RUS||ca. 8,250–7,800||on deposit||Middle Holocene (HCO)||5,915 assorted elements (130 individuals)||Pitulko et al. 2015|
|87.||Mordy-Yahk River mouth RUS †||not dated||on deposit||Pleistocene?||1 R. ulna (M)||Vereshchagin 1969; Harington 2008|
|88.||Pechora River mouth RUS †||not dated||on deposit||Pleistocene?||1 molar tooth||Harington 2008|
|89.||Iceland ICE †||ca. 13000||on deposit||Late Pleistocene (YD)||present (not quantified)||Áskelsson 1938; Petersen 2010|
|90.||Asdal DEN †||12,900–12,400||K-3741||Late Pleistocene (YD)||1 L. mandible (M)||Aaris-Sørensen 2009; Berglund et al. 1992|
|91.||Kuröd Bohuslän SWE †||10,170 ± 125 uncal||Lu-1075||Late Pleistocene (YD)||1 dist. femur + 4 other elements||Kurtén 1988; Berglund et al. 1992|
|92.||Nedre Kuröd Bohuslän SWE †||10,360 ± 130 uncal||Lu-1074||Late Pleistocene (YD)||1 rib fragment + 2 other elements||Kurtén 1988; Berglund et al. 1992|
|93.||Hisingen SWE †||not dated||on deposit||Late Pleistocene (YD)?||1 L. maxilla (M)||Kurtén 1988; Berglund et al. 1992|
|94.||Kärraberg Vekkinge parish SWE †||not dated||on deposit||Late Pleistocene (YD)?||1 skull (F?)||Kurtén 1988; Berglund et al. 1992|
|95.||Östra Karup Bastad SWE †||12,230 ± 130 uncal||Lu-1076||Late Pleistocene||1 R. ulna (F)||Berglund et al. 1992; Aaris-Sørensen 2009|
|96.||Kullaberg Scania SWE †||12,320 ± 125 uncal||Lu-602||Late Pleistocene||1 R. femur||Berglund et al. 1992; Aaris-Sørensen 2009|
|97.||Svenskøya Svalbard NO †||7,760 ± 50||T-4167||Middle Holocene (HCO)||1 bone||Harington 2008; Ingolfsson & Wiig 2009|
|98.||Svalbard NO †||ca. 8,200||on deposit||Middle Holocene (HCO)||>1 bone||Harington 2008|
|99.||Finnøy NOR †||10,925 ± 110 uncal||T-4724||Late Pleistocene (YD)||1 almost complete skeleton (M)||Blystad et al. 1983; Berglund et al. 1992|
|100.||Nordcemgrotta Kjæpsvik NOR †||ca. 22,000 uncal||direct date||Late Pleistocene 1||ulna + others||Lauritzen et al. 1996; Hufthammer 2001|
|101.||Hamnsundhelleren NOR †||36,000–28,000 uncal||direct date||Late Weichselian (MIS 3) 2||>1 bones||Valen et al. 1996; Hufthammer 2001|
|102.||Nordcemgrotta Kjæpsvik NOR †||ca. 115,000||on deposit||Early Weichselian||1 rib (mtDNA) + 2 other elements||Lauritzen et al. 1996; Davison et al. 2011|
|103.||Poolepynten Svalbard NOR †||ca. 130,000–110,000||LuS-6155||Eemian Interglacial/MIS 5e||1 L. mandible (M)(mtDNA)||Ingolfsson & Wiig 2009; Lindqvist et al. 2010|
|104.||Kew Bridge, Thames River UK †3||ca. 70,000||on deposit||Early Weichselian||1 R. ulna (M)||Kurtén 1988; Harington 2008|
This historical compilation presents, with some caveats, the entire record of ancient polar bear remains from fossil, archaeological, and ethnographic contexts prior to AD 1910 as recorded in the English scientific literature, presented by country in approximate chronological order (Table 1). Some specimens may have been missed because reports were never published, were reported in an inaccessible format (i.e. so-called ‘grey literature’) or published in a foreign language. Two well-known Russian-language archaeological reports were consulted but there was no attempt to make a comprehensive search of the Russian literature or to access records published in Norwegian, Swedish, Finnish, or Icelandic. However, in many cases, specimens initially reported in a language other than English or in unpublished reports have been cited by other authors in English papers, in which case, I refer to both sources.
The ‘fossil’ remains reported here are in most cases not actually mineralized and are technically ‘subfossils’, as is true for the archaeological remains. However, for the purpose of this report, all natural-death remains are referred to as fossils. The table includes information on location, chronological date or dates (if available), approximate geological time period, type of specimen, and abundance information (if available), and sources (references). All geological and climatological time periods used in this paper are defined in Table 1 and the approximate geographical location of the specimen finds are shown in Figure 1.
Some single polar bear finds have been dated directly and where this has been done, the date is reported as given and the lab number for the date provided. However, this level of precision is rare for most archaeological remains except for some specimens from Canada and Greenland (e.g., #51, 71). Specimens from archaeological sites are in most cases given as approximate dates for associated deposits using a range of dating methods (including artifact styles, depth of deposit, and 14C dates on other material, including charcoal) and therefore, lab numbers for dates are not provided. Because they are a marine mammal, direct dates on polar bear bone have been corrected for the carbon reservoir effect, the phenomenon that makes 14C dates on marine material appear older than they actually are by up to about 400 years (depending on the region). Unfortunately for the use of charcoal for dating, the prevalent use of long-dead driftwood by ancient human hunters in the Arctic has a similar effect on accuracy. In addition, charcoal and bone from terrestrial species from Arctic sites may be contaminated in situ by oils from marine mammals. With these caveats in mind, modern archaeologists are usually careful in their selection of datable material and choose terrestrial mammal bone such as musk ox or caribou, or fast-growing wood like willow where ever possible (e.g., Friesen, Finkelstein & Medeiros 2020; McGhee 2000), may pre-treat terrestrial mammal bone to test for the presence of sea-mammal lipids (e.g., Desjardins 2018), and/or test terrestrial species together with a marine species to arrive at a local marine-reservoir correction factor (e.g., Dyke et al. 2018). The dating accuracy in the polar bear data presented here therefore varies considerably and makes all but broadly-defined chronological patterns untenable. However, it is considered better to know the true nature of the record than to impose arbitrary limits for inclusion that might discard important records that could, if re-examined, yield more useful information in the future.
In addition to the record of ancient polar bear remains, an Arctic map of the approximate location of known polynyas is provided (Figure 2) based on regional studies of this phenomenon (Barber et al. 2001; Grønnow et al. 2011; Jackson et al 2020; Kassens & Thiede 1994; Kern 2008; Morales Maqueda, Willmott & Biggs 2004; Pedersen et al. 2010; Smedsrud et al. 2006; Speer et al. 2017; Stirling & Cleator 1981; Stringer & Groves 1991). Some polynyas are not only important areas of biological productivity and air to breathe for seals, walrus, and whales but contribute extensively to sea ice formation in the Arctic. For example, severe continental weather in Siberia generates cold winds that blow across the shallow Laptev Sea from October to April, which create almost constant upwelling that generates a large flaw polynya about 1,800 km long and 10–15 km wide, called the Great Siberian polynya, which is largely responsible for the almost continuous production of Arctic sea ice every winter (Buckley et al. 1979; de Vernal et al. 2020; Tamura & Ohshima 2011; Wakefield 2020). For polar bears, polynyas offer critical ice-edge hunting opportunities that may otherwise exist only at the periphery of consolidated pack ice. Changes in size and productivity have been documented for a number of polynyas since the end of the LGM that may have influenced polynya availability and thus polar bear distribution during the Holocene: e.g., Northeast Water (Hjort 1997); Kara Sea polynyas (Hörner, Stein & Fahl 2018); North Water (Jackson et al. 2021); and Storfjorden (Rasmussen & Thomsen 2014). Some polynyas may not have existed at all before a certain time: for example, one analysis (Dyke & England 2003) suggested that the polynyas that currently form due to high water flow between the channels that separate Ellesmere and Devon Island in the Central Canadian Arctic (Hell Gate-Cardigan Strait and Penny Strait) probably did not exist before 4,000 BP due to postglacial isostatic uplift. In contrast, some polynyas may have existed in the past that are no longer present today due to sea level and sea ice changes, as I suggest may have existed in the North Atlantic during the LGM and its immediate aftermath.
Most ancient remains of polar bears come from archaeological sites and ethnographic locations within the modern range of the species that date within the Holocene. Extralimital polar bear specimens have been documented in the north Atlantic during the late Pleistocene and in the Bering Sea during the middle Holocene (Figure 1, Table 1). These extralimital records indicate that sea ice extended beyond the present maximum extent (currently reached in March every year) at two particular points in time: in the Bering Sea during the mid-Holocene Neoglacial cold period (Crockford 2008; Crockford & Frederick 2007; Caissie et al. 2010; Davis 2001) and in the North Atlantic during the Younger Dryas (YD) cold period. The YD was a rapid return to cold conditions that briefly interrupted the warming that began ca. 19,700 a BP and which eventually brought the LGM to an end ca. 11,700 a BP (Alley 2000; Bradley & England 2008; Cheng et al. 2020).
In the Bering Sea, there are both fossil and historic era records that date to the mid-to-late Holocene: an old bear shot on St. Paul Island in the Pribilof Islands in 1875 (#1) dates to the Little Ice Age (Ray 1971), and two assemblages on the same island found in vertical caves (Qagnax and Bogoslov), which functioned as lethal ‘death traps’ (#9 and 10), date to the early part of the Neoglacial. The 250 polar bear bones from Qagnax Cave constitute the largest fossil assemblage found in the Arctic and represent at least eight bears, two of which were dated directly (Veltre et al. 2008). The material from nearby Bogoslov Cave (n = 15, three individuals) has not been dated but presumably comes from a similar period (Ray 1971).
Iceland, southern Norway, southern Sweden, and Denmark have generated nine fossil polar bear remains (#89–99), seven of which date within the brief YD cold period and two (#93, 96) date to a slightly earlier time when the region was undergoing active deglaciation (Aaris-Sorensen 2009; Aaris-Sorensen & Petersen 1984; Áskelsson 1938; Berglund et al. 1992; Bylstad et al. 1983; Harington 2008; Ingolfsson & Wiig 2009; Petersen 2010). During both periods, the Skagerrak Strait between Norway and Denmark was essentially a dead-end fjord of the North Sea with ice cover in winter and spring which probably had an associated polynya due to cold winds blowing off the thick ice sheet that still covered Norway and Sweden (Berglund et al. 1992; Gyllencreutz 2005; Stroeven et al. 2016). Most of these extralimital fossil remains are isolated bones or a small cluster of bones that have been dated directly, although there is also one almost complete skeleton of an old male approximately 28 years old (#99) (Berglund et al. 1992; Næss 2018). The complete mandible from an adult male recovered in Denmark (#90) is shown in Figure 3.
The sheer number of natural death remains of polar bears recovered in Scandinavia that date within a narrow time frame is unique. It suggests strongly that the climatic conditions during the late LGM that created suitable habitat for polar bears so far south of their modern range were associated with unusual circumstances that have not existed elsewhere in time or space. Either death rates from starvation or bone survival rates—or both—were unusually high. It is possible that polar bears existed at high densities due to limited suitable habitat in the region, resulting in greater competition and higher overall death rates, and/or that abrupt sea level changes and rapid sediment accumulation during deglaciation preserved a greater number of bones than usual. I suggest the clustering of remains along that ancient shoreline indicate that polynya formation was likely a feature of the sea ice in the region at that time, similar to those that develop in Frobisher Bay and Cumberland Sound on Baffin Island today (Figure 2) (Gyllencreutz 2005), although no geophysical evidence of such a phenomenon has been reported.
Three additional Scandinavian specimens pre-date the end of the LGM and also lie outside the current range of the species on the Norwegian coast. The specimen from Nordcemgrotta (#102), on a small island on the northwest coast, has been dated to the beginning of the Early Weichselian glacial period (ca. 115k cal a BP) and has had mitochondrial DNA (mtDNA) extracted and reported (Davison et al. 2011; Hufthmammer 2001; Lauritzen et al. 1996). Specimen #101 was found in a coastal cave farther south and dates to the Late Weichselian (‘Ålesund Interstadial’, aka MIS 3 interstadial 3.1, ca. 36,000–28,000 cal a BP), an LGM ice retreat documented in this region (Hufthammer 2001; Lambeck et al. 2010; Valen et al. 1996). Another specimen found at the Nordcemgrotta site (#100) has a date of 22,000 14C a BP (Hufthammer 2001; Lauritzen et al. 1996) and is associated with the so-called ‘Hamnsund Interstadial’ which was another, but short-lived ice retreat dated to 22,000–19,000 14C a BP in western Norway (Winguth et al. 2005: 181). All three specimens are associated with ice sheet formation and expansion over Svalbard and Scandinavia during the last Glacial period. Ice sheet formation pushed Barents Sea polar bears and other Arctic marine mammals to the southern North Sea (Post 2005), except during short periods when suitable habitat existed along the Norwegian coast during temporary ice retreat.
A fourth pre-LGM polar bear specimen (#104, ca. 70k cal a BP) also lies outside the current range of the species but its taxonomic identity has been disputed. It was originally identified as polar bear several decades ago (Kurtén 1988), with a note it was large even for that species. However, while C.R Harington (2008: S25) argued that the identification of polar bear is plausible based on sea level changes and ice conditions in the North Sea during that time (e.g., Bennike et al. 2014; Post 2005), he also stated:
‘Andy Currant of the Natural History Museum – London (personal communication) believes that the Kew Bridge bear ulna represents a huge brown bear rather than a polar bear, based on faunas similar to that at Kew Bridge from many British sites containing dominant steppe bison (Bison priscus) and reindeer (Rangifer tarandus) with wolves (Canis lupus) and gigantic brown bears moderately represented’.
This opinion that the Kew Bridge specimen is not polar bear, also expressed in an interview with the BBC in 2007 (Amos 2007) and a note in a 2009 scientific paper (Ingolfsson & Wiig 2009), awaits the official verification of a published note by Currant that corrects the record.
A small third lower molar tooth (not included in Table 1), reported to resemble polar bear in size and shape, was recovered amongst remains of black bear (Ursus americanus) from an archaeological site in coastal New England called Crouch’s Cove, apparently of Late Holocene age (perhaps LIA), that was excavated in the mid-1800s (Packard 1886; Wyman 1868). The tentative nature of the original identification precluded its inclusion in this record, although if confirmed it would represent an extralimital record. Similarly, the report of a cluster of bones (right humerus, left femur, right fibula, some ribs, plus vertebrae 1 and 2) of undetermined chronological age from Lough Gur near Limerick, Ireland in 1858 identified as polar bear (Denny 1859) would also be an extralimital occurrence but do not appear in any other record and is therefore considered an identification error.
Only seven polar bear fossils have been found within the current range of the species and all were found in proximity to modern polynyas. Four are from the Barents Sea (#88, 97, 98, 103): three from Svalbard (#97, 98, 103), a short distance from the central Storfjorden Bay polynya, and one from the southern Barents Sea coast of Russia (#88) where small coastal flaw polynyas routinely form (not shown in Figure 2) (Harington 2008; Ingolfsson & Wiig 2009). One Pleistocene-aged specimen was found in the Kara Sea (#87) where coastal polynyas are also common (Harington 2008; Vereshchagin 1969). A right ulna from an adult bear dated to 1,971 ± 25 BP, recovered from Vaygach Island (#83) in the same area, is presumed to be from a natural deposit as it predates the known occupation of the region by Nenets people (Boeskorov et al. 2018). Another Pleistocene-aged specimen was recovered from the eastern Beaufort Sea (#52) at the edge of the modern Bathurst polynya (Harington 2003; Vincent 1989).
Aside from the Vaygach Island bone, only three of these specimens have been dated more precisely than ‘Pleistocene’. One is the oldest dated fossil (#103), a complete mandible with canine tooth from a male bear with a chronological age that falls within the warm Eemian Interglacial, ca. 130–110 ka BP. This specimen has also yielded a complete mtDNA sequence that has been critical for inferring polar bear evolutionary history (Ingolfsson & Wiig 2009; Lan et al. 2022; Lindqvist et al. 2010). The other two specimens from Svalbard (#97, 98) date to the Holocene, ca. 8,000 a BP (Harington 2008; Ingolfsson & Wiig 2009) and are the earliest reported polar bear remains from the Eastern Arctic after the end of the LGM and the melting of the Svalbard ice sheet, ca. 10,000–8,200 a BP (Rasmussen & Thomsen 2014).
In the Aleutian Islands, archaeological remains of polar bear (n = 102, 24 confidently identified as polar bear, plus an additional 78 presumed to be polar bear rather than brown bear as both species were confidently identified) were recovered from Margaret Bay on Unalaska Island near Dutch Harbour (#11) (Davis 2001). The dates of the deposits (based on charcoal) have a similar range to the Pribilof fossil specimens (#9, 10) mentioned above (ca. 4,700–4,100 a BP). The slightly younger but still Neoglacial-aged deposit at the Amaknak Bridge site (UNL-50), lies adjacent to Margaret Bay, and while it lacks polar bear remains it does have faunal indicators (especially foetal and newborn ringed and bearded seal remains) used as evidence that late spring sea ice extended much farther south than it does today (Crockford & Frederick 2007; Crockford & Frederick 2011). The historic era specimen shot on the Pribilofs indicates that sea ice expanded that far south during the LIA (as it has done occasionally in recent times), but as far as is known, not as far south as the eastern Aleutians as it did during the Neoglacial (Brown, van Dijken & Arrigo 2011; Crockford & Frederick 2007; Frey et al. 2015).
One unique archaeological assemblage stands out from all others with regards to polar bear remains: the faunal material from the Zhokhov Island site (record #86), at 76°N where the Laptev Sea meets the East Siberian Sea. The site was excavated in 1989–1990 (Pitulko 2003; Pitulko 1993; Pitulko & Kasparov 1996) and again in 2000–2005 (Pitulko et al. 2015). It is not only the oldest archaeological site in the Arctic with polar bear bones but also contains by far the most polar bear remains of any human occupation (n = 5,915). In contrast to most sites, where they represent at most 3.5% (usually less) of the total mammalian remains recovered (Table 2), polar bear bones at the Zhokhov Island site comprised 28.4% of the total and represent at least 130 individuals. Domestic dogs were also recovered and were assumed to have chewed many of the damaged polar bear bones (Pitulko & Kasparov 2017). The site was inhabited for at most 450 years between ca. 8,250 and 7,800 a BP (Pitulko et al. 2019), although most of the deposits date to a brief period ca. 8,000–7,900 a BP. It is known that the initial flooding of Beringia by rising sea levels at the end of the LGM began before 10,000 a BP, which made the Arctic accessible again to marine mammals that had taken refuge in the North Pacific during the LGM (Crockford, Frederick & Wigen 2002; Dyke, Hooper & Savelle 1996; Dyke et al. 1999; de Vernal et al. 2020; Guthrie 2004; Heaton & Grady 2003; Polyak et al. 2010). Therefore, the large assemblage of skeletal remains recovered from Zhokhov Island marks the first evidence known of the return of polar bears to the western Arctic after being driven out by extraordinarily thick ice cover during LGM.
|MAP REF. & SPECIMEN LOCATION||COUNT||NISP||PERCENTAGE|
|11. Margaret Bay (UNL-48) Unalaska Is. AK||102||12,548||<1|
|36. Tiktalik (NkRi-3, H5) CAN||28||6216||<1|
|14. Nelson River CAN||?||70†||3.5|
|15. Co-Op (OdPp-2, H1, H5) Victoria Is. CAN||193||22,200||<1|
|37. Bell site (NiNg-2) Victoria Is. CAN||4||5,791||<1|
|17. Pingiqqalik (NgHd-1) Foxe Basin CAN||55||10,753||<1|
|19. Sadlermiut (KkHh-1), CAN||38||2,818||1.3|
|29. Hazard Inlet group, Somerset Is. CAN||24||10,235||<1|
|43. Cape Garry (PcJq-5) Somerset Is. CAN||21||2,658||<1|
|30. Learmonth (PeJr-1) Somerset Is. CAN||146||4,892||3.0|
|39. Hornby Head (RbJq-1, H2, H3) CAN||17||1,820||<1|
|41a. Skraeling (SfFk-4, H2–12, 17–23) CAN||235||9625||2.4|
|41b. Eskimobyen (SgFm-4, H25–26) CAN||53||3185||1.7|
|41c. Sverdrup (SfFk-5, H6) CAN||13||391||3.3|
|42. Skraeling Is. (SfFk-4, H14–16) CAN||66||2,810||2.3|
|61. Qeqertaaraq, (H1 + midden) GRE||19||2,249||<1|
|56. Walrus Is. (caches/shelters) GRE||16||1,044||1.5|
|58. Fladstrand (site 41) GRE||91||4,642||2.0|
|59. Dødemandsbugten (sites 45–47) GRE||66||2,625||2.5|
|53. Scoresby Sound (House of Beads) GRE||2||522||<1|
|67. Solbakken, GRE||9||60||15.0|
|76c. Tiutei-Sale 1 Early-Late RUS (total sample)||295||3,423||8.6|
|76b. Tiutei-Sale 1 Early only (DAC) RUS||42||159||26.4|
|76a. Tiutei-Sale 1 Late only (MWP) RUS||89||1,931||4.6|
|86. Zhokhov Island RUS||5,915||20,855||28.4|
The Zhokhov site occupants were primarily reindeer hunters and apparently treated polar bears as a terrestrial resource, as there were few other marine mammals remains present (e.g., only six seal bones, no walrus, no whale). This is a pattern not seen elsewhere in the Arctic, regardless of time period. The bears appear to have been primarily females (some with newborn young) taken on land in winter or early spring with spears from their winter maternity dens although mixed sexes were perhaps taken in traps on land during the ice-free season (Pitulko et al. 2015). The range of total length of intact mandibles recovered (n = 37, sex/age unknown; mean 223.1 mm, range 206–268 mm) indicates at least a few adult males as well as females were taken, based on measurements of modern adult bears from Svalbard and East Greenland (female, n = 47: mean 217.8 ± 6.6, range 203.1–232.9 mm; male, n = 58: 243.2 ± 11.2, range 216.1–265.9 mm) (Bechshøft et al. 2008).
Approximately 8,300 years ago, the slightly elevated terrain of Zhokhov Island was part of a low coastal plain that extended ca. 100 km north of the present coastline. It remained above sea level after Beringia was inundated. Today few areas of the eastern Laptev Sea and the East Siberian Sea are deeper than 50 m (Pitulko et al. 2019). However, as sea levels continued to rise, the region was transformed ca 7,800 a BP into an archipelago—the New Siberian Islands—which put an end to the human occupation. The Great Siberian flaw polynya first developed after the end of the LGM at about 14–16 cal ka BP (Taldenkova et al. 2008). Today, it extends as far east as the New Siberian Islands (Kassens & Thiede 1994; Speer et al. 2017). Given that Siberian winters 8,000 a BP were cold (Kokorowski et al. 2008; Nazarova et al. 2013) but with reduced summer sea ice cover offshore compared to today (Taldenkova et al. 2008), it seems likely that polynya formation documented since the 20th century also occurred to some degree at the time of the site’s occupation (Andreev et al. 2009; Hörner et al. 2018; Kassens & Thiede 1994; Timokhov 1994). Since Zhokhov Islanders were not marine mammal hunters, the faunal remains from this site are unhelpful in determining whether the Pacific walrus, which currently over-winter in the Great Siberian polynya, were present at that time (Fay 1982; Lindqvist et al. 2009). However, the presence of polar bear is consistent with ecological conditions similar to today, including the reliable off-shore presence of breeding ringed seals in spring which make land-based denning by females possible ((Amstrup & Gardner 1994; Pitulko et al., 2015; Ramsay & Stirling 1988; Stirling 1997; Stirling 2002).
All other Holocene-aged archaeological sites are within the modern range of polar bears. Archaeological sites with more than ten polar bear elements are primarily near modern major open-water polynyas, including the one south of St. Lawrence Island, and in Peard Bay (off Utqiaġvik, Alaska – formerly known as Barrow), the Cape Bathurst polynya, the North Water, the Sirius Water, as well as those in the Kara, Laptev Sea and East Siberian Seas, Frobisher Bay, Bellot Strait, and Hell Gate/Cardigan Strait (between Ellesemere and Devon Islands) (Table 3). As Table 2 indicates, sample sizes for virtually all of these are so much smaller than Zhokhov Island that they are best compared to each other. Of these, the Tiutei-Sale 1 site on the Yamal Peninsula (#76), where polar bear bones comprised 42 of 159 bones (i.e., n = 42/159) or 26.4% of the early occupation during the Dark Ages Cold period (DAC) component, had the highest relative abundance after Zhokhov Island. However, for all periods combined bear remains at Tiutei-Sale 1 represent only 8.6% of the sample (and 21 individuals). In only one other site did polar bear remains comprise more than 5% of the sample: the Neoglacial-aged site of Solbakken in Greenland opposite the northeastern end of Ellesmere Island (#67), where polar bear remains made up 15.0% of the mammalian sample (n = 9/60). However, this metric is skewed because most of the polar bear remains appear to be from one individual (Darwent 2003) and the total sample size is small. Sites with the next highest abundance of polar bear remains were in the Canadian Arctic Archipelago: at Sverdrup (#41) on Ellesmere Island at 3.3% (n = 13/391) (adjacent to the North Water) and Learmonth (#30) on Somerset Island, at 3.0% (n = 146/4,892) (near the Bellot Strait polynya). At the Nelson River site (#14) adjacent to the Cape Bathurst polynya, the material was reported only as minimum number of individuals (MNI) rather than bone count but a minimum of 70 individuals accounted for 3.5% of the mammalian MNI remains reported (Moody & Hodgetts 2013).
|SITE #||POLYNYA CODE||POLYNYA NAME||POLAR BEAR COUNT|
|97, 98, 103||A||Storfjorden Bay||1 each|
|74, 75?, 76||C||Kara Sea group||>10 each|
|83, 87||C||Kara Sea group||1 each|
|86||D||Great Siberian flaw||>10 each|
|81, 82||D||Great Siberian flaw||P (at least 1 each)|
|3||G||St. Matthew Island||9|
|2, 6||H||St. Lawrence Island||>10 each|
|5, 7||H||St. Lawrence Island||P (at least 1 each)|
|4a, 4c||L||Peard Bay||>10|
|13, 14, 36a||M||Cape Bathurst||>10 each|
|36b, 52||M||Cape Bathurst||1–4 each|
|29, 30, 43||N||Bellot Strait||>10 each|
|40||O||Penny Strait/Queens Channel||1|
|31, 39||P||Hell Gate/Cardigan Strait||>10 each|
|32, 38, 49, 50||P||Hell Gate/Cardigan Strait||1–3 each|
|17||Q||Fury and Hecla Strait||>10|
|18, 34||Q||Fury and Hecla Strait||1–2 each|
|20||S||Hudson Bay flaw||2|
|27, 33a, 33b||T||Frobisher Bay||>10 each|
|41a-41c, 42, 61||V||North Water||>10 each|
|62, 63, 72||V||North Water||1 each|
|60, 71||W||NE Water||1each|
|56, 57, 58, 59||X||Sirius Water||>10 each|
|53, 54||Y||Scoresby Sound Water||2 each|
|64, 65, 68, 69||Z||Wandel Water (proposed)||<4 each|
Four sites with fewer than four polar bear bones each were found in northeast Greenland at Peary Land that date to several periods (#64, 65, 68, 69) (Bennike 1991; Bennike 1997; Darwent 2003; Grønnow & Jensen 2003). Sites here are closest to the geographic North Pole (ca. 82° N) of any archaeological sites with faunal remains (from both terrestrial and marine species). The large polynya that developed in that region in 2018 and again in 2020 (Ludwig et al. 2018; Moore et al. 2018; Schweiger et al. 2021) (called here the ‘Wandel Water’, Figure 2) may not be an entirely new phenomenon but a recurrent feature that has formed historically to some degree under particular climatic conditions. Alternatively, it may also be that this area is close enough to the NE Water for both people and polar bears to access seals. Polar bears are rare in this area because the thick offshore ice precludes the survival of the seals they need to survive (Bennike 1991), but they do occur. In 1992, a female bear with a satellite collar travelled from the Beaufort Sea, across the Arctic Ocean to an area off the northeast coast of Greenland, then moved west across the Peary Land coast of northern Greenland, and eventually made her way into Kane Basin at the North Water (Figure 2) (Durner & Amstrup 1995). Such transient occurrences may be more common than has been documented. In addition, in 2018 and 2019, three polar bear maternity dens made in snow banks around icebergs grounded in land-fast ice were observed in the Peary Land area and females with cubs were also sighted (Laidre & Stirling 2020). These records indicate a small resident population of polar bears and therefore, a reliable source of breeding ringed seals nearby.
Despite polar bears being abundant in Hudson Bay today because of the flaw polynya that develops every winter between the shorefast ice and the central pack ice (Henderson et al. 2021; Stirling 1997), only two archaeological sites in the region have bear remains (Desrosiers et al. 2010; Nash 1976) and no polar bear fossil remains at all have been recovered (Harington 2003). The area was covered by remnants of the Laurentide Ice Sheet until about 8,000 a BP (Condron and Winsor 2012) and Hudson Bay as we know it today did not exist until about 7,800 a BP. At that time sea level was about 165 m above present sea level at Churchill (Dredge 1992). Due to changes in the shoreline and currents, it may not have been suitable ringed seal and polar bear habitat until about 6,500 a BP (Bilodeau et al. 1990; Harington 1988; Harington 2008). By about 2,000 a BP, sea level was still about 25m above present levels and the shoreline several kilometers inland from its present position. This means any coastal sites occupied by ancient people (and any terrestrial maternity dens of polar bears) would be of recent age and well inland from the present coastline unless they were located on elevated terrain (Murray 2008; Nash 1976).
The relative dearth of archaeological sites reporting polar bear remains from across the huge expanse of the Russian Arctic coast is almost certainly a reflection of my inability to read or access the Russian literature and because some regions may be better surveyed than others. The Yamal Peninsula and the coast of Chukotka, in particular, appear to have been relatively well surveyed and reported by archaeologists and ethnologists. Work by ethnologists in the 1800s, for example, indicate the Nenets people considered polar bears to have strong spiritual qualities and polar bear ‘monuments’ discovered during the 1800s and early 1900s (#73–76) are evidence of this belief system (Kishchinskiy 1976; Kochneva 2007; Vdoving 1977). Such features are composed of large numbers of polar bear skulls that appear to have accumulated over centuries and span the Russian Arctic from Chukotka to the Barents Sea. Similar finds, but with no other details provided, have been reported from Wrangel Island and adjacent to the villages of Vankarem, Inchoun, Enormino, Akkani and others in Chukotka (Kochneva 2007). Archaeological reports of polar bear skulls associated with human burials (#78) and a prehistoric ritual feature (#79) involving multiple polar bear skulls associated with a shaman, come from much older time periods at Cape Schmidt (opposite Wrangel Island on the Chukotka coast), support the suggestion that this spiritual role for polar bears was long-standing (Dikov 1988).
This belief seems to have travelled with ancient peoples of Siberia east to St. Lawrence Island in the Bering Sea. There is little detail available on the polar bear remains from sites on this prominent island (e.g., #5, 7), which were excavated in the early 20th century when faunal remains were of little interest to archaeologists (e.g., Rainey 1941). In the reports that are available (e.g., Collins 1937), species are listed only as ‘present’ and could be almost 2,000 years old or only a few centuries. However, two caches of polar bear skulls excavated by Dr. Otto Geist in the 1930s eventually made their way to the American Museum of Natural History along with his field notes and were later catalogued for repatriation to their ancestral communities (NPS 2013a; NPS 2013b). These consisted of 89 skulls collected at Cape Chibulak (near Gambell) from the grave of a hunter named Kowarin who died in 1910 (#2) and another 287 skulls from prehistoric human burials near Kukilik (near Savoonga), some of which may be almost 2,000 years old (#6). These finds extend the Russian pattern of a strong and long-standing spiritual role for polar bears into the Bering Sea at St. Lawrence Island.
There is only one archaeological site with polar bear remains recorded on St. Matthew Island in the southern Bering Sea (#3) (Table 3) (Frink et al. 2001), but this is the only prehistoric site ever excavated (Griffin 2008). However, it is known from historic records that as late as 1875, hundreds of polar bears used the island as a summer refuge and winter denning area but were exterminated by the 1890s by indiscriminate hunting (Elliott 1875; Elliott & Coues 1875; Klein & Sowls 2011). Bears have not recolonized the island since, but as illustrated (Figure 2), St. Matthew Island develops a prominent polynya on its south coast in spring similar to St. Lawrence Island, which almost certainly made it as suitable a denning area as Wrangel Island in the Chukchi Sea is today (Garner et al. 1994; Voorhees et al. 2014).
Only the Tiutei-Sale 1 site on the Yamal Peninsula provided data adequate to addressing whether relative polar bear abundance might have changed between distinct short-term climatic changes at the same location over time (Table 2) (e.g., Briffa et al. 2013; Connolly & Connolly 2014). At Tiutei-Sale 1, the Medieval Warm Period (MWP) deposits yielded relatively fewer polar bear bones (4.6% of the sample) than the preceding DAC (26.4%). Although the DAC results may be skewed by the much smaller sample size compared to the MWP sample (159 vs. 1,931), it does suggest the possibility that polar bears may have been hunted more frequently during the DAC period at this location but cannot tell us unequivocally that this was because the animals were more abundant.
As far as it has been possible to determine, there are no fossil polar bear remains reported from Ireland, although it has been suggested polar bears evolved nearby and abundant brown bear remains have been recovered (Edwards et al. 2011; Edwards et al. 2014). In addition, although there are fossil remains reported, no archaeological remains of polar bears have been found anywhere in the UK or Scandinavia. No archaeological or fossil remains have been recovered from the Barents Sea coasts of northern Norway or Finland (Rankama 2003). Similarly, there were no ancient polar bear remains of any kind found in the Sea of Okhotsk or the Gulf of Alaska in the western Arctic although the presence of ringed seal bones dated to the LGM on Prince of Wales Island, Southeast Alaska suggest there was almost certainly suitable ice-edge habitat for polar bears in the region (Heaton & Grady 2003). Furthermore, although bowhead whales apparently returned to the western Canadian Arctic via Bering Strait soon after it was physically possible to do so (Atkinson 2009; Dyke & England 2003; Fisher et al. 2006), I was informed by geologist Art Dyke (pers. comm., 2007) that no natural-death assemblages of polar bears were found during the shoreline surveys of both eastern and western portions of the Canadian Arctic Archipelago that recovered early to mid-Holocene bowhead and walrus fossil remains (Dyke, Hooper & Savelle 1996; Dyke et al. 1999; Dyke et al. 2011; Dyke & Savelle 2001).
Within the past 130 ka, sea ice conditions have at times been very different than they are today and this has affected where polar bears have been able to live. The thick perennial ice that developed during the LGM pushed polar bears south and out of the Arctic entirely. They returned when warmer conditions prevailed during the HCO. The Eemian Interglacial and the HCO, although both were warmer than today with less summer ice, apparently provided adequate habitat for polar bears to survive around Svalbard and in the East Siberian Sea. During the Neoglacial cold period of the Middle and Late Holocene, sea ice extended farther south into the Bering Sea than it does today, which allowed polar bears to temporarily reach the Pribilof Islands and the Eastern Aleutians.
The oldest dated polar bear fossil (ca. 130–110k a BP) was found within the modern range of the species, which is also true for virtually all Holocene-age archaeological sites with polar bear remains (one exception). Extralimital polar bear fossil specimens have been documented in the north Atlantic from the late Pleistocene (13 records) and in the southern Bering Sea during the mid-Holocene (three records). Prevailing sea level, ice sheet, and sea ice conditions surrounding the ancient Skagerrak fjord between Norway and Denmark during the YD support a suggestion that the region probably had an associated polynya, although this has not been confirmed by geophysical evidence.
The enormous assemblage of polar bear bones found at the Zhokhov Island archaeological site in the East Siberian Sea (ca. 8.2–7.8k a BP) and two fossil specimens recovered from Svalbard, Norway (also ca. 8.2–7.8k a BP) are so far the earliest evidence of the return of polar bears to the Arctic after the end of the LGM and all date to the same period of the HCO. The Zhokhov assemblage is the only archaeological site dating to the HCO and has by far the highest proportion of polar bear remains, as well as the greatest number of remains, recovered from any time period across the Arctic. Prevailing climatic conditions in the East Siberian Sea region during the HCO indicate that a polynya in some form probably existed about 8,000 years ago as it does today.
Except for the Zhokhov site and one Neoglacial-aged site in the southern Bering Sea, archaeological sites older than 2,000 years have relatively few polar bear remains. Only one archaeological site with deposits that span a complete climatic shift within the last 2,000 years (Tiutei-Sale 1 site on the Yamal Peninsula) has data that are indicative of a shift from hunting more bears during a cold period (DAC) to fewer during a subsequent warm period (MWP), but no broad conclusions can be drawn from this example. Except for two Neoglacial-aged natural-trap sites in the Bering Sea and one almost-complete late Pleistocene skeleton from southern Norway, fossil remains are predominantly single element finds.
Archaeological sites with more than ten polar bear elements are primarily near modern open-water polynyas, as are most of the isolated fossil remains. Polar bear remains from sites near the Hell Gate-Cardigan Strait and Penny Strait polynyas all date well after the postglacial uplift 4,000 years ago that created the polynyas. On St. Lawrence Island in the Bering Sea, evidence from historic- and prehistoric-era ritual burials of polar bears indicate that polar bears have been relatively abundant there for at least the last 2,000 years, as expected due to the prominent polynya that today forms along the southern coast. It is also possible that at times during the past 4,000 years, a polynya of some size formed off northern Greenland in the Wandel Sea, making it possible for humans living in Peary Land to add seals and polar bears to their usual diet of terrestrial species such as Arctic hare (Lepus arcticus), Arctic fox (Vulpes lagopus), and muskox (Ovibos moschatus) (Darwent 2003). However, it is also possible that historically, both bears and people in northern Greenland travelled to the nearby NE Water to hunt seals.
In contrast, there are few archaeological bones and no fossil remains of polar bears found in Hudson Bay but this dearth of records is consistent with the dynamic geological and sea level history of the region.
Most ancient polar bear remains from fossil and archaeological contexts before A.D. 1910 date within the Holocene and derive from human habitation sites within the current range of the species. Extralimital specimens have been documented in the north Atlantic during the late Pleistocene and in the southern Bering Sea during the middle Holocene, both of which were cold periods when Arctic sea ice expanded to the south of modern limits in winter. The earliest evidence for the return of polar bears to the Arctic after the end of the LGM dates to the early HCO (ca. 8,000 a BP) in both the Atlantic and Pacific sectors, even though bowhead whales in the Pacific returned almost 2,000 years earlier. Unfortunately, none of the skeletal evidence is adequate for determing if changes occurred in abundance of polar bears in response to short-term climatic changes. However, the geographic distribution of ancient remains, from both fossil and archaeological contexts, indicates that polynyas have been important ice-edge habitats for polar bears since the last Interglacial period, as they are today.
The author gratefully acknowledges feedback on an earlier draft of this paper by James Woollet, Kim Aaris-Sørensen, and James Savelle, as well as for the additional references they supplied. Thanks also to Diane Hanson for constructive comments on the submitted manuscript, Aaris-Sørensen and the Natural History Museum of Denmark (University of Copenhagen) for permission to reprint the Asdal mandible photo (Figure 3), Rob Losey for providing the Russian archaeological report for the Tiutei-Sale 1 site, and Anne Birgitte Gotfredsen for Scoresby Sound data.
Some funding for researching and writing this paper came from Pacific Identifications Inc. of Victoria, British Columbia, Canada and is gratefully acknowledged.
The author has no competing interests to declare.
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