Polar Bear Fossil and Archaeological Records from the Pleistocene and Holocene in Relation to Sea Ice Extent and Open Water Polynyas

Authors: {'first_name': 'Susan J.', 'last_name': 'Crockford'}


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.

Keywords: Ursus maritimusArcticZhokov Islandextralimital recordsecologyskeletal remains 
 Accepted on 25 Jan 2022            Submitted on 14 Aug 2021

1. Introduction

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).

Fossil and archaeological polar bear remains, as per Table 1
Figure 1 

Fossil and archaeological polar bear remains across the Arctic, within the current species range (blue line) vs. extralimital finds as per International Union for the Conservation of Nature, Species Survival Commission, Polar Bear Specialist Group (IUCN/SSC PBSG). See text for explanation of ‘fossil’ remains. Legend of site numbers (see Table 1 for more details):

1. NE Point, St. Paul Is.; 2. St. Lawrence Is.; 3. Pottery House, St. Matthew Is.; 4. Walakpa; 5. St. Lawrence Is. (3 sites); 6. St. Lawrence Is. (Kukulik); 7. St. Lawrence Is. (Hillside site); 8. Cape Espenberg; 9. Qagnax Cave, St. Paul Is.; 10. Bogoslov Cave, St. Paul Is.; 11. Margaret Bay (UNL-48); 12. Washount (NjVi-2, H3); 13. Agvik (OkRn1); 14. Nelson River site; 15. Co-Op (OdPp-2, H1, H5); 16. Lady Franklin Pt. (NdPd-2); 17. Pingiqqalik (NgHd-1); 18. Naujan (MdHs-1); 19. Sadlermiut (KkHh-1); 20. Qijurittuq (IbGk-31); 21. Staffe Is.; 22. Nachvak Fjord group; 23. Oakes Bay (HeCg-8); 24. Iglosiatik Is.; 25. JfEl-10, Quebec; 26. Talaguak, Baffin Is.; 27. Outer Frobisher Bay (3sites); 28. Cumberland Sound (LlDj-1); 29. Hazard Inlet group (3 sites); 30. Learmonth (PeJr-1); 31. Porden Pt. group (3 sites); 32. Porden Pt. (RbJq-6); 33. Peale Pt. (KkDo-1); 34. Sanirajak (NeHd-1); 35. Kuukpak (NiTs-1, H1); 36. Amundsen Gulf group (4 sites); 37. Bell site (NiNg-2); 38. Port Refuge (Snowdrift); 39. Hornby Head (RbJq-1); 40. Brooman Point; 41. Bache Peninsula, 3 sites; 42. Skraeling Is. (SfFk-4, H 14–16); 43. Cape Garry (PcJq-5); 44. Victoria Is.; 45. Victoria Is.; 46. Cape Richard Collinson; 47. Seahorse Gully (IeKn 6); 48. Port Refuge (upper beach); 49. Port Refuge (Gull Cliff); 50. Port Refuge (Lower Beach); 51. Gulf of Boothia; 52. Baillie Island; 53. Scoresby Sound (‘House of Beads’); 54. Scoresby Sound (Skærgårdshalvøen 1); 55. Nugarsuk; 56. Walrus Is.; 57. Clavering Is. (4 sites); 58. Fladstrand; 59. Dødemandsbugten (3 sites); 60. Sephus Müller Næs; 61. Qeqertaaraq; 62. Washington Land; 63. Washington Land; 64. Kolnæs; 65. Vandfeldsnaes; 66. Saqqaq; 67. Solbakken; 68. Adam C. Knuth; 69. Pearylandville; 70. Sønderland; 71. Norde Eskimonœsset; 72. Nuulliit; 73. Cape Schmidt; 74. Yamal Peninsula; 75. Vaygach Is.; 76. Tiutei-Sale 1; 77. Dezhnevo; 78. Cape Schmidt; 79. Cape Schmidt; 80. Cape Baranov; 81. Mainland south of Laptev Strait; 82. Mainland, near Tikai; 83. Vaygach Island; 84. Ekven; 85. Devil’s Gorge; 86. Zhokhov Is.; 87. Mordy-Yahk River; 88. Pechora River; 89. Iceland; 90. Asdal DEN; 91. Kuröd Bohuslän; 92. Nedre Kuröd Bohuslän; 93. Hisingen; 94. Kärraberg; 95. Östra Karup; 96. Kullaberg; 97. Svenskøya; 98. Svalbard; 99. Finnøy; 100. Nordcemgrotta; 101. Hamnsundhelleren; 102. Nordcemgrotta; 103. Poolepynten; 104. Kew Bridge.

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).

Table 1

Fossil and archaeological polar bear remains by approximate chronological date, by country; site numbers as in Figure 1. Abbreviations: USA, United States of America; CAN, Canada; GRE, Greenland; RUS, Russia; ICE, Iceland; DEN, Denmark; SWE, Sweden; NOR, Norway; UK, United Kingdom; F, female; M, male; R, right, L, left; H, house; Is., Island; Pt., Point; ca., approximately; LIA, Little Ice Age; MWP, Medieval Warm Period; DAC, Dark Age Cold; RWP, Roman Warm Period; HCO, Holocene Climatic Optimum; NEO, Neoglacial; YD, Younger Dryas. † indicates fossil specimens, see text for explanation.


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

† Indicates ‘fossil’ specimens, see text.

‡ These are calibrated radiocarbon years BP unless indicated otherwise. Carbon 14 dates on polar bear bone are corrected for marine reservoir effect unless indicated otherwise; one historic specimen (#1) is a calendar date (e.g., AD 1875) and one (#103) is an IRSL (‘infrared-stimulated luminescence’) date on sediments.

§ Relative geological and climatological time periods (a BP) defined as: Pleistocene 2,500,000–11,700; MIS 5e/Eemian Interglacial 130,000–115,000; Last Glacial Maximum (LGM) 30,000–19,700; Holocene 11,700–1950; Younger Dryas (YD) 12,900–11,700; Early Holocene 11,700–8,200; Middle Holocene 8,300–4,200; Holocene Climatic Optimum (HCO) 9,000–5,500; Neoglacial 5,500–2,000; Roman Warm Period (RWP) 2,000–1,500; Dark Ages Cold (DAC) 1,500–1,050; Medieval Warm Period (MWP) 1,050–650; Little Ice Age (LIA) 650–50 (Alley 2000; Cohen et al. 2013; Kaufman et al. 2004; Lamb 1982; Marcott et al. 2009; Polyak et al. 2010; Soon & Baliunas 2004).

1. The ‘Hamnsund Interstadial’ was a short-lived ice retreat originally dated to 22–19k uncal a BP in W. Norway. See text for discussion.

2. The Ålesund Interstadial was a short- lived ice treat dated to 30k a BP in W. Norway (Hufhammer 2001).

3. The species identification of this specimen has been disputed but not resolved, see text for details.

Known and proposed polynyas across the Arctic
Figure 2 

Known recurrent major and flaw polynyas across the Arctic (after Barber et al. 2001; Grønnow et al. 2011; Jackson et al 2020; Kern 2008; Morales Maqueda et al. 2004; Pedersen et al. 2010; Smedsrud et al. 2006; Speer et al. 2017; Stirling and Cleator 1981; Stringer and Groves 1991). The ‘Wandel Water’ (Z) is a proposed polynya that forms under certain climatic conditions in the Wandel Sea (e.g., Schweiger et al. 2021).

2. Materials & Methods

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.

3. Results

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).

3.1 Extralimital fossil records

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.

Polar bear mandible from Denmark, dated 12.4-12.9k cal a BP
Figure 3 

Left mandible of polar bear found at Kjul Å near Asdal in northern Jylland (Denmark). Age: ca. 12.4–12.9k cal a BP (Photo by Geert Brovad, courtesy Natural History Museum of Denmark).

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.

3.2 Fossil records within modern range

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).

3.3 Extralimital archaeological records

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).

3.4 Archaeological records within range

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.

Table 2

Select archaeological assemblages with polar bear remains expressed as the relative proportion of the total number of identified specimens (NISP) of all mammals not including whale. † indicates percentage based on minimum number of individuals rather than bone count. Map reference information as in Table 1.


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).

Table 3

Fossil and archaeological sites near polynyas, by site number (as per Table 1) and polynya code (as per Figure 2) according to count of polar bear remains (those with 1–9 vs. >10 bones). P indicates ‘present’.


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)

85 E Wrangel Island 1

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

28 U Cumberland Sound 3

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).

3.5 Change through time

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.

3.6 Absence of data

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).

4. Discussion

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.

5. Conclusion

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.

Funding Information

Some funding for researching and writing this paper came from Pacific Identifications Inc. of Victoria, British Columbia, Canada and is gratefully acknowledged.

Competing Interests

The author has no competing interests to declare.


  1. Aaris-Sørensen, K. 2009. Diversity and dynamics of the mammalian fauna in Denmark throughout the last glacial-interglacial cycle, 115–0 kry BP. Copenhagen: Fossils and Strata 57. 

  2. Aaris-Sørensen, K and Petersen, KS. 1984. A Late Weichselian find of polar bear (Ursus maritimus Phipps) from Denmark and reflections on the paleoenvironment. Boreas, 13(1): 29–33. DOI: 

  3. Alley, RB. 2000. The Younger Dryas cold interval as viewed from central Greenland. Quaternary Science Reviews, 19(1–5): 213–226. DOI: 

  4. Amos, J. 2007. Ancient polar bear jawbone found. BBC News, 10 December. URL 

  5. Amstrup, SC. 2003. Polar bear (Ursus maritimus). In Feldhamer, GA, Thompson, BC and Chapman, JA (eds.) Wild Mammals of North America. Baltimore: Johns Hopkins University Press. 587–610. 

  6. Amstrup, SC and Gardner, C. 1994. Polar bear maternity denning in the Beaufort Sea. The Journal of Wildlife Management, 58(1): 1–10. DOI: 

  7. Andersen, M, Derocher, AE, Wiig, Ø, et al. 2012. Polar bear (Ursus maritimus) maternity den distribution in Svalbard, Norway. Polar Biology, 35: 499–508. DOI: 

  8. Andreasen, C. 1997. The prehistory of the coastal areas of Amdrup Land and Holm Land adjacent to the northeast water polynya: an archaeological perspective. Journal of Marine Systems, 10(1–4): 41–46. DOI: 

  9. Andreev, AA, Grosse, G, Schirrmeister, L, et al. 2009. Weischelian and Holocene palaeoenvironmental history of the Bol’shoy Lyakhovsky Island, New Siberian Archipelago, Arctic Siberia. Boreas, 38: 72–110. DOI: 

  10. Arnold, CD. 1986. In search of Thule pioneers. In Bielawski, E, Kobelka, C and Janes, RR (eds.) Thule Pioneers. Yellowknife: Occasional Papers of the Prince of Wales Northern Heritage Centre 2. 1–93. 

  11. Arnold, CD. 2000. The Hillside Site, St. Lawrence Island, Alaska: An examination of collections from the 1930s, by Don E. Dumond. Arctic, 53(2): 196–198. DOI: 

  12. Áskelsson, J. 1938. Um íslenzk dýr og jurtir frá jökultíma. Náttúrufræðingurinn, 8(1): 6–7. [Icelandic] 

  13. Atkinson, N. 2009. A 10,400-year-old bowhead whale (Balaena mysticetus) skull from Ellef Ringnes Island, Nunavut: implications for sea-ice conditions in high Arctic Canada at the end of the last glaciation. Arctic, 62(1): 38–44. DOI: 

  14. Barber, DG, Hanesiak, JM, Chan, W, et al. 2001. Sea-ice and meteorological conditions in northern Baffin Bay and the North Water polynya between 1979 and 1996. Atmosphere-Ocean, 39(3): 343–359. DOI: 

  15. Barnes, I, Matheus, P, Shapiro, B, et al. 2002. Dynamics of Pleistocene population extinctions in Beringian brown bears. Science, 295(5563): 2267–2270. DOI: 

  16. Bechshøft, TØ, Sonne, C, Rigét, FF, et al. 2008. Differences in growth, size and sexual dimorphism in skulls of East Greenland and Svalbard polar bears (Ursus maritimus). Polar Biology, 31: 945–958. DOI: 

  17. Bennike, O. 1991. Marine mammals in Peary Land, northern Greenland. Polar Record, 27(163): 357–359. DOI: 

  18. Bennike, O. 1997. Quaternary vertebrates from Greenland: a review. Quaternary Science Reviews, 16(8): 899–909. DOI: 

  19. Bennike, O. 2002. Late Quaternary history of Washington Land, North Greenland. Boreas, 31(3): 260–272. DOI: 

  20. Bennike, O, Leth, JO, Jensen, JB, et al. 2014. Arctic plant remains of Weichselian age from the Danish North Sea. GEUS Geological Survey of Denmark and Greenland Bulletin, 31: 43–45. DOI: 

  21. Berglund, BE, Håkansson, S and Lepiksaar, J. 1992. Late Weichselian polar bear (Ursus maritimus Phipps) in southern Sweden. Sveriges Geologiska Undersökning, Series Ca, 81: 31–42. 

  22. Betts, MW and Friesen, TM. 2006. Declining foraging returns from an inexhaustible resource? Abundance indices and beluga whaling in the western Canadian Arctic. Journal of Anthropological Archaeology, 25(1): 59–81. DOI: 

  23. Bilodeau, G, de Vernal, A, Hillaire-Marcel, C, et al. 1990. Postglacial paleoceanography of Hudson Bay: stratigraphic, microfaunal, and palynological evidence. Canadian Journal of Earth Science, 27(7): 946–963. 

  24. Bland, RL. 2008. The early sites of Cape Baranov. In Okladnikov, AP and Beregovaya, NA (eds./trans.) Drevnie poseleniya Baranova Mysa. Novosibirsk: Nauka, 1971 [including appendix by N.K. Vereshchagin, ‘The remains of animals from houses at Cape Baranov, east of the mouth of the Kolyma River’] Anchorage: Alaska. US National Parks Service. 

  25. Blystad, P, Thomsen, H, Simonsen, A, et al. 1983. Find of a nearly complete Late Weichselian polar bear skeleton, Ursus maritimus Phipps, at Finnøy, southwestern Norway: a preliminary report. Norsk Geologisk Tidskrift, 63: 193–197. 

  26. Boeskorov, GG, Tikhonov, AN, Protopopov, AV, et al. 2018. New records of Holocene polar bear and walrus (Carnivora) in the Russian Arctic. Russian Journal of Theriology, 17(2): 68–77. DOI: 

  27. Bradley, RS and England, JH. 2008. The Younger Dryas and the sea of ancient ice. Quaternary Research, 70(1): 1–10. DOI: 

  28. Briffa, KR, Melvin, TM, Osborn, TJ, et al. 2013. Reassessing the evidence for tree-growth and inferred temperature change during the Common Era in Yamalia, northwestern Siberia. Quaternary Science Reviews, 72: 83–107. DOI: 

  29. Brown, ZW, van Dijken, GL and Arrigo, KR. 2011. A reassessment of primary production and environmental change in the Bering Sea. Journal of Geophysical Research, 116(C8): C08014. DOI: 

  30. Buckley, JR, Gammelsrod, T, Johannessen, OM, et al. 1979. Upwelling: Oceanic structure at the Arctic ice pack in winter. Science, 203(4376): 165–167. DOI: 

  31. Caissie, BE, Brigham-Grette, J, Lawrence, KT, et al. 2010. Last Glacial Maximum to Holocene sea surface conditions at Umnak Plateau, Bering Sea, as inferred from diatom, alkenone, and stable isotope records. Paleoceanography and Paleoclimatology, 25(1): PA1206. DOI: 

  32. Calvert, W, Stirling, I, Schweinsburg, RE, et al. 1986. Polar bear management in Canada 1982–84. In Polar Bears: Proceedings of the 9th meeting of the Polar Bear Specialists Group IUCN/SSC, Edmonton, Canada on 9–11 August, 1985. pp 19–34. 

  33. Castro de la Guardia, L, Myers, PG, Derocher, AE, et al. 2017. Sea ice cycle in western Hudson Bay, Canada, from a polar bear perspective. Marine Ecology Progress Series, 564: 225–233. DOI: 

  34. Cheng, H, Zhang, H, Spötl, C, et al. 2020. Timing and structure of the Younger Dryas event and its underlying climate dynamics. Proceedings of the National Academy of Science USA, 117(38): 23408–23417. DOI: 

  35. Cohen, KM, Finney, SC, Gibbard, PL, et al. 2013. The ICS International Chronostratigraphic Chart (updated version 2018–07). Episodes, 36: 199–204. DOI: 

  36. Collins, HB. 1937. Archaeology of St. Lawrence Island, Alaska. Washington: Smithsonian Institution. 

  37. Collins, HB. 1956. The T1 site at Native Point, Southampton Island, N.W.T. Anthropological Papers of the University of Alaska, 4: 63–89. 

  38. Collins, HB. 1981. Record of animal bones recovered at Sadlermiut, 1954–1955. Ottawa: Canadian Museum of History 1922. 

  39. Condon, A and Winsor, P. 2012. Meltwater routing and the Younger Dryas. Proceedings of the National Academy of Science USA, 109(49): 19928–19933. DOI: 

  40. Connolly, R and Connolly, M. 2014. Global temperature changes of the last millennium. Open Peer Review Journal, 16 (Clim. Sci.), ver. 1.0. DOI: 

  41. Crockford, SJ. 2008. Be careful what you ask for: Archaeozoological evidence of mid-Holocene climate change and implications for the origins of Arctic Thule. In Clark, G, Leach, F and O’Connor, S (eds.) Islands of Inquiry: Colonization, Seafaring, and the Archaeology of Marine Landscapes. Canberra: ANU E Press Terra Australis 29. 113–131. DOI: 

  42. Crockford, SJ and Frederick, G. 2007. Sea ice expansion in the Bering Sea during the Neoglacial: evidence from archaeozoology. The Holocene, 17(6): 699–706. DOI: 

  43. Crockford, SJ and Frederick, G. 2011. Neoglacial sea ice and life history flexibility in ringed and fur seals. In Braje, T and Torrey, R (eds.) Human and Marine Ecosystems: Archaeology and Historical Ecology of Northeastern Pacific Seals, Sea Lions, and Sea Otters. Los Angeles: University of California Press. 65–90. DOI: 

  44. Crockford, SJ, Frederick, G and Wigen, R. 2002. The Cape Flattery fur seal: an extinct species of Callorhinus in the Eastern North Pacific? Canadian Journal of Archaeology, 26: 152–174. 

  45. Darwent, CA. 2003. Appendix 1. The zooarchaeology of Peary Land and adjacent areas. In Grønnow, B and Jensen, JF (eds.) The northernmost ruins of the globe: Eigil Knuth’s archaeological investigations in Peary Land and adjacent areas. (Meddelelser om Grønland/Man & Society 29). Copenhagen: Danish Polar Center. 342–347. 

  46. Davis, B. 2001. Sea mammal hunting and the Neoglacial: an archaeofaunal study of environmental change and subsistence technology at Margaret Bay, Unalaska. In Dumond, DE (ed.) Recent archaeology in the Aleut Zone of Alaska. (University of Oregon Anthropology Papers 58). Eugene: University of Oregon. 71–85. 

  47. Davison, J, Ho, SYW, Bray, SC, et al. 2011. Late-Quaternary biogeographic scenarios for the brown bear (Ursus arctos), a wild mammal model species. Quaternary Science Reviews, 30(3–4): 418–430. DOI: 

  48. Degerbøl, M. 1936. The former Eskimo habitation in Kangerdlugssuak District East Greenland. Meddelelser om Grønland, 104(10): 40–45. 

  49. DeMaster, DP, Kingsley, MCS and Stirling, I. 1980. A multiple mark and recapture estimate applied to polar bears. Canadian Journal of Zoology, 58(4): 633–638. DOI: 

  50. Denny, ALS. 1859. Observations on the distribution of the extinct bears of Britain, with especial reference to a supposed new species of fossil bear from Ireland. Proceedings of the Yorkshire Geological Society, 4(Jan): 338–358. DOI: 

  51. Derocher, AE and Stirling, I. 1992. The population dynamics of polar bears in western Hudson Bay. In McCullough, DR and Barrett, RH (eds.) Wildlife 2001: Populations. London: Elsevier. 1150–1159. DOI: 

  52. Derocher, AE and Stirling, I. 1995. Temporal variation in reproduction and body mass of polar bears in western Hudson Bay. Canadian Journal of Zoology, 73(9): 1657–1665. DOI: 

  53. Desjardins, SPA. 2013. Evidence for intensive walrus hunting by Thule Inuit, northwest Foxe Basin, Nunavut, Canada. Anthropozoologica, 48(1): 37–51. DOI: 

  54. Desjardins, SPA. 2018. Neo-Inuit strategies for ensuring food security during the Little Ice Age climate change episode, Foxe Basin, Arctic Canada. Quaternary International, 549: 163–175. DOI: 

  55. Desrosiers, PM, Lofthouse, S, Bhiry, N, et al. 2010. The Qijurittuq site (IbGk-3), Eastern Hudson Bay: An IPY Interdisciplinary Study. Geografisk Tidsskrift-Danish Journal of Geography, 110: 227–243. DOI: 

  56. de Vernal, A, Hillaire-Marcel, C, Le Duc, C, et al. 2020. Natural variability of the Arctic Ocean sea ice during the present interglacial. Proceedings of the National Academy of Sciences USA, 117(42): 26069–26075. DOI: 

  57. Dikov, NN. 1988. The earliest sea mammal hunters of Wrangell Island. Arctic Anthropology, 25: 80–93. 

  58. Dredge, LA. 1992. Field guide to the Churchill Region, Manitoba: Glaciations, sea level changes, permafrost landforms, and archaeology of the Churchill and Gillam areas. (Miscellaneous Report 53). Ottawa: Geological Survey of Canada. DOI: 

  59. Dumond, DE. 1998. The Hillside site, St. Lawrence Island, Alaska. (University of Oregon Anthropology Papers 55). Eugene: University of Oregon. 

  60. Dyke, AS and England, JH. 2003. Canada’s most northerly postglacial bowhead whales (Balaena mysticetus): Holocene sea-ice conditions and polynya development. Arctic, 56(1): 14–20. DOI: 

  61. Dyke, AS, Hooper, J and Savelle, JM. 1996. A history of sea ice in the Canadian Arctic Archipelago based on postglacial remains of bowhead whale (Balaena mysticetus). Arctic, 49(3): 235–255. DOI: 

  62. Dyke, AS, Hooper, J, Harington, CR, et al. 1999. The late Wisconsinan and Holocene record of walrus (Odobenus rosmarus) from North America: a review with new data from arctic and Atlantic Canada. Arctic, 52(2): 160–181. DOI: 

  63. Dyke, AS, Savelle, JM and Johnson, DS. 2011. Paleoeskimo demography and Holocene sea-level history, Gulf of Boothia, Arctic Canada. Arctic, 64(2): 151–168. DOI: 

  64. Dyke, AS, Savelle, JM, Szpak, P, et al. 2018. An assessment of marine reservoir corrections for radiocarbon dates on walrus from the Foxe Basin region of Arctic Canada. Radiocarbon, 61(1): 1–15. DOI: 

  65. Durner, GM and Amstrup, SC. 1995. Movements of a polar bear from northern Alaska to northern Greenland. Arctic, 48(4): 338–341. DOI: 

  66. Edwards, CJ, Suchard, MA, Lemey, P, et al. 2011. Ancient hybridization and an Irish origin for the modern polar bear matriline. Current Biology, 21(15): 1251–1258. DOI: 

  67. Edwards, CJ, Ho, SYW, Barnes, R, et al. 2014. Continuity of brown bear maternal lineages in northern England through the Last-glacial period. Quaternary Science Reviews, 96: 131–139. DOI: 

  68. Elliott, HW. 1875. Polar bears on St. Matthew Island. Harper’s Weekly Journal of Civilization, May 1. New York: Harper and Brothers. 

  69. Elliott, HW and Coues, E. 1875. A report upon the condition of affairs in the territory of Alaska. Washington: US Government Printing Office. DOI: 

  70. Fay, FH. 1982. Ecology and biology of the Pacific walrus, Odobenus rosmarus divergens Illiger. (North American Fauna, Number 74). Washington: US Fish and Wildlife Service. DOI: 

  71. Fisher, D, Dyke, A, Koerner, R, et al. 2006. Natural variability of arctic sea ice over the Holocene. EOS, Transactions of the American Geophysical Union, 87(28): 273–280. DOI: 

  72. Frey, KE, Moore, GWK, Cooper, LW, et al. 2015. Divergent patterns of recent sea ice cover across the Bering, Chukchi, and Beaufort seas of the Pacific Arctic Region. Progress in Oceanography, 136: 32–49. DOI: 

  73. Friesen, TM, Finkelstein, SA and Medeiros, AS. 2020. Climate variability of the Common Era (AD 1–2000) in the eastern North American Arctic: Impacts on human migrations. Quaternary International, 549: 142–154. DOI: 

  74. Friesen, TM and Hunston, JR. 1994. Washout – the final chapter: 1985–86 NOGAP salvage excavations on Herschel Island. Canadian Archaeological Association Occasional Papers, 2: 39–60. 

  75. Frink, L, Corbett, D, Rosebrough, A, et al. 2001. The archaeology of St. Matthew Island, Bering Sea. Alaska Journal of Anthropology, 1(1): 131–137. 

  76. Fedorova, NV, Kosintsev, PA and Fitzhugh, WW. 1998. ‘Ushedshie v Kholmy’: Kul’tura Naselenii Poberezhii Severo-Zapadnogo Iamala v Zheleznom Veke [‘Gone into the Hills’: Culture of the Northwestern Iamal Coast Population in the Iron Age]. Ekaterinburg: Izd-vo Ekaterinburg. [Russian] 

  77. Garner, GW, Belikov, SE, Stishov, MS, et al. 1994. Dispersal patterns of maternal polar bears from the denning concentration on Wrangel Island. International Conference on Bear Research and Management, 9(1): 401–410. DOI: 

  78. Gotfredsen, AB. 1992. Nyt fra Saqqaq-kulturen. Forskning i Grønland/Tusaat, 92: 41–45. [Danish] 

  79. Gotfredsen, AB. 2010. Faunal remains from the Wollaston Forland – Clavering Ø region, northeast Greenland – Thule culture subsistence in a High Arctic polynya and ice-edge habitat. Geografisk Tidsskrift/Danish Journal of Geography, 110(2): 175–200. DOI: 

  80. Gotfredsen, AB, Appelt, M and Hastrup, K. 2018. Walrus history around the North Water: Human – animal relations in a long-term perspective. Ambio, 47(Suppl 2): S193-S212. DOI: 

  81. Griffin, D. 2008. The archaeology of St. Matthew Island, Bering Sea. Alaska Journal of Anthropology, 2(1–2): 84–99. 

  82. Grønnow, B. 2016. Living at a High Arctic polynya: Inughuit settlement and subsistence around the North Water during the Thule Station Period, 1910- 53. Arctic, 69(5) (Suppl. 1): 1–15. DOI: 

  83. Grønnow, B, Gulløv, HC, Jakobsen, BH, et al. 2011. At the edge: high Arctic walrus hunters during the little ice age. Antiquity, 85(329): 960–977. DOI: 

  84. Grønnow, B and Jensen, JF. 2003. The northernmost ruins of the globe: Eigil Knuth’s archaeological investigations in Peary Land and adjacent areas. (Meddelelser om Grønland. Man & Society 29). Copenhagen: Danish Polar Center. 

  85. Gusev, SV, Zaqoroulko, AV and Porotov, AV. 1999. Sea mammal hunters of Chukotka, Bering Strait: recent archaeological results and problems. World Archaeology, 30(3): 354–369. DOI: 

  86. Guthrie, RD. 2004. Radiocarbon evidence of mid-Holocene mammoths stranded on an Alaskan Bering Sea Island. Nature, 429: 746–749. DOI: 

  87. Gyllencreutz, R. 2005. Holocene and Latest Glacial Paleoceanography in the North-Eastern Skagerrak. Unpublished thesis (PhD), Stockholm University. 

  88. Harington, CR. 1988. Marine mammals of the Champlain Sea, and the problem of whales in Michigan. Geological Association of Canada Special Paper, 35: 225–240. 

  89. Harington, CR. 2003. Annotated Bibliography of Quaternary Vertebrates of Northern North America. Toronto: University of Toronto Press. 

  90. Harington, CR. 2008. The evolution of Arctic marine mammals. Ecological Applications, 18(sp2): S23-S40. DOI: 

  91. Hastrup, K, Mosbech, A and Grønnow, B. 2018. Introducing the North Water: Histories of exploration, ice dynamics, living resources, and human settlement in the Thule Region. Ambio, 47: S162-S174. DOI: 

  92. Heaton, TH and Grady, F. 2003. The Late Wisconsin vertebrate history of Prince of Wales Island, Southeast Alaska. In Schubert, BW, Mead, JI and Graham, RW (eds.) Ice Age Cave Faunas of North America. Bloomington: Indiana University Press. 17–53. 

  93. Heide-Jørgensen, MP, Richard, P, Ramsay, M, et al. 2002. Three recent ice entrapments of Arctic cetaceans in West Greenland and the eastern Canadian High Arctic. NAMMCO Scientific Publications, 4: 143–148. DOI: 

  94. Heide-Jørgensen, MP, Sinding, M-HS, Nielsen, NH, et al. 2016. Large numbers of marine mammals winter in the North Water polynya. Polar Biology, 39: 1605–1614. DOI: 

  95. Henderson, EM, Derocher, AE, Lunn, NJ, et al. 2021. Polar bear Ursus maritimus use of the western Hudson Bay flaw lead. Marine Ecology Progress Series, 664: 227–242. DOI: 

  96. Henshaw, AS. 1995. Central Inuit Household Economies: Zooarchaeology, Environmental, and Historical Evidence from Outer Frobisher Bay, Baffin Island, Canada. Unpublished thesis (PhD), Harvard University. 

  97. Henshaw, AS. 2003. Polynyas and ice edge habitats in cultural context: archaeological perspectives from southeast Baffin Island. Arctic, 56(1): 1–13. DOI: 

  98. Hjort, C. 1997. Glaciation, climate history, changing marine levels and the evolution of the Northeast Water polynya. Journal of Marine Systems, 10(1–4): 23–33. DOI: 

  99. Hörner, T, Stein, R and Fahl, K. 2018. Paleo-sea ice distribution and polynya variability on the Kara Sea shelf during the last 12 ka. Arktos, 4: 1–16. DOI: 

  100. Howse, LR. 2019. Hunting technologies and archaeofaunas: societal differences between hunter-gatherers of the Eastern Arctic. Journal of Archaeological Method and Theory, 26: 88–111. DOI: 

  101. Hufthammer, AK. 2001. The Weichselian (c. 115,000–10,000 B.P.) vertebrate fauna of Norway. Bollettino della Societá Paleontologica Italiana, 40: 201–208. 

  102. Ingolfsson, Ó and Wiig, Ø. 2009. Late Pleistocene fossil find in Svalbard: the oldest remains of a polar bear (Ursus maritiumus Phipps, 1744) ever discovered. Polar Research, 28(3): 455–462. DOI: 

  103. Jackson, R, Bang Kvorning, A, Limoges, A, et al. 2021. Holocene polynya dynamics and their interaction with ocean heat transport in northernmost Baffin Bay. Nature Science Reports, 11: 10095. DOI: 

  104. Jackson, R, Bang Kvorning, A, Pearce, C, et al. 2020. High Arctic polynas in a changing climate. European Geosciences Union, 22nd EGU General Assembly, 4–8 May, id.17959. DOI: 

  105. Jeppesen, E, Appelt, M, Hastrup, K, et al. 2018. Living in an oasis: Rapid transformations, resilience, and resistance in the North Water Area societies and ecosystems. Ambio, 47: 296–309.DOI: 

  106. Kaplan, S and Woollett, J. 2016. Labrador Inuit: Thriving on the Periphery of the Inuit World. In Friesen, M and Mason, O (eds.) The Oxford Handbook of the Prehistoric Arctic. Oxford: Oxford University Press. 851–872. DOI: 

  107. Kassens, H and Thiede, J. 1994. Climatological significance of Arctic sea ice at present and in the past. In Kassens, H et al. (eds.) Russian-German Cooperation in the Siberian Shelf Seas: Geo-System Laptev Sea. Bremerhaven: Berichte zur Polarforschung 144. 81–85. 

  108. Kaufman, DS, Ager, TA, Anderson, NJ, et al. 2004. Holocene thermal maximum in the western Arctic (0-1800W). Quaternary Science Reviews, 23(5–6): 529–560. DOI: 

  109. Kavry, VI, Kochnev, AA, Nikiforov, VV, et al. 2006. Cape Vankarem – nature-ethnic complex at the Arctic coast of Chukotka (northeastern Russia). In Marine Mammals of the Holarctic, Papers of the Fourth International Conference, Saint Petersburg on 10–14 September 2006. 227–230. 

  110. Kern, S. 2008. Polynya area in the Kara Sea, Arctic, obtained with microwave radiometry for 1979–2003. IEEE Geoscience and Remote Sensing Letters, 5(2): 171–175. DOI: 

  111. Kishchinskiy, AA. 1976. The polar bear. In Large Carnivores. Moscow. [no page number cited] 

  112. Klein, DR and Sowls, A. 2011. History of polar bears as summer residents on the St. Matthew Islands, Bering Sea. Arctic, 64(4): 429–436. DOI: 

  113. Knuth, E. 1978. The ‘Old Nûgdlît Culture’ site at Nûgdlît Peninsula, Thule District, and the ‘Mesoeskimo’ site below it. Folk, 19–20: 15–47. 

  114. Kochnev, AA. 2002. Factors causing Pacific mortality on the coastal haulouts of Wrangel Island. In Aristov, AA et al. (eds.), O Romanenko (trans.) Marine Mammals: Results of Research conducted in 1995–1998. Moscow: Marine Mammal Council. 191–215. 

  115. Kochneva, S. 2007. Polar bear in material and spiritual culture of the native peoples of Chukotka: bibliographic information. Report prepared for the Association of Traditional Marine Mammal Hunters of Chukotka, Alaska Nanuuq Commission and Pacific Fisheries Research Center, Chukotka Branch, Anadyr. [translated by O Romanenko, 2008] 

  116. Kokorowski, HD, Anderson, PM, Sletten, RS, et al. 2008. Late Glacial and early Holocene climatic changes based on a multiproxy lacustrine sediment record from northeast Siberia. Arctic, Antarctic, and Alpine Research, 40(3): 497–505. DOI:[KOKOROWSKI]2.0.CO;2 

  117. Kotar, K. 2016. Variability in Thule Inuit Subsistence Economy: A Faunal Analysis of OkRn-1, Banks Island, NWT. Unpublished thesis (PhD), University of Western Ontario. 

  118. Kroon, A, Jakobsen, BH and Pedersen, JBT. 2010. Coastal environments around Thule settlements in Northeast Greenland. Geografisk Tidsskrift-Danish Journal of Geography, 110: 143–154. DOI: 

  119. Kurtén, B. 1968. On Pleistocene Mammals of Europe. London: Weidenfeld and Nicolson. 

  120. Kurtén, B. 1988. On Evolution and Fossil Mammals. New York: Columbia University Press. 

  121. Laidre, KL and Stirling, I. 2020. Grounded icebergs as maternity denning habitat for polar bears (Ursus maritimus) in North and Northeast Greenland. Polar Biology, 43: 937–943. DOI: 

  122. Laidre, KL, Stirling, I, Estes, JA, et al. 2018. Historical and potential future importance of large whales as food for polar bears. Frontiers in Ecology and the Environment, 16(9): 515–524. DOI: 

  123. Lamb, HH. 1982. Climate, History, and the Modern World. London: Methuen. 

  124. Lambeck, K, Purcell, A, Zhao, J, et al. 2010. The Scandinavian Ice Sheet: from MIS 4 to the end of the Last Glacial Maximum. Boreas, 39: 410–435. DOI: 

  125. Lamy, P and Spitery, J. 1991. Analyses Des Vestiges Osseux de OdPp-2, Ile Victoria, N.W.T., Étude Préliminaire. (Unpublished MS 4329). Ottawa: Canadian Museum of Civilization. 

  126. Lan, T, Leppälä, K, Tomlin, C, et al. 2022. Insights into bear evolution from a Pleistocene bear genome. Biorxiv Preprint, April 20. DOI: 

  127. Lauritzen, SE, Nese, H, Lie, RW, et al. 1996. Interstadial/interglacial fauna from Norcemgrotta, Kjøpsvik, north Norway. In Lauritzen, SE (ed.) Climate change: The Karst Record. Charles Town: Karst Waters Institute Special Publication 2. 89–92. 

  128. Lindqvist, C, Bachmann, L, Andersen, LW, et al. 2009. The Laptev Sea walrus Odobenus rosmarus laptevi: an enigma revisited. Zoologica Scripta, 38(2): 113–127. DOI: 

  129. Lindqvist, C, Schuster, SC, Sun, Y, et al. 2010. Complete mitochondrial genome of a Pleistocene jawbone unveils the origin of polar bear. Proceedings of the National Academy of Sciences USA, 107(11): 5053–5057. DOI: 

  130. Lofthouse, SE. 2003. A Taphonomic Treatment of Thule Zooarchaeological Materials from Diana Bay, Nunavik (Arctic Quebec). Unpublished thesis (PhD), McGill University. 

  131. Ludwig, V, Spreen, G, Haas, C, et al. 2019. The 2018 North Greenland polynya observed by a newly introduced merged optical and passive microwave sea-ice concentration dataset. Cryosphere, 13: 2051–2073. DOI: 

  132. Marcott, SA, Fountain, AG, O’Connor, JE, et al. 2009. A latest Pleistocene and Holocene glacial history and paleoclimate reconstruction at Three Sisters and Broken Top volcanoes, Oregon, USA. Quaternary Research, 71(2): 181–189. DOI: 

  133. McCullough, KM. 1989. The Ruin Islanders: Thule culture pioneers in the Eastern High Arctic. (Mercury Series 141). Hull: Canadian Museum of Civilization. DOI: 

  134. McGhee, R. 1979. The Paleoeskimo occupations at Port Refuge, High Arctic Canada. (Mercury Series 92). Ottawa: National Museum of Canada. DOI: 

  135. McGhee, R. 1981. The Dorset occupations in the vicinity of Port Refuge, High Arctic Canada. (Mercury Series 105). Ottawa: National Museum of Canada. DOI: 

  136. McGhee, R. 1984. The Thule village at Brooman Point, High Arctic Canada. (Mercury Series 125). Ottawa: National Museum of Canada. DOI: 

  137. McGhee, R. 2000. Radio carbon dating and the timing of the Thule migration. In Appelt, M, Berglund, J and Gulløv, HC (eds.) Identities and Cultural Contacts in the Arctic: Proceedings from a Conference at the Danish National Museum. Copenhagen: Danish National Museum & Danish Polar Centre. 181–191. 

  138. Mathiassen, T. 1927. Archaeology of the Central Eskimos. Copenhagen: Gyldendal. 

  139. Møhl, U. 1979. Description and analysis of the bone material from Nugarasuk, an Eskimo settlement representative of the Thule culture in West Greenland. In McCartney, AP (ed.) Thule Eskimo Culture: an Anthropological Retrospective. (Mercury Series 88). Ottawa: National Museum of Canada. 380–394. DOI: 

  140. Moody, JF and Hodgetts, LM. 2013. Subsistence practices of pioneering Thule-Inuit: a faunal analysis of Tiktalik. Arctic Anthropology, 50(2): 4–24. DOI: 

  141. Moore, GWK, Schweiger, A, Zhang, J. et al. 2018. What caused the remarkable February 2018 North Greenland Polynya? Geophysical Research Letters, 45(24): 13, 342–13, 350. DOI: 

  142. Morales Maqueda, MA, Willmott, AJ and Biggs, NRT. 2004. Polynya dynamics: A review of observations and modeling. Review of Geophysics, 42(1): RG1004. DOI: 

  143. Morrison, DA. 2000. The arrival of the Inuit: Amundsen Gulf and the Thule Migration. In Identities and Cultural Contacts in the Arctic: Proceedings from a Conference at the Danish National Museum, Appelt, M, Berglund, J and Gulløv, HC (eds.). Copenhagen: Danish National Museum & Danish Polar Centre. pp 221–228. 

  144. Murray, MS. 2008. Zooarchaeology and arctic marine mammal biogeography, conservation and management. Ecological Applications, 18 (sp2): S41-S55. DOI: 

  145. Næss, RN. 2018. The Finnøy polar bear., 23 January. 

  146. Nash, RJ. 1976. Cultural systems and culture change in the central Arctic. Memoirs of the Society for American Archaeology, 31: 150–155. DOI: 

  147. National Parks Service (NPS). 2013a. Notice of intent to repatriate cultural items: U.S. Department of the Interior, Bureau of Land Management, Alaska State Office, Anchorage, AK. Federal Register, 78 (232): 72711–72712. 

  148. National Parks Service (NPS). 2013b. Notice of intent to repatriate cultural items: U.S. Department of the Interior, Bureau of Land Management, Alaska State Office, Anchorage, AK. Federal Register, 78 (232): 72710–72711. 

  149. Nazarova, L, Lüpfert, H, Subetto, D, et al. 2013. Holocene climate conditions in central Yakutia (eastern Siberia) inferred from sediment composition and fossil chironomids of Lake Temje. Quaternary International, 290–291: 264–274. DOI: 

  150. Nomokonova, T, Losey, RJ, Plekhanov, AV, et al. 2018. Iarte VI and Late Holocene reindeer remains from the Iamal Peninsula of Arctic Siberia. Arctic Anthropology, 55(2): 56–75. DOI: 

  151. Nordensheldt, AE. 1881. Travels of A.E. Nordensheldt around Europe and Asia on the ship ‘Vega’ in 1878–1880. Part 1, St. Petersburg. [Russian]. 

  152. Packard, AS. 1886. The former southern limits of the white or polar bear. American Naturalist, 20(7): 655–659. DOI: 

  153. Park, RW. 1989. Porden Point: An Intrasite Approach to Settlement System Analysis. Unpublished thesis (PhD), University of Alberta. 

  154. Pedersen, JBT, Kaufmann, LH, Kroon, A, et al. 2010. The Northeast Greenland Sirius Water Polynya dynamics and variability inferred from satellite imagery. Geografisk Tidsskrift-Danish Journal of Geography, 110(2): 131–142. DOI: 

  155. Pereverzev, AA and Kochnev, AA. 2012. Marine mammals in the Cape Schmidt vicinity (Chukotka) in September-October 2011. In Marine Mammals of the Holarctic, Papers of the Sixth International Conference, Moscow on 11–15 October 2010 (Vol. 2). pp176–181. 

  156. Petersen, Æ. 2010. Hvítabjarnakomur á Íslandi, einkum á Norðurlandi, ásamt almennum upplýsingum. In Þorsteinn Sæmundsson, Helgi P. Jónsson and Þórdís V. Bragadóttir (eds). Húnvetnsk náttúra 2010. Málþing um náttúru Húnavatnssýslna á Gauksmýri 10. apríl 2010. Náttúrustofa Norðurlands vestra, NNV-2010-003. Sauðárkrókur: Náttúrustofa Norðurlands vestra. 21–23 [Icelandic] Summarized at URL 

  157. Pitulko, VV. 1993. An early Holocene site in the Siberian high Arctic. Arctic Anthropology, 30(1): 13–21. 

  158. Pitulko, VV. 2003. The bear-hunters of Zhokhov Island, east Russian Arctic. Senri Ethnological Studies, 63: 141–152 

  159. Pitulko, VV and Kasparov, AV. 1996. Ancient Arctic hunters: material cultural and survival strategy. Arctic Anthropology, 33(1): 1–36. 

  160. Pitulko, VV and Kasparov, AV. 2017. Archaeological dogs from the Early Holocene Zhokhov site in the eastern Siberian Arctic. Journal of Archaeological Science: Reports, 13: 491–515. DOI: 

  161. Pitulko, VV, Ivanova, VV, Kasparov, A, et al. 2015. Reconstructing prey selection, hunting strategy and seasonality of the early Holocene frozen site in the Siberian High Arctic: A case study on the Zhokhov site faunal remains, De Long Islands. Environmental Archaeology, 20(2): 120–157. DOI: 

  162. Pitulko, VV, Kuzmin, YV, Glascock, MD, et al. 2019. ‘They came from the ends of the earth’: long-distance exchange of obsidian in the High Arctic during the Early Holocene. Antiquity, 93(367): 28–44. DOI: 

  163. Polyak, L, Alley, RB, Andrews, JT, et al. 2010. History of sea ice in the Arctic. Quaternary Science Reviews, 29(15–16): 1757–1778. DOI: 

  164. Post, K. 2005. A Weichselian marine mammal assemblage from the southern North Sea. Deinsea, 11: 21–27. 

  165. Rainey, FG. 1941. Eskimo Prehistory: The Okvik Site on the Punuk Islands. Anthropological Papers of the American Museum of Natural History, XXXVII (Part IV): 453–569. 

  166. Ramsay, MA and Stirling, I. 1988. Reproductive biology and ecology of female polar bears (Ursus maritimus). Journal of Zoology London, 214(4): 601–624. DOI: 

  167. Rankama, T. 2003. The colonization of Northernmost Finnish Lapland and the inland areas of Finnmark. In Larsson, L, et al. (eds.) Mesolithic on the Move. Oxford: Oxbow Books. 37–46. 

  168. Rasmussen, KL. 1996. Carbon-14 datings from northern East Greenland. In Grønnow, B and Pind, J (eds.) The paleo-eskimo cultures of Greenland. Copenhagen: Danish Polar Center. 188–189. 

  169. Rasmussen, TL and Thomsen, E. 2014. Brine formation in relation to climate changes and ice retreat during the last 15,000 years in Storfjorden, Svalbard, 76–78°N. Paleoceanography and Paleoclimatology, 29(10): 911–929. DOI: 

  170. Ray, CE. 1971. Polar bear and mammoth on the Pribilof Islands. Arctic, 24(1): 9–18. DOI: 

  171. Rick, AM. 1980. Non-cetacean vertebrate remains from two Thule winter houses on Somerset Island, N.W.T. Canadian Journal of Archaeology, 4: 99–117. 

  172. Rode, KD, Wilson, RR, Regehr, EV, et al. 2015. Increased land use by Chukchi Sea polar bears in relation to changing sea ice conditions. PLoS One, 10(11): e0142213. DOI: 

  173. Sabo III, G. 1981. Thule Culture Adaptations on the South Coast of Baffin Island, NWT. Unpublished thesis (PhD), Michigan State University. DOI: 

  174. Saleeby, B. 1994. Appendix II: Results of the faunal analysis. In Harritt, RK Eskimo prehistory on the Seward Peninsula. Unpublished Resources Report (NPS/ARO/RCR/CRR-93/21). Anchorage: U.S. National Park Service. 317–382 

  175. Sandell, HT and Sandell, B. 1991. Archaeology and environment in the Scoresby Sund fjord: Ethno-archaeological investigations of the last Thule culture of Northeast Greenland. (Meddelelser om Grønland/Man & Society 15). Copenhagen: Danish Polar Center. 

  176. Savelle, JM, Dyke, AS, Whitridge, PJ, et al. 2012. Paleoeskimo demography on western Victoria Island, Arctic Canada: Implications for social organization and longhouse development. Arctic, 65(2): 167–181. DOI: 

  177. Savinetsky, AB, Kiseleva, NK and Khassanov, BF. 2004. Dynamics of sea mammal and bird populations of the Bering Sea regionover the last several millennia. Palaeogeography, Palaeoclimatology and Palaeoecology, 209(1–4): 335–352. DOI: 

  178. Schledermann, P. 1975. Thule Eskimo prehistory of Cumberland Sound, Baffin Island, Canada. Mercury Series 38. Ottawa: National Museum of Man. 

  179. Schledermann, P. 1980. Polynyas and prehistoric settlement patterns. Arctic, 33(2): 292–302. DOI: 

  180. Schweiger, AJ, Steele, M, Zhang, J, et al. 2021. Accelerated sea ice loss in the Wandel Sea points to a change in the Arctic’s Last Ice Area. Communications Earth & Environment, 2: 122. DOI: 

  181. Smedsrud, LH, Budgell, WP, Jenkins, AD, et al. 2006. Fine-scale sea-ice modelling of the Storfjorden polynya, Svalbard. Annals of Glaciology, 44: 73–79. DOI: 

  182. Soon, W and Baliunas, S. 2003. Proxy climatic and environmental changes of the past 1000 years. Climate Research, 23: 89–110. DOI: 

  183. Sørensen, M and Gulløv, HC. 2012. The prehistory of Inuit in Northeast Greenland. Arctic Anthropology, 49(1): 88–104. DOI: 

  184. Sørensen, M, Gotfredsen, AB, Pedersen, JT, et al. 2009. GeoArk 2008. Archaeological and Zoo-archaeological Report on Investigations of the Southern Coast of Clavering Ø, the Revet area, Hvalros Ø and the Estuary of Young Sund. (Report 28). Copenhagen: National Museum of Denmark. 

  185. Speer, L, Nelson, R, Casier, R, et al. 2017. Natural Marine World Heritage in the Arctic Ocean, Report of an Expert Workshop and Review Process. Gland, Switzerland: IUCN. 

  186. Stanford, DJ. 1976. The Walakpa Site, Alaska: its place in the Birnirk and Thule cultures. Washington: Smithsonian Institution Press. DOI: 

  187. Stenton, DR. 1987. Recent archaeological investigations in Frobisher Bay, Baffin Island, NWT. Canadian Journal of Archaeology, 11: 13–48. 

  188. Stirling, I. 1974. Midsummer observations on the behavior of wild polar bears (Ursus maritimus). Canadian Journal of Zoology, 52(9): 1191–1198. DOI: 

  189. Stirling, I. 1980. The biological importance of polynyas in the Canadian Arctic. Arctic, 33(2): 303–315. DOI: 

  190. Stirling, I. 1997. The importance of polynyas, ice edges, and leads to marine mammals and birds. Journal of Marine Systems, 10(1–4): 9–21. DOI: 

  191. Stirling, I. 2002. Polar bears and seals in the eastern Beaufort Sea and Amundsen Gulf: a synthesis of population trends and ecological relationships over three decades. Arctic, 55(5) (Suppl. 1–93): 59–76. DOI: 

  192. Stirling, I and Cleator, HJ. (eds.). 1981. Polynyas in the Canadian Arctic. (Canadian Wildlife Service Occasional Paper 45). Ottawa: Environment Canada. 

  193. Stirling, I, Cleator, HJ and Smith, TG. 1981. Marine mammals. In Stirling, I and Cleator, HJ (eds.) Polynyas in the Canadian Arctic. (Canadian Wildlife Service Occasional Paper 45). Ottawa: Environment Canada. 45–58. 

  194. Stirling, I and Lunn, NJ. 1997. Environmental fluctuations in arctic marine ecosystems as reflected by variability in reproduction of polar bears and ringed seals. In Woodin, SJ and Marquiss, M (eds.) Ecology of Arctic Environments. Oxford: Blackwell Science. 167–181. 

  195. Stringer, WJ and Groves, JE. 1991. Location and areal extent of polynyas in the Bering and Chukchi Seas. Arctic, 44(5) (Suppl. 1–171): 164–171. DOI: 

  196. Stroeven, AP, Hättestrand, C, Kleman, J, et al. 2016. Deglaciation of Fennoscandia. Quaternary Science Reviews, 147: 91–121. DOI: 

  197. Swinarton, LE. 2008. Animals and the Precontact Inuit of Labrador: An Examination Using Faunal Remains, Space and Myth. Unpublished thesis (PhD), Memorial University. 

  198. Taldenkova, E, Bauch, HA, Stepanova, A, et al. 2008. Postglacial to Holocene history of the Laptev Sea continental margin: Paleoenvironmental implications of benthic assemblages. Quaternary International, 183(1): 40–60. DOI: 

  199. Taylor, WE, Jr.. 1972. An archaeological survey between Cape Parry & Cambridge Bay, NWT, Canada. (Mercury Series 1). Ottawa: National Museum of Man. 

  200. Taylor, WE Jr. and McGhee, R. 1979. Archaeological material from Creswell Bay, NWT, Canada. (Mercury Series 85). Ottawa: National Museum of Man. DOI: 

  201. Tamura, T and Ohshima, KI. 2011. Mapping of sea ice production in the Arctic coastal polynyas. Journal of Geophysical Research, 116(C7): C07030. DOI: 

  202. Tein, TS. 1977. The Investigations in Chyortov Ovrag (Devil’s Gorge) on Wrangell Island. In [no authors] Archaeological Discoveries of 1976. Moscow: Nauka. 246–247. [Russian]. 

  203. Tein, TS. 1978. The Investigations of Paleo-Eskimo Culture on Wrangell Island. In [no authors] Archaeological Discoveries of 1977. Moscow: Nauka. 278. [Russian]. 

  204. Thiemann, GW, Budge, SM, Iverson, SJ, et al. 2007. Unusual fatty acid biomarkers reveal age- and sex-specific foraging in polar bears (Ursus maritimus). Canadian Journal of Zoology, 85(4): 505–517. DOI: 

  205. Timokhov, LA. 1994. Regional characteristics of the Laptev and the East Siberian Seas: Climate, topography, ice phases, thermonhaline regime, circulation. In Kassens, H et al. (eds.) Russian-German Cooperation in the Siberian Shelf Seas: Geo-System Laptev Sea. Bremerhaven: Berichte zur Polarforschung 144. 15–31. 

  206. Vibe, C. 1950. The marine mammals and the marine fauna in the Thule District (Northwest Greenland) with observations on ice conditions in 1939–41. Copenhagen: Meddelelser om Grønland 150(6). 

  207. Vibe, C. 1967. Arctic animals in relation to climatic fluctuations. Copenhagen: Meddelelser om Grønland I70(5). 

  208. Valen, V, Mangerud, J, Larsen, E, et al. 1996. Sedimentology and stratigraphy in the cave Hamnsundhelleren, western Norway. Journal of Quaternary Science, 11(3): 185–201. DOI:<185::AID-JQS247>3.0.CO;2-Y 

  209. Vdovin, IS. 1977. Chukchi religious cults: Cultural monuments of the peoples of Siberia and the North (second half of the XIX- early XX centuries). Leningrad. [Russian]. 117–138. 

  210. Veltre, DW, Yesner, DR, Crossen, KJ, et al. 2008. Patterns of faunal extinction and paleoclimatic change from mid-Holocene mammoth and polar bear remains, Pribilof Islands, Alaska. Quaternary Research, 70: 40–50. DOI: 

  211. Vereshchagin, NK. 1969. The origin and evolution of the polar bear. In Bannikov, AG, Kishchinskii, AA, and Uspenskii, SM (eds.) The polar bear and its conservation in the Soviet Arctic. Leningrad: Hydrometeorological Publishing House. [Russian, English summary and figure captions]. 25–53. 

  212. Vincent, J-S. 1989. Quaternary geology of the northern Canadian Interior Plains. In Fulton, RJ (ed.) Quaternary Geology of Canada and Greenland. (Geology of Canada 1). Ottawa: Geological Survey of Canada. 100–137. DOI: 

  213. Voorhees, H, Sparks, R, Huntington, HP, et al. 2014. Traditional knowledge about polar bears (Ursus maritimus) in Northwestern Alaska. Arctic, 67(4): 523–536. DOI: 

  214. Wakefield, R. 2020. ‘The birthplace of Arctic ice’. Alfred-Wegener Institute, 20 November. URL 

  215. Winguth, C, Mickelson, DM, Larsen, E, et al. 2005. Thickness evolution of the Scandinavian Ice Sheet during the Late Weichselian in Nordfjord, western Norway: evidence from ice-flow modelling. Boreas, 34(2): 176–185. DOI: 

  216. Whitridge, PJ. 1992. Thule Subsistence and Optimal Diet: A Zooarchaeological Test of a Linear Programming Model Unpublished thesis (PhD), McGill University. 

  217. Woollett, JM. 2010. ‘Oakes Bay 1: A preliminary reconstruction of a Labrador Inuit seal hunting economy in the context of climate change.’ Geografisk Tidsskrift/Danish Journal of Geography, 110: 245259. DOI: 

  218. Woollett, JM, Henshaw, AS and Wake, CP. 2000. Palaeoecological implications of archaeological seal bone assemblages: case studies from Labrador and Baffin Island. Arctic, 53(4): 395–413. DOI: 

  219. Wyman, J. 1868. An account of some kjoekkenmoeddings, or shell-heaps, in Maine and Massachusetts. American Naturalist, 1: 561–584. DOI: