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The white-colored brown bears of the Southern Kurils

Yoshikazu Sato1,5, Hidetsugu Nakamura2, (Rounds 1987, Marshall and Ritland 2002). Inter- Yuka Ishifune3, and Noriyuki Ohtaishi4 and intra-population variation of pelage color and relationships with habitat, behavior, body size, and 1College of Bioresource Sciences, Nihon University, litter size have been studied for American black Fujisawa 252-0880, Japan bears (Jonkel and Cowan 1971; Rogers 1976, 1980; 2Graduate School of Bioresource Sciences, Nihon Rounds 1987; Klinka and Reimchen 2009). Howev- University, Fujisawa 252-0880, Japan er, only a few studies of the relationship between 3Graduate School of Veterinary Medicine, Hokkaido pelage type and ecological characteristics have been University, Sapporo 060-0818, Japan conducted for brown bears (Schwartz et al. 2003, 4 Hokkaido University Museum, Hokkaido University, Bochkin 2005). Sapporo 060-0810, Japan A white-colored brown bear lives in the Southern Kuril Islands. The earliest documentation of this Abstract: We investigated the distribution of brown white bear is its inclusion in the 1791 Japanese bears (Ursus arctos) with white pelage in Kunashiri painting titled Ininkari-zu (Fig. 1, Ohtsuka 2005), of and Etorofu Islands, Southern Kuril Islands; we here the Ainu chief Ininkari. We hereby name this white name this white pelage form the Ininkari bear. The pelage form of the brown bear Ininkari. The Ainu fur color of the brown bear varies considerably are an indigenous people of Japan. In the painting, throughout its range, and many pelage variations Chief Ininkari is seen leading 2 young bears. One of have been reported. Ininkari bears are unique in the bears has white fur with black eyes, nose, and having white fur only on the upper half of the body. claws; the other has dark brown fur. These bears There are no reports of bears with the Ininkari-type might be from the Kunashiri or Etorofu Islands, markings in other regions of the world. According to because Chief Ininkari is believed to have been literature and interview surveys, Ininkari bears have engaged in hunting and trade in the Southern Kuril been recognized since at least the late 1800s on Islands, as well as in eastern Hokkaido (Ohtaishi and Kunashiri and Etorofu. We surmise the reasons that Honma 2008, Taishi and Sato 2010). The painting is distribution of Ininkari bears is restricted to these one of a series of 12 portraits of Ainu chiefs, entitled islands are the lack of predators and the low hunting Ishu-retsuzo, painted by Hakyo Kakizaki (1764– pressure on brown bears there; these factors may 1826). allow the bears to maintain such a unique pelage. To ascertain the past and present status of the distribution of Ininkari bears, we surveyed the Key words: brown bear, coat color polymorphism, literature for records related to white-colored brown Etorofu, Ininkari, Ininkari bear, Kunashiri, Ursus bears. Moreover, we interviewed local people on the arctos, white pelage Southern Kuril Islands during 2008–09 and exam- Ursus 22(1):84–90 (2011) ined photographs to complement the literature survey. We confirm a white morph of brown bears of Kunashiri (Kunashir) and Etorofu (Iturup) Islands, Pelage color polymorphism is well known in the Southern Kuril Islands. We believe this is the first American black bear (Ursus americanus) and the report of brown bears with white coats on Kunashiri brown bear (U. arctos) (Garshelis 2009). Variation and Etorofu Islands, and probably from anywhere in and distribution of color type have been well the world. documented for these species (black bear: Rounds 1987, Larivie`re 2001; brown bear: LeFranc et al. 1987, Pasitschniak-Arts 1993, Schwartz et al. 2003). Methods However, observations of frequent color variations We surveyed the literature beginning with the among cubs in the same litter render color phases Meiji Era (1868–1912) for descriptions of Ininkari somewhat unreliable as criteria for classification bears in the Southern Kuril Islands and in Japan, using mainly the Digital Library of the National [email protected] Diet Library, Japan (http://kindai.ndl.go.jp/index.

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mappu, Kunashiri, and included in the survey only those photographs in which the location and the date the photograph was taken were available. We also conducted camera-trapping to document Ininkari bears. Five camera traps with an infrared triggering system (Sensor Camera FieldNote DCs700, Marif Co. Ltd., Yamaguchi, Japan) were set up at a total of 8 camera stations near the animal trails along the Onnebetsu River in Kunashiri Island for 6 days during 22–27 October 2009 (total of 23 camera-nights).

Results From the literature survey, we found several references that described white-colored brown bears in Japan in addition to the Ininkari-zu painting. Komiya (2008) reported that a white-colored brown bear born on Etorofu Island had been reared in the Ueno Zoo, Tokyo, during 1878–81. Although we found other references to white brown bears reared in the Ueno Zoo, we could neither authenticate the birth place for the bears nor rule out the possibility that those bears were albinos. In his notes on the numbers of hunted animals and the prices of animal Fig. 1. Ininkari-zu by Hakyo Kakizaki; this 1791 products in 1890 in the Chishima (Kuril) Islands, painting is the earliest documentation of the white Ohtashiro (1893), a resident of Etorofu Island, bear. Original drawing is owned by the Museum of recorded the color of Etorofu brown bears as golden, Fine Arts and Archeology, Besanc¸ on, France. This black, and white. Tarao (1893), a Japanese explorer, photo of the painting was provided by the National Museum of Ethnology, Japan. in a report of an exploration in the Chishima (Kuril) Islands during 1891–1892, also recorded the presence of white-colored brown bears on Etorofu Island html). We used only articles in which a location and along with black and golden-colored bears. Other a date of the record for a white bear were explicitly sources include mention of white bears, but because mentioned or appropriately estimated. We also the origin of those bears was unknown, these examined articles about coat color variation in references were not included in our survey. Our brown bears throughout the world with special literature search results confirmed that Ininkari reference to white color. bears existed by no later than the late 1800s, and During our preliminary study in the Seoi and probably as early as the late 1700s, based on the Onnebetsu River area of Kunashiri Island, 2008–09, depiction in Ininkari-zu. The literature survey yielded we conducted questionnaire surveys on Ininkari no clear evidence of the existence of Ininkari bears bears through interviews with local hunters, the staff on Kunashiri Island. of Kurilsky State Nature Reserve, and amateur We collected reliable information from 7 local naturalists, as well as at meetings about bears in people (A–E), including hunters in Kunashiri (A, B), Furukamappu, Kunashiri (Fig. 2). We used only staff members of the nature reserve of Kunashiri information from witnesses who directly observed Island (C, D, E), a former staff member of the nature the bears. reserve on Etorofu Island (F), and an amateur We collected photographs of Ininkari bears taken naturalist on Kunashiri Island (G). by staff members of the nature reserve, photogra- Local hunter A estimated that Ininkari bears phers, and amateur naturalists in Kunashiri. We made up approximately 10% of the total brown bear evaluated all photographs in 2008–09 in Furuka- population on Kunashiri Island based on his

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Fig. 2. Map of the Kunashiri and Etorofu Islands in the Southern Kuril Islands and locations where white- colored brown bears were recorded on Kunashiri Island (shot:m, photo: X). experience observing and hunting bears. Three of 30 white bear sightings had increased recently. Ininkari bears that he had shot were Ininkari bears: a bears were observed more often in the northern part subadult male in May 2001 between the Chikuni of Kunashiri Island, where the bear population River and Okitsuus, an adult male in October 2003 density is higher than in the southern part (infor- at the Ruyobetsu River, and an adult male bear in mants A, B, and C). The former staff member of the October 2007 at the Chikappunai River. In the last nature reserve on Etorofu Island (F) observed instance, we confirmed the white fur of the brown Ininkari bears in Etorofu. bear hide. Our examination of photographs taken since 1990 The second hunter (B) and a staff member of the in Kunashiri and Etorofu Islands by 8 local people nature reserve (C) pointed out that the number of revealed that there were various patterns of colora-

Ursus 22(1):84–90 (2011) SHORT COMMUNICATION N Sato et al. 87 tion among the bears from dark brown to white. The local photographers included 5 people who we contacted directly and 3 whose photographs were owned by a staff member at the nature reserve. Most of the bears in the photographs have dark brown fur with gold tips on the guard hairs on their heads, necks, and backs. This pattern is similar to that of typical brown bears living on Hokkaido (Sato 2009). Even so, the area of the golden tips varies among individual bears. Our interviews revealed some disparity about the usage of the term ‘‘white bear.’’ Some local people in Kunashiri Island seem to regard a bear with broad swaths of gold-tipped pelage on the head, neck, and back as a ‘white bear.’ However, as some authors have pointed out (Swenson et al. 2000, Schwartz et al. 2003, Garshelis 2009), pelage color may differ according to the level of sunshine, as well as the direction from which the observer sees the bear. Therefore, we excluded any suspect descriptions from the study. Fortunately, we obtained clear photographs of Ininkari bears from 4 people on both Islands (Fig. 3; Anatolii 2008); in these photos, the difference between normal brown bears and Ininkari bears was clear. Even when the bear was photographed with the sun behind, the Ininkari bear with its wide bands of white fur was quite distinctive. Respondents A, B, C, and G pointed out the Fig. 3. White-colored (Ininkari) brown bears in characteristic coloration in the Ininkari bears: white Kunashiri (Kunashir) Island, in the Southern Kuril fur covers only the upper half of the body (head, Islands. Photos by T. Shpilenok (top, 2005) and E. shoulders, and forelegs), and dark brown fur covers Grigoriev (bottom, 2004). the lower half thinly (Fig. 3). Unlike brown bears whose fur on the face and back may change with age, neck and forelegs, inhabit both Kunashiri and Ininkari bears have white fur from birth (A). A total Etorofu Islands. of 14 brown bear photographs were taken by camera-traps in 2009, and an Ininkari bear was confirmed in one of these (unpublished data). Discussion Although our interviews revealed observations of The coat color of brown bears varies considerably both young and adult Ininkari bears, there was no throughout their range, from pale tan, blond, mention of sightings of adult Ininkari bears accom- golden, silver, cinnamon, and all shades of brown panied by young. Respondents A, B, C, and G to almost black (LeFranc et al. 1987, Pasitschniak- reported having observed an adult dark brown bear Arts 1993, Vaisfield and Chestin 1993, Schwartz et accompanied by both a young dark brown bear and al. 2003, Garshelis 2009, Sato 2009). Color morph a young Ininkari bear, but did not observe any adult variation seems to be greater in North America and Ininkari bears with young bears. Likewise, the Asia than in Europe (Swenson et al. 2000, Garshelis former staff member of the nature reserve on 2009). Some individuals may appear light or dark Etorofu (F) also observed an adult dark brown bear from different angles due to variegated guard hairs with a young Ininkari bear on Etorofu Island. (Swenson et al. 2000, Schwartz et al. 2003, Garshelis Hunter A said that all 3 of the Ininkari bears he 2009). Furthermore, the coat of some individuals shot were male. From the interviews, we confirmed may change from a lighter brown to a darker brown that Ininkari bears, with their unique white head, as they age, or coat color may change to silver

Ursus 22(1):84–90 (2011) 88 SHORT COMMUNICATION N Sato et al. because of inadequate nutrition (Rogers 1980, difference in coat color is caused by a single recessive Bochkin 2005). Additionally, the coat color of allele (Ritland et al. 2001). However, ecological or individuals may be lightened by the sun during the evolutionary explanations for the restricted distribu- animals’ active season (Bochkin 2005, Garshelis tion and persistence of the white bear have not yet 2009). Generally, though variation in color is high, been fully developed. Marshall and Ritland (2002) and the head and shoulders are paler than the sides, suggested that white-phase black bears are main- belly, and legs (LeFranc et al. 1987, Pasitchniak-Arts tained in populations by a combination of genetic 1993, Schwartz et al. 2003). isolation and small population sizes caused by an Some brown bear populations in Asia and cubs in insular habitat, with the possible contribution of their first year in North America have prominent selective pressure, nonrandom mating, or both. On white markings that extend from the chest over the the other hand, Klinka and Reimchen (2009) pointed shoulders and sometimes connect at the back out the direct fitness benefits of white fur color as an (Schwartz et al. 2003, Garshelis 2009, Sato 2009). alternative or a supplemental mechanism for the Another brown bear subspecies, Tibetan blue bears origin and persistence of the polymorphism, because (U. arctos pruinosus), have a light colored coat, a white morph is more efficient at capturing salmon called blue or white, over their torso with dark legs, (Oncorhynchus spp.) during daylight than in the face, and hump (Schaller 1998, Garshelis 2009). dark. Possibly, there are additional costs associated Additionally, some people refer to grizzly bears (U. with the white phase, such as increased risk of arctos horribilis) as ‘white bears’ because of the sheen predation. Klinka and Reimchen (2009) suggested of their white-tipped guard hairs (Garshelis 2009). that the absence of brown bears and low frequency Despite these examples of ‘white’ brown bears, we of wolves (Canis lupus) would influence the presence were unable to find any reference to bears with a of the white morph bear only in these islands. white coat on their head, shoulders, and forelegs In the Kunashiri and Etorofu Islands, bears fed on except for the Ininkari bears of Kunashiri and early vegetation, berries, nuts of the previous years, Etorofu Islands that we report in this study. and invertebrates after leaving the den (Yudin 1993). Although the white bear in the painting in Ininkari- Because most of their diet in spring and summer was zu (Fig. 1) looks different from the white-colored plant material, the white pelage of Ininkari bears bears in the wild (Fig. 3), especially for the would not provide an advantage. However, pink and coloration of the posterior part, we could not chum salmon (Oncorhynchus gorbuscha and O. keta) explain whether the difference was caused by an are known to be their major food items in autumn artistic license in the painting, or some other reason (Yudin 1993). White pelage may provide an advan- such as changes with age. Additional observation of tage for catching salmon as Klinka and Reimchen white-colored bears in various cohorts would con- (2009) have reported for the white morph of the tribute to an understanding of the difference. American black bear. Variation in coloration among and within species Although further study of genetic and behavioral is known for mammals, but its function is not yet differences between Ininkari bears and other brown understood (Caro 2005). In addition to the brown bears in Kunashiri and Etorofu Islands is clearly bear, the American black bear presents one of the needed, we may infer the costs of having white most striking examples of variation in color among pelage. White fur seems to be a disadvantage because wild mammals (Rounds 1987). Several authors have it may increase visibility to predators. However, suggested that there are relationships between color there have been no wolves (which might prey on and habitat, weight, and litter size (Jonkel and young brown bears) on either island for at least Cowan 1971; Rogers 1976, 1980; Rounds 1987). 100 years (Inukai 1934). Possibly wolves are now Among these factors, habitat is considered the extinct on the islands, although there exist a few dominant influence in color phenotypes in bears records of their fur in both islands from more than (Rounds 1987). 100 years ago (Schrenk 1862, Kishida 1960). A white morph of American black bears (U. Cannibalism within bears was not found in the americanus kermodei) occurs on the Kermode Kunashiri and Etorofu Islands by Yudin (1993). Islands off the Pacific coast of British Columbia, Indigenous Ainu people conserved brown bears, and Canada, where they co-exist with the more common hunting by Ainu seems not to have been a major black morph (Hornaday 1905, Larivie`re 2001). The cause of mortality among bears on these islands

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(Mano 2006). In our interviews, local Russian JONKEL, C.J., AND I.M. COWAN. 1971. The black bear in the hunters (A and B) said that the number of active spruce–fir forest. Wildlife Monograph 27. bear hunters in Kunashiri in recent years was low, KISHIDA, K. 1960. Mammalian fauna of Kuril Islands. and they reported that they had never selectively (1932). Japan Wildlife Bulletin 17:283–306. (In Japanese.) killed white bears while hunting. Consequently, we KLINKA, D.R., AND T.E. REIMCHEN. 2009. Adaptive coat colour polymorphism in the Kermode bear of coastal suspect that one reason for the restricted distribution British Columbia. Biological Journal of the Linnean and persistence of Ininkari bears is a lack of Society 98:479–488. predators and low hunting pressure, a view consis- KOMIYA, T. 2008. History of raising bears in Ueno Zoo. tent with the example of the white morph of the Pages 227–254 in S. Matsunaga, editor. Environment. American black bear. Culture and Policy. Toshindo, Tokyo, Japan. (In Additional study of genetic and behavioral differ- Japanese.) ences between Ininkari and normal brown bears LARIVIE` RE, S. 2001. Ursus americanus. Mammalian Species would contribute to an understanding of the 647:1–11. persistence of polymorphisms in the coat color in LEFRANC, M.N., JR., M.B. MOSS, K.A. PATNODE, AND large vertebrates and to conservation of the bear W.C. SUGG III. 1987. Grizzly bear compendium. population in Kunashiri and Etorofu Islands. Interagency Grizzly Bear Committee, Washington, DC, USA. MANO, T. 2006. The status of brown bears in Japan. Pages 111–121 in Japan Bear Network, editor. Under- Acknowledgments standing Asian bears to secure their future. Japan Bear We thank the staff of the Ministry of Environment Network, Ibaraki, Japan. (MOE) and the Ministry of Foreign Affairs (MOFA) MARSHALL, H.D., AND K. RITLAND. 2002. Genetic diversity of Japan for arranging our field study in Kunashiri and differentiation of Kermode bear populations. and Etorofu Islands. We also are grateful to I. Molecular Ecology 11:685–697. Nevedomskaya, A. Loguntzev, and other staff of the OHTAISHI, N., AND H. HONMA. 2008. Shiretoko and the Kurilsky State Nature Reserve for their collabora- Four Northern Islands. A world natural heritage tive research in the field. As well, we thank S.D. natured by floating ice. Iwanami Shoten, Tokyo, Japan. Ohdachi and A. Kameyama for their valuable (In Japanese.) comments on an earlier draft of this paper. This OHTASHIRO, J. 1893. A geographical description of industry research was supported in part by a Grant-in-Aid for in Chishima. Komeikan, Tokyo, Japan. (In Japanese.) OHTSUKA, K. 2005. Keynote lecture: Objectives of the Scientific Research 21405009 from the Japan Society symposium. Pages 3–5 in Abstract of the symposium for the Promotion of Science and by the Mitsui & ‘‘Close reading of Ishu-retsuzo’’ in memory of retire- Company, Ltd. Environment Fund (2007 and 2009). ment for Professor Kazuyoshi Otsuka. National Museum of Ethnology, 19–20 March 2005, Osaka, Japan. (In Japanese.) Literature cited PASITSCHNIAK-ARTS, M. 1993. Ursus arctos. Mammalian ANATOLII, S. 2008. Kuril. Fascinated islands. Polikor, Species 439:1–10. Krasnojorsk, Russia. (In Russian.) RITLAND, K., C. NEWTON, AND H.D. MARSHALL. 2001. BOCHKIN, E. 2005. Bear of Kolyma region. OxoTa, Inheritance and population structure of the white- Hunting 9:14–17. (In Russian.) phased ‘‘Kermode’’ black bear. Current Biology CARO, T. 2005. The adaptive significance of coloration in 11:1468–1472. mammals. Bioscience 55:125–136. ROGERS, L.L. 1976. Effects of mast and berry crop failures GARSHELIS, D. 2009. Family Ursidae. Pages 448–497 in on survival, growth, and reproductive success for black D.E. Wilson and R.A. Mittermeier, editors. Handbook bears. Proceedings of North American Wildlife and of mammals of the world. Volume 1. Carnivora. Lynx Natural Resource Conference 41:431–438. Editions, Barcelona, Spain. ———. 1980. Inheritance of coat color and changes in HORNADAY, W.T. 1905. A new white bear, from British pelage coloration in black bears in northeastern Columbia. Annual Report of New York Zoological Minnesota. Journal of Mammalogy 61:324–327. Society 9:81–86. ROUNDS, R.C. 1987. Distribution and analysis of colour INUKAI, T. 1934. A general survey of fauna in Chishima. morphs of the black bear (Ursus americanus). Journal Pages 87–89 in Hokkaido Government, editor. Chishima of Biogeography 14:521–538. Gaikyo. An outlook of Chishima. Hokkaido Govern- SATO, Y. 2009. Ursus arctos Linnaeus, 1758. Pages 232–234 ment, Sapporo, Japan. (In Japanese.) in S.D. Ohdachi, Y. Ishibashi, M.A. Iwasa, and T.

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