Brown bear

From Wikipedia, the free encyclopedia
Jump to: navigation, search
This article is about the animal. For the athletics teams at Brown University, see Brown Bears.
Brown bear
Temporal range: 0.5–0 Ma
Middle Pleistocene-Holocene
Brown bear.jpg
Brown bear (U. arctos) in Hallo Bay, Katmai National Park, Alaska
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Ursidae
Genus: Ursus
Species: U. arctos
Binomial name
Ursus arctos
Linnaeus, 1758
Subspecies

16, see text

Ursus arctos range map.svg
Brown bear range map

The brown bear (Ursus arctos) is a large bear distributed across much of northern Eurasia and North America and is the largest terrestrial carnivoran.[2] There are several recognized subspecies within the brown bear species.

While the brown bear's range has shrunk and it has faced local extinctions, it remains listed as a least concern species by the IUCN with a total population of approximately 200,000. As of 2012, this and the American black bear are the only bear species not classified as threatened by the IUCN. However, the Californian, North African (Atlas bear), and Mexican subspecies, as well as brown bear populations in the Pacific Northwest were hunted to extinction in the nineteenth and early twentieth centuries, and many of the southern Asian subspecies are highly endangered. The smallest subspecies, the Himalayan brown bear, is critically endangered, occupying only 2% of its former range and threatened by uncontrolled poaching for its parts.[3] The Marsican brown bear in central Italy is believed to have a population of just 30 to 40 bears.

The brown bear's principal range includes parts of Russia, Central Asia, China, Canada, the United States (mostly Alaska), Scandinavia and the Carpathian region (especially Romania), Anatolia, and Caucasus.[1][4] The brown bear is recognized as a national and state animal in several European countries. It is the most widely distributed of all bears.

Evolution and taxonomy[edit]

The brown bear is sometimes referred to as the bruin, from Middle English. This name originated in the fable, History of Reynard the Fox, translated by William Caxton, from Middle Dutch bruun or bruyn, meaning brown (the color).[5] In the mid-19th century United States, the brown bear was termed "Old Ephraim" and sometimes as "Moccasin Joe".[6] The scientific name of the brown bear, Ursus arctos, comes from the Latin "ursus", meaning "bear",[7] and Άρκτος "arctos", from the Greek word for bear.[8]

Generalized brown bear names and evolution[edit]

Brown bears are thought to have evolved from Ursus etruscus in Asia.[9] The brown bear Kurten believed to be "clearly derived from the Asian population of Ursus savini about 800,000 years ago; spread into Europe, to the New World."[10] The oldest fossils occur in China from about 0.5 million years ago.[9] They entered Europe about 250,000 years ago, and North Africa shortly after.[9] Brown bear remains from the Pleistocene period are common in the British Isles, where it is thought they might have outcompeted cave bears.[9] The species entered Alaska 100,000 years ago, though they did not move south until 13,000 years ago.[9] It is speculated that brown bears were unable to migrate south until the extinction of the much larger Arctodus simus.[11]

Several paleontologists suggest the possibility of two separate brown bear migrations: inland brown bears, also known as grizzlies, are thought to stem from narrow-skulled bears which migrated from northern Siberia to central Alaska and the rest of the continent, while Kodiak bears descend from broad-skulled bears from Kamchatka, which colonized the Alaskan peninsula.[9] Brown bear fossils discovered in Ontario, Ohio, Kentucky and Labrador show the species occurred farther east than indicated in historic records.[9]

Scientific brown bear taxonomy[edit]

There are many methods used by scientists to define bear species and subspecies as no one method is always effective. Genetic testing is now perhaps the most important way to scientifically define brown bear relationships and names. Generally genetic testing uses the word clade rather than species because a genetic test alone cannot define a biological species. Most genetic studies report on how closely related the bears are (or their Genetic distance). There are hundreds of obsolete brown bear subspecies, each with its own name, and this can become confusing; Hall (1981) lists 86 different types.[12] The exact number of overall brown subspecies and its precise relationship to the polar bear remains in debate. The polar bear is a recent offshoot of the brown bear, having diverged approximately 400,000 years ago.[13]

In North America, two ecotypes of the single species Ursus arctos horribilis are generally recognized—the coastal brown bear and the inland grizzly; these two types broadly define the range of sizes of all brown bear subspecies.

Genetic clades[edit]

"The genetic diversity of present-day brown bears (Ursus arctos) has been extensively studied over the years and appears to be geographically structured into five main clades."[14]

Brown bear names[edit]

There is little agreement on the classification of brown bears because there are so many ways to name and group them.

Some systems have proposed as many as 90 subspecies,[15] while recent DNA analysis has identified as few as five clades.[16]

There are several recognized subspecies within the brown bear species. In North America, two types of the subspecies Ursus arctos horribilis are generally recognized—the coastal brown bear and the inland grizzly bear; these two types broadly define the range of sizes of all brown bear subspecies. An adult grizzly living inland in Yukon may weigh as little as 80 kg (180 lb), while an adult coastal brown bear in nearby coastal Alaska living on a steady, nutritious diet of spawning salmon may weigh as much as 680 kg (1,500 lb).[17] The exact number of overall brown subspecies remains in debate.

DNA analysis recently revealed that the brown bears in North America are genetically quite homogeneous except for the one subspecies Kodiak bear and the enigmatic ABC Islands Bears that have the mtDNA of the polar bear (Ursus maritimus).[18] As of 2005, 16 living global subspecies have been recognized.[19][20] The subspecies have been listed as follows:

Eurasian[edit]

Subspecies name Image Distribution Description/Comments
Ursus arctos arctosEurasian brown bear European Brown Bear.jpg Europe, Caucasus, Siberia (except the east) and Mongolia[21] A predominantly dark colored (rarely light colored), moderately sized subspecies with dark claws, the Eurasian browns occurring in Siberia are larger than their European counterparts, as they are hunted less.[21]
Ursus arctos beringianusKamchatka brown bear (or Far Eastern brown bear) Brown-bear-in-spring.jpg Shantar Islands, Kolyma Peninsula, Kamchatka Peninsula and Paramushir Island[21] A very large, dark colored bear. It is thought to be the ancestor of the polar bear and the Kodiak bear.[9] and the Peninsular brown bears of Alaska. Middendorf described it from Greater Shantar Island with its distribution range comprising the eastern coast of Siberia and Japan.[22]
Ursus arctos collarisEast Siberian brown bear Ucollaris.jpg East Siberia from the Yenisei River to the Altai Mountains, also found in northern Mongolia A predominantly dark form, it is intermediate in size between U. a. arctos and U. a. beringianus, with a proportionately larger skull.[21]
Ursus arctos crowtheri – †Atlas bear (extinct) Atlasbear.jpg Habitat while still extant was the Atlas Mountains and adjacent areas in North Africa, from Morocco to Libya. Last surviving bear is thought to have been killed by hunters in 1890.[23]
Ursus arctos isabellinusHimalayan brown bear Medvěd plavý (Ursus arctos isabellinus).jpg Northern Nepal, Northern and Eastern India, and Northern Pakistan Has a reddish-brown or sandy coat color and large ears, this bear is smaller than most other brown bears found on the Asian continent. Prefers high altitude forest and alpine meadow. Critically Endangered.
Ursus arctos marsicanusMarsican brown bear or Apennine brown bear[24] Marsica, central Italy There are an estimated 30 to 40 bears remaining in the Marsica area. This is an unrecognized subspecies that is considered to be a member of the nominate subspecies.[19]
Ursus arctos pruinosusTibetan blue bear Tibetan Blue Bear - Ursus arctos pruinosus - Joseph Smit crop.jpg Tibetan plateau [21] an isolated sub-population lives in the Gobi Desert. This is a moderately sized subspecies with long and shaggy fur. Both dark and light variants are encountered, with intermediate colors predominating. The fur around the neck is light, and forms a "collar". The skull is distinguished its relatively flattened choanae, an arch-like curve of the molar row and large teeth.[21]
Ursus arctos lasiotusUssuri brown bear (or Amur brown bear, black grizzly or horse bear) Ursus arctos lasiotus - Beijing Zoo 3.JPG Russia: Southern Kuril Islands, Sakhalin, Maritime territory, and the Ussuri/Amur River region south of the Stanovoy Range, China (former Manchuria): Heilongjiang, Japan: Hokkaidō, Honshu (last glacial period), Korean Peninsula: North Korea[21] Became extinct on Rebun and Rishiri Islands in 13th century.[25] Ursus arctos lasiotus in Hokkaido is one of the smallest form of brown bears, however, individuals growing larger and reaching 400 kg or up to 550 kg[26] in weight by feeding on cultivations.[27]

This bear is thought to be the ancestor of U. a. horribilis.[9]

Ursus arctos syriacusSyrian brown bear Ursus arctos syriacus.jpg The trans-Caucasus, Syria, Iraq, Turkey (Asia Minor), Iran, Afghanistan, Pakistan, western Himalayas and the Pamir-Alai and Tien Shan mountains,[21] probable historical presence in Israel The Syrian is a light colored, moderate to small-sized subspecies with light claws.[21]

North American[edit]

Subspecies name Image Distribution Description/Comments
Ursus arctos alascensis[28][29] – Alaska brown bear Coastal Alaska
Ursus arctos californicus – †California grizzly (extinct) Monarch the bear.jpg California The last known bear was shot in California in 1922.
Ursus arctos dalliDall Island brown bear Dall Island
Ursus arctos gyasPeninsular brown bear Brown bear.jpg Alaska Peninsula Considered by some biologists to be same subspecies as U. a. middendorffi.[30]
Ursus arctos horribilisGrizzly bear Grizzly Bear Yellowstone.jpg Northern and Western Canada, Alaska, and the northwestern United States, historically existed in Great Plains Grizzlies are identified by a medium to dark brown coat with gray or blond "grizzled" tips on the fur. Smaller than coastal bears, a large male grizzly can weigh up to 364 kilograms (802 lb) in inland areas, with bears in the Yukon region weighing as little as 80 kg (180 lb)[citation needed]. Coastal bears may be nearly twice a mountain grizzly's weight. Highly adaptable: it can live in montane pine forests, temperate rainforest, semi-arid scrubland, and shortgrass prairie.
Ursus arctos middendorffiKodiak bear Bear Square.JPG Kodiak, Afognak, Shuyak Islands (Alaska) This is the largest subspecies of brown bear, with other coastal brown bears potentially reaching nearly as large.
Ursus arctos nelsoni – †Mexican grizzly bear (extinct) Mexico grizzlies.png The smallest North American bear, formerly from northern Mexico, including Chihuahua, Coahuila, and Sonora, southwestern United States including southern ranges of Texas, Arizona, and New Mexico This bear is believed extinct due to its interference with cattle ranching in both the United States and Mexico. Distinct in its ability to survive arid conditions, it could live in both montane pine forests of Mexico and canyonlands of adjacent to the Sonoran Desert.
Ursus arctos sitkensis Admiralty Island, Baranof Island, and Chichagof Island the ABC Islands of Alaska. Appearing to be more closely related to the polar bear than to other brown bears,[18] this species is called "clade II" by Waits, and others, and is part of the former subspecies identified as U. a. sitkensis, by Hall and as U. a. dalli by Kurtén.
Ursus arctos stikeenensisStikine brown bear Northwestern British Columbia[31] From the Stikine River to the Skeena River.[32] Considered by biologists to be U. a. horribilis,[18][33] it once was thought a subspecies.[31]

Hybrids[edit]

A grizzly–polar bear hybrid (known as a pizzly bear or grolar bear) is a rare ursid hybrid resulting from a union of a brown bear and a polar bear. It has occurred both in captivity and in the wild. In 2006, the occurrence of this hybrid in nature was confirmed by testing the DNA of a strange-looking bear that had been shot in the Canadian arctic.[34][35][36] Previously, the hybrid had been produced in zoos, and was considered a "cryptid" (a hypothesized animal for which there is no scientific proof of existence in the wild).

Because they can interbreed these bears are not different biological species.[dubious ]

Analyses of the genomes of bears have recovered multiple instances of introgressive hybridization between various bear species,[37][38] including introgression of polar bear DNA intro brown bears during the Pleistocene.

Formerly considered subspecies[edit]

Former
Subspecies Name
Image Distribution Description
Ursus arctos pyrenaicusIberian brown bear, sometimes called Cantabrian brown bear, now considered Ursus arctos arctosEuropean brown bear See photographs in Eroski article (in Spanish, also available in Catalan, Basque and Galician) and in Fauna Ibérica. Oso pardo ibérico (Ursus arctos pyrenaicus), in Spanish Iberian Peninsula, primarily the Cantabrian Mountains and hills in Galicia, and the Pyrenees[39] Until recently, this bear was considered a separate subspecies. Today, it is considered to belong to the U. arctos arctos subspecies. Scientific evidence based on DNA studies would furthermore indicate the European brown bear can be divided into two distinct lineages. "There is a clear division into two main mitochondrial lineages in modern European brown bear populations. These populations are divided into those carrying an eastern lineage (clade IIIa, Leonard et al. 2000), which is composed of Russian, northern Scandinavian and eastern European populations, and those carrying a western lineage (clade I, Leonard et al. 2000), which is composed of two subgroups, one believed to originate from the Iberian Peninsula, including southern Scandinavian bears and the Pyreneean populations; and the other from the Italian–Balkan peninsulas (Taberlet et al. 1994; see however Kohn et al. 1995). In addition, based on the subfossil record in northwestern Moldova and mitochondrial DNA data from modern populations, a Carpathian refuge has also been proposed (Sommer & Benecke 2005; Saarma et al. 2007)."[40]

The brown bear is the largest wild animal on the Iberian Peninsula, although one of the smallest of the brown bears, weigh between 130 and 180 kg (290 and 400 lb) as adults. Their fur varies from a pale cream color to dark brown, but always with a distinctively darker, nearly black tone at the paws and a yellowish tinge at the tip of each hair. The brown bear population is considered endangered in Spain.

The brown bear population in the Pyrenees stems mostly from bears reintroduced from Slovenia, with one or two remaining original males.[39][40][41][42][43]

Description[edit]

Brown bear claws are longer and less curved than those of black bears
Brown bear skull

Color of brown bears[edit]

Brown bears are often not brown. Brown bears have long, thick fur, with a moderately long mane at the back of the neck. In India, brown bears can be reddish with silver tips, while in China, brown bears are bicolored with a yellow-brown or whitish cape across the shoulders. North American grizzlies can be dark brown (almost black) to cream (almost white) or yellowish brown. Black hairs usually have white tips.[23] The winter fur is very thick and long, especially in northern subspecies, and can reach 11 to 12 centimetres (4 to 5 in) at the withers. The winter hairs are thin, yet rough to the touch. The summer fur is much shorter and sparser, and its length and density varies geographically.[21]

Large claws for digging[edit]

Brown bears have very large and curved claws, those present on the forelimbs being longer than those on the hind limbs. They may reach 5 to 6 centimetres (2.0 to 2.4 in) and sometimes 7 to 10 centimetres (2.8 to 3.9 in) along the curve.[44] They are generally dark with a light tip, with some forms having completely light claws.[21] Brown bear claws are longer and straighter than those of American black bears (Ursus americanus).[23] The claws are blunt, while those of a black bear are sharp. Due to their claw structure, in addition to their excessive weight, adult brown bears cannot climb trees as can both species of black bear. The paws of the brown bear are quite large. The rear feet of adult bears have been found to typically measure 21 to 36 cm (8.3 to 14.2 in) long, with huge Kodiak bears having measured up to 46 cm (18 in) along their rear foot.[45][46]

Massive brown bear skulls and jaws[edit]

Adults have massive, heavily built concave skulls, which are large in proportion to the body. The forehead is high and rises steeply.[23] The projections of the skull are well developed when compared to those of Asian black bears (Ursus thibetanus): the latter have sagittal crests not exceeding more than 19–20% of the total length of the skull, while the former have sagittal crests comprising up to 40–41% of the skull's length. Skull projections are more weakly developed in females than in males. The braincase is relatively small and elongated. There is a great deal of geographical variation in the skull, and presents itself chiefly in dimensions.[21] Grizzlies, for example, tend to have flatter profiles than European and coastal American brown bears.[47] Skull lengths of Russian bears tend to be 31.5 to 45.5 centimetres (12.4 to 17.9 in) for males, and 27.5 to 39.7 centimetres (10.8 to 15.6 in) for females. The width of the zygomatic arches in males is 17.5 to 27.7 centimetres (6.9 to 11 in), and 14.7 to 24.7 centimetres (5.8 to 9.7 in) in females.[21] Brown bears have very strong teeth: the incisors are relatively big and the canine teeth are large, the lower ones being strongly curved. The first three molars of the upper jaw are underdeveloped and single crowned with one root. The second upper molar is smaller than the others, and is usually absent in adults. It is usually lost at an early age, leaving no trace of the alveolus in the jaw. The first three molars of the lower jaw are very weak, and are often lost at an early age.

Body size[edit]

The size of brown bears is the most variable of modern bears. The typical size depends upon which population it is from. A large barren ground brown bear might seem quite small elsewhere.

The brown bear (grizzly bear in Europe, Yellowstone National Park or interior Alaska) seasonally weigh on average between 115 and 360 kg (254 and 794 lb). Bears from the Yukon Delta, interior British Columbia, Jasper National Park and southern Europe can weigh from 55 to 155 kg (121 to 342 lb) on average.[17][48] Bears from the Syrian (U. a. syriacus) and the Gobi Desert (U. a. gobiensis) subspecies are around the same mass as the smaller Eurasian brown and grizzly bears and can exceptionally measure as small as 1 m (3.3 ft) in head-and-body length.[17] On the other end of the scale among interior brown bears, exceptional grizzly, Eurasian brown bears, and East Siberian brown bears (U. a. collaris) have been weighed up to 680 kg (1,500 lb), 481 kg (1,060 lb) and 600 kg (1,300 lb), respectively.[46]

Interior brown bears are generally smaller than is often perceived, being around the same weight as an average African lion at an estimate average of 180 kg (400 lb) in males and 135 kg (298 lb) in females.[49]

The largest inland brown bear subspecies appears to be the Ussuri brown bear (U. a. lasiotus), likely the ancestor of the modern-day American grizzly, which can obtain sizes comparable to those of the coastal bears as described below.[21]

Due to the lack of genetic variation within subspecies, the environmental conditions in a given area likely plays the largest part in such weight variations.[17]

The largest subspecies, the Kodiak bear (U. a. middendorffi), rivals the polar bear as the largest member of the bear family (Ursidae).[17][50][51][51][52] The Ussuri brown bear and also the extinct California grizzly[15][53] approached the Kodiak brown bear in size.[21] Leopold (1959) described the Mexican grizzly that, according to Rausch (1963), qualifies as the smallest form of U. a. horribilis in North America.[15] One California grizzly weighed 2,200 lb (1,000 kg).[53]

Size fluctuates depending upon sex, age, individual, geographic location, and season. The normal range of physical dimensions for a brown bear is a head-and-body length of 1.4 to 2.8 m (4.6 to 9.2 ft) and a shoulder height of 70 to 153 cm (28 to 60 in). Males within one area are larger than females, typically weighing around 30% more.

The tail is relatively short, ranging from 6 to 22 cm (2.4 to 8.7 in) in length.[54]

Generally speaking, brown bears weigh the least when they emerge from hibernation in the spring and then reach peak weights when preparing for hibernation in the fall (when they often gorge on large food stuffs).[17]

Ecotype or regional populations[edit]

An adult grizzly living inland in Yukon may weigh as little as 80 kg (180 lb), while an adult coastal brown bear in nearby coastal Alaska living on a steady, nutritious diet of spawning salmon may weigh as much as 680 kg (1,500 lb).[17]

Young of the year typically weigh 2–27 kg (4.4–59.5 lb), while yearlings typically weigh 9–37 kg (20–82 lb).[17]

The brown bears found in coastal regions of Alaska and far eastern Russia are the largest. The largest subspecies is the Kodiak bear (U. a. middendorffi), followed closely by the Kamchatka brown bear (U. a. beringianus), although bears from other coastal regions of eastern Asia and western North America can be comparably large. In these areas, the female averages from 181 to 318 kg (399 to 701 lb) and the male averages from 272 to 635 kg (600 to 1,400 lb).[17][51] The maximum size of a Kodiak or brown bears in general is not certain. Exceptionally huge male Kodiaks can stand over 3 m (9.8 ft) tall while on their hind legs and may exceed 1.5 m (5 ft) high at the shoulder.[55] The weight of these enormous bruins is less certain. The heaviest verified wild Kodiak bear weighed about 751 kg (1,656 lb), as per the Gerard Wood's 1983 publication for the Guinness Book of World Records.[46] Gary Brown, in his 1996 publication "The Great Bear Alamanc", claimed that the heaviest recorded brown bear weighed over 1,150 kilograms (2,500 lb), without going into further detail beyond mentioning it was of the Kamchatka brown bear subspecies rather than the Kodiak.[23] David W. MacDonald, a well-respected biologist claimed in his guide to the "Mammals of Europe" (2002) while reviewing the species that it is not unusual for Kodiak bear males to weigh up to 680 kg (1,500 lb) during autumn hypophagia and some specimens can attain 780 kg (1,720 lb) or more in mass. MacDonald also mentions in this account that the Kodiak can attain a maximum mass of 1,500 kg (3,300 lb) although he again does not specify verifiable details.[56]

Distribution and habitat[edit]

The brown bear currently occurs in Afghanistan, Albania, Andorra (recently reoccupied), Armenia, Azerbaijan, Belarus, Bhutan (possibly extinct), Bosnia and Herzegovina, Bulgaria, Canada, China, Croatia, Czech Republic (possibly only vagrants), Estonia, Finland, France, Georgia, Greece, India, Iran, Iraq, Italy, Japan, Kazakhstan, Kyrgyzstan, Latvia, Macedonia, Mongolia, Montenegro, Nepal, North Korea, Norway, Pakistan, Poland, Romania, Russian Federation, Serbia, Slovakia, Slovenia, Spain, Sweden, Tajikistan, Turkey, Turkmenistan, Ukraine, United States, and Uzbekistan.[1]

Brown bear at Brooks Falls

Brown bears were once native to much of Asia, the Atlas Mountains of Africa, Europe, and North America,[57] but are now extinct in some areas, and their populations have greatly decreased in other areas. There are approximately 200,000 brown bears left in the world.[58] The largest populations are in Russia with 120,000,[59] the United States with 32,500, and Canada with around 25,000. In Europe, there are 14,000 brown bears in ten fragmented populations, from Spain (estimated at only 20–25 animals in the Pyrenees in 2010,[60][61] in a range shared between France, Spain and Andorra, and some 210 animals in Asturias, Cantabria, Galicia and León, in the Picos de Europa and adjacent areas in 2013 [62] in the west, to Russia in the east, and from Sweden and Finland in the north to Romania (4000–5000), Bulgaria (900–1200), Slovakia (with about 600–800 animals), Slovenia (500–700 animals) and Greece (with about 200 animals) in the south. They are extinct in the British Isles, extremely threatened in France and Spain, and in trouble over most of Central Europe.

The Carpathian brown bear population of Romania is the largest in Europe outside Russia, estimated at 4,500 to 5,000 bears, although declining alarmingly due to overhunting.[63] There is also a smaller brown bear population in the Carpathian Mountains in Ukraine (estimated at about 200 in 2005), Slovakia and Poland (estimated at about 100 in 2009 in the latter country).[64] The total Carpathian population is estimated at about 8,000.[65]

Northern Europe is home to a large bear population, with an estimated 2,500 (range 2,350–2,900) in Sweden, about 1,600 in Finland,[66] about 700 in Estonia and 70 in Norway. Another large and relatively stable population of brown bears in Europe, consisting of 2,500–3,000 individuals, is the Dinaric-Pindos (Balkans) population, with contiguous distribution in northeast Italy, Slovenia, Croatia, Bosnia and Herzegovina, Serbia, Montenegro, Macedonia, Albania, Bulgaria and Greece.[67] Brown bears inhabited the mountains of Austria until as recently as 2011, after a reintroduction effort failed and the species became extinct again. There is currently no effort to reintroduce the species into Austria.[68]

This species inhabits the broadest range of habitats of any living bear species.[45] They seem to have no altitudinal preferences and have been recorded from sea-level to an elevation of 5,000 m (16,000 ft) (the latter in the Himalayas).[45] In most of their range, brown bears generally seems to prefer semiopen country, with a scattering of vegetation that can allow them a resting spot during the day. However, they have been recorded as inhabiting every variety of northern temperate forest known to occur.[45] North American brown bears, or grizzly bears, generally seem to prefer open or semi-open landscapes, with the species once having been common on the Great Plains and continues to occur in sizeable numbers in tundra and coastal estuaries and islands. Variable numbers still occur in prairie areas of the northern Rocky Mountains (mostly in Canada but some in the contiguous United States).[45]

In western Eurasia, they inhabit mostly mountainous woodlands, in ranges such as the Alps, the Pyrenees and the Caucasus, though they may have been driven into more wooded, precipitous habitats due to the prior extensive persecution of the species in some regions.[56][69] Desolate parts of northern and eastern Europe, like large patches of Scandinavia and the Carpathian Mountains, have always been quite heavily forested and have maintained relatively stable populations of bears, indicating that the brown bears here are well-adapted to forest-dwelling.[45]

In Central Asia, human disturbances are minimal as this area has a harsher environment and is more sparsely populated. In this part of the world, bears may be found in steppe, alpine meadows and even desert edge. In Siberia, the species seems well-adapted to living in denser pine forests. Eastern Russian forests hold arguably the largest number of brown bears in the world outside of possibly Alaska and northeastern Canada.[45] It is thought the Eurasian bears which colonized America were tundra-adapted and the species is sometimes found around sub-Arctic ice fields. This is indicated by brown bears in the Chukotka Peninsula on the Asian side of Bering Strait, which are the only Asian brown bears to live year-round in lowland tundra like their North American cousins.[70]

North America[edit]

The brown bear is usually called the grizzly bear in North America. It once ranged throughout much of the entire continent.[71]

Canada[edit]

As many as 20,000 brown bears range throughout the Yukon, Northwest Territories, and British Columbia, and in the western half of Alberta.[71]

United States[edit]

The brown bear has lost 98% of its habitat in the lower 48 states. About 95% of the brown bear population in the United States is in Alaska, though in the lower 48 states, they are repopulating slowly but steadily along the Rockies and the western Great Plains.

Almost all U.S. bears live in Alaska. The Alaskan population is estimated at 32,000 individuals.[72] The largest other populations of brown bears are found in the 23,300-km2 Greater Yellowstone Ecosystem and the 24,800-km2 Northern Continental Divide Ecosystem.[73] Small populations exist in the Greater Yellowstone Ecosystem of northwest Wyoming (with about 674–839 animals), the Northern Continental Divide Ecosystem of northwest Montana (with about 765 animals), the Cabinet-Yaak Ecosystem of northwest Montana and northeast Idaho (with about 42–65 animals), the Selkirk Ecosystem of northeast Washington and northwest Idaho (with about 40–50 animals), and the North Cascades Ecosystem of northcentral Washington (with about 5–10 animals). These five ecosystems combine for a total of a maximum 1,729 wild grizzlies still persisting in the contiguous United States. Unfortunately, these populations are isolated from each other, inhibiting any genetic flow between ecosystems.[73] This poses one of the greatest threats to the future survival of the grizzly bear in the contiguous United States.[73]

Mexico[edit]

Although many people hold the belief some brown bears may be present in Mexico they are almost certainly extinct.[74] The last Mexican grizzly bear was shot in 1976.[75] None had been seen since 1960 prior.

Asia[edit]

In Asia, brown bears are found throughout Russia, thence more spottily southwest to parts of the Middle East, to as far south as southwestern Iran, and to the southeast in a small area of Northeast China, Western China, and parts of North Korea, Pakistan, Afghanistan and India.

Three distinct lineages of the Hokkaido brown bear (Ursus arctos yesoensis L.) can also be found on the Japanese island of Hokkaidō.[76][77] Hokkaido has the largest number of non-Russian brown bears in eastern Asia with about 2,000–3,000 animals,[45] although, in 2015, the Biodiversity Division of the Hokkaido government estimated the population as being as high as 10,600.[78]

Africa[edit]

Many people hold the belief some brown bears may be present in the Atlas Mountains of Morocco but there have been none sighted in the last century.

In addition to the native Atlas bear the Romans apparently imported bears from Spain for spectacles with some escaping and founding a population in Africa.[14]

Europe and Asia[edit]

As of 2015 a stable population of around 250 individuals is present in the Cantabrian mountains of North West Spain.[citation needed] However, the population of brown bears in the Pyrenees mountain range between France and Spain is extremely low, estimated at 14 to 18, with a shortage of females.[1] Their rarity in this area has led biologists to release bears, mostly female, from Slovenia in spring 2006 to reduce the imbalance and preserve the species' presence in the area. The bears were released despite protests from French farmers. A small population of brown bears (Ursus arctos marsicanus) still lives in central Italy (Apennine Mountains, Abruzzo and Latium), with no more than 70 individuals, protected by strong laws, but endangered by the human presence in the area.[citation needed]

In Arctic areas, the potential habitat of the brown bear is increasing. The warming of that region has allowed the species to move farther north into what was once exclusively the domain of the polar bear. In non-Arctic areas, habitat loss is blamed as the leading cause of endangerment, followed by hunting.

Brown bear populations in Northern Italy, particularly in the Trentino areas as well as Slowenia and Bosnia-Herzegovina, have recovered remarkably through natural protection of viable sub-groups and enduring international restrictions on illegal hunting and poaching. The European population in those areas is expanding its range into Austria and Switzerland. The current estimate of the number of European brown bears in the Croatian segment of the Dinara-Pindos population is about 1000 individuals.[79]

Iran[edit]

The brown bear occurs in Iran, Iraq and Turkey.[80] At least 20–30 were present in northern Iran a few years ago[when?].[81]

Conservation status[edit]

While the brown bear's range has shrunk and it has faced local extinctions, it remains listed as a least concern species by the IUCN with a total population of approximately 200,000. As of 2012, this and the American black bear are the only bear species not classified as threatened by the IUCN. However, the Californian, North African (Atlas bear), and Mexican subspecies were hunted to extinction in the nineteenth and twentieth centuries, and many of the southern Asian subspecies are highly endangered. The Syrian brown bear (Ursus arctos syriacus) is very rare.[82] The smallest subspecies, the Himalayan brown bear, is critically endangered, occupying only 2% of its former range and threatened by uncontrolled poaching for its parts.[3] The Marsican brown bear in central Italy is believed to have a population of just 30 to 40 bears.

The brown bear's principal range includes parts of Russia, India, China, Canada, the United States (mostly Alaska), Japan, Scandinavia and the Carpathian region (especially Romania),[4]

Regional extinctions[edit]

The brown bear is extinct in: Algeria, Belgium, Denmark, Egypt; Germany; Hungary; Ireland, Israel; Jordan, Lebanon; Liechtenstein; Lithuania, Luxembourg, Mexico; Moldova; Monaco, Morocco; Netherlands, Oregon, Portugal; San Marino; Germany; Syrian Arab Republic, Tunisia, United Kingdom, Vatican. (Possibly extinct in Bhutan.)[1]

Behavior[edit]

Although the brown bear is primarily nocturnal, it is frequently seen in morning and early evening hours.[83] In summer through autumn, it can double its weight, gaining up to 180 kg (400 lb) of fat, on which it relies to make it through winter, when it becomes very lethargic. Although they are not full hibernators and can be woken easily, both sexes like to den in a protected spot, such as a cave, crevice, or hollow log, during the winter months. Brown bears are mostly solitary, although they may gather in large numbers at major food sources (e.g., moth colonies, open garbage dumps or rivers holding spawning salmon) and form social hierarchies based on age and size.[84] Adult male bears are particularly aggressive and are avoided by adolescent and subadult males. Female bears with cubs rival adult males in aggression, and are more intolerant of other bears than single females. Young adolescent males tend to be least aggressive, and have been observed in nonantagonistic interactions with each other. Dominance between bears is asserted by making a frontal orientation, showing off canines, muzzle twisting and neck stretching to which a subordinate will respond with a lateral orientation, by turning away and dropping the head and by sitting or lying down.[85] During combat, bears use their paws to strike their opponents in the chest or shoulders and bite the head or neck. In his Great Bear Almanac, Gary Brown lists 11 different sounds bears produce in 9 different contexts. Sounds expressing anger or aggravation include growls, roars, woofs, champs and smacks, while sounds expressing nervousness or pain include woofs, grunts, and bawls. Sows will bleat or hum when communicating with their cubs.[23]

On Shiretoko Peninsula, especially in the area called "Banya", many females with cubs often approach fishermen and spend time nearby mankind. This unique behavior was firstly noted more than a half century ago, and not a single casualties or accidents ever recorded. It is speculated that females take cubs and approach fishermen to avoid encountering with adult males.[86]

Home ranges[edit]

Brown bears usually occur over vast home ranges, however they are not highly territorial. Several adult bears often roam freely over the same vicinity without issue unless rights to a fertile female or food sources are being contested. Males always cover more area than females each year and will try to mate with as many females as they can (although females are not monogamous either). In areas where food is abundant and concentrated, such as coastal Alaska, home ranges for females are up to 24 km2 (9.3 sq mi) and for males are up to 89 km2 (34 sq mi). Similarly, in British Columbia, bears of the two sexes travel relatively compact home ranges of 115 km2 (44 sq mi) and 318 km2 (123 sq mi). In Yellowstone National Park, home ranges for females are up to 281 km2 (108 sq mi) and up to 874 km2 (337 sq mi) for males. In the central Arctic of Canada, where food sources are quite sparse, home ranges range up to 2,434 km2 (940 sq mi) in females and 8,171 km2 (3,155 sq mi) in males.[45]

A study of male-inherited Y chromosome DNA sequence found that brown bears, over the past few 10,000 years, have shown strong male-biased dispersal[87] That study found surprisingly similar Y chromosomes in brown bear populations as far apart as Norway and coastal Alaska, indicating extensive gene flow across Eurasia and North America. Notably, this contrasts with genetic signals from female-inherited mitochondrial DNA (mtDNA), where brown bears of different geographic regions typically show strong differences in their mtDNA, a result of female philopatry.

Reproduction[edit]

The mating season is from mid-May to early July.[45] Being serially monogamous, brown bears remain with the same mate from several days to a couple of weeks.[88] Females mature sexually between the age of 4 and 8 years of age, while males first mate about a year later on average, when they are large and strong enough to successfully compete with other males for mating rights.[45]

Pair of mating brown bears, at Ähtäri Zoo in Ähtäri, Finland

Males, however, take no part in raising their cubs – parenting is left entirely to the females.

Through the process of delayed implantation, a female's fertilized egg divides and floats freely in the uterus for six months. During winter dormancy, the fetus attaches to the uterine wall. The cubs are born eight weeks later, while the mother sleeps. If the mother does not gain enough weight to survive through the winter, the embryo does not implant and is reabsorbed into the body. The average litter has one to four cubs, usually two. There have been cases of bears with as many as six cubs.[45] There are records of females sometimes adopting stray cubs or even trading cubs when they emerge from hibernation.[45] Older females tend to give birth to larger litters. The size of a litter also depends on factors such as geographic location and food supply. At birth, the cubs are blind, toothless, hairless, and weigh less than 450 grams (1 lb). They feed on their mother's milk until spring or even early summer, depending on climate conditions. At this time, the cubs weigh 7 to 9 kg (15 to 20 lb) and have developed enough to follow her and begin to forage for solid food.

Cubs remain with their mother from two to four years (exceptionally to 4 and a half years), during which time they learn survival techniques, such as which foods have the highest nutritional value and where to obtain them; how to hunt, fish, and defend themselves; and where to den. The cubs learn by following and imitating their mother's actions during the period they are with her.[89] Brown bears practice infanticide.[90] An adult male bear may kill the cubs of another bear either to make the female sexually receptive or simply for consumption. Cubs flee up a tree when they see a strange male bear, and the mother often successfully defends them, even though the male may be twice as heavy as she.

Longevity[edit]

The brown bear is naturally a long-living animal. Wild females have been known to be able to reproduce at as old as 28 years of age, the oldest known age for reproduction in the wild of any ursid.[45] Males commonly can live to 25 years, with the oldest female having been 37 years old. The species can live to 48 years of age in captivity. Annual mortality for bears of any age is estimated at around 10% in most protected areas.[45] However, in hunted populations, an estimated average mortality rate of 38% is given.[45] Around 13% to 44% of cubs die within their first year. Beyond predation by other large predators (rarely by wolf packs or Siberian tigers (Panthera tigris altaica)) and brown bears, starvation and accidents claim a few cubs. Even in populations living in protected areas without legal, non-governmental hunting, though, humans are still the leading cause of mortality for brown bears.[91] The largest number of legally hunting on the species occurs in Canada, Finland, Russia, Slovakia and Alaska.[45]

Dietary habits[edit]

Brown bear feeding on salmon

The brown bear is one of the most omnivorous animals in the world and has been recorded as consuming the greatest variety of foods of any bear.[45] Throughout life, this species is regularly curious about the potential of eating virtually any organism or object that they encounter. Food that is both abundant and easily accessed or caught is preferred. Their jaw structure has evolved to fit their dietary habits. Their diet varies enormously throughout their differing areas based on opportunity.

Brown bears are capable of obtaining practically all forms of the mammals: from the mouse-like rodents to the tiger.[92] Despite their reputation, most brown bears are not highly carnivorous, as they derive up to 90% of their dietary food energy from vegetable matter.[93] They often feed on a variety of plant life, including berries, grasses, flowers, acorns (Quercus ssp.) and pine cones as well as fungi such as mushrooms.[17] Among all bears, brown bears are uniquely equipped to dig for tough foods such as roots and shoots. They use their long, strong claws to dig out earth to reach the roots and their powerful jaws to bite through them.[17] In spring, winter-provided carrion, grasses, shoots, sedges and forbs are the dietary mainstays for brown bears internationally.[45] Fruits, including berries, become increasingly important during summer and early autumn. Roots and bulbs become critical in autumn for some inland bear populations if fruit crops are poor.[45] They will also commonly consume animal matter, which in summer and autumn may regularly be in the form of insects, larvae and grubs, including beehives. Bears in Yellowstone eat an enormous number of moths during the summer, sometimes as many as 40,000 army cutworm moths (Euxoa auxiliaris) in a single day, and may derive up to half of their annual food energy from these insects.[94] Brown bears living near coastal regions will regularly eat crabs and clams. In Alaska, bears along the beaches of estuaries regularly dig through the sand for clams.[23] This species may eat birds and their eggs, including almost entirely ground- or rock-nesting species, ranging from ptarmigan (Lagopus ssp.) to Steller's sea eagles (Haliaeetus pelagicus).[23][95][96] The diet may be supplemented by rodents or similar smallish mammals, including hares (Lepus ssp.), marmots (Marmota ssp.), ground squirrels, mice, rats, lemmings and voles.[17][23][95] With particular regularity, bears in Denali National Park will wait at burrows of Arctic ground squirrels (Spermophilus parryii) hoping to pick off a few of the 1 kg (2.2 lb) rodents.[97]

In the Kamchatka peninsula and several parts of coastal Alaska, brown bears feed mostly on spawning salmon, whose nutrition and abundance explain the enormous size of the bears in these areas. The fishing techniques of bears are well-documented. They often congregate around falls when the salmon are forced to breach the water, at which point the bears will try to catch the fish in mid-air (often with their mouths). They will also wade into shallow waters, hoping to pin a slippery salmon with their claws. While they may eat almost all the parts of the fish, bears at the peak of spawning, when there is usually a glut of fish to feed on, may eat only the most nutritious parts of the salmon (including the eggs and head) and then indifferently leave the rest of the carcass to scavengers, which can include red foxes, bald eagles, common ravens and gulls. Despite their normally solitary habits, brown bears will gather rather closely in numbers at good spawning sites. The largest and most powerful males claim the most fruitful fishing spots and bears (especially males) will sometimes fight over the rights to a prime fishing spot.[23]

Beyond the regular predation of salmon, most brown bears are not particularly active predators.[23] While perhaps a majority of bears of the species will charge at large prey at one point in their lives and most eat carrion, many predation attempts start with the bear clumsily and half-heartedly pursuing the prey and end with the prey escaping alive.[23] On the other hand, some brown bears are quite self-assured predators who habitually pursue and catch large prey items. Such bears are usually taught how to hunt by their mothers from an early age.[23] Large mammals preyed on can include various deer species such as elk (Cervus canadensis), red deer (Cervus elaphus), axis deer (Axis axis), European roe deer (Capreolus capreolus), Siberian roe deer (Capreolus pygargus), fallow deer (Dama dama), mule deer (Odocoileus hemionus), white-tailed deer (Odocoileus virginianus), moose (Alces alces) and caribou (Rangifer tarandus). Bovids are also regular prey including various sheep, goats, antelope, bison (Bison ssp.) and muskoxen (Ovibos moschatus), as are wild boars (Sus scrofa).[17] When brown bears attack these large animals, they usually target young or infirm ones, as they are easier to catch. Typically when hunting (especially with young prey), the bear pins its prey to the ground and then immediately tears and eats it alive.[98] In northeastern Norway, it was found that moose were the most important single food item (present in up to 45% of scats and locally comprising more than 70% of the bear's dietary energy) for local brown bears and several local bears appear to be specialized moose hunters, most often picking off sickly yearling moose and pregnant but healthy cows.[99] In a study from Yellowstone National Park, grizzly bears who derived much of the food energy from ungulates were studied, 30% of the ungulates consumed were through predation, the remaining amount from scavenging of carcasses, elk, bison and moose (the 3 largest native ungulates in the region) each nearly comprised a quarter each of the overall ungulate diet. 13% of the total ungulates actively hunted and killed per that study in Yellowstone were elk calves, while 8% of the actively and successfully hunted prey there were adult cow elk.[100] On one occasion in Yellowstone, a female grizzly bear with two cubs was exceptionally observed to successfully prey on an adult bull bison.[101] Brown bears will also bite or swipe some prey in order to stun it enough to knock it over for consumption.[56] To pick out young or infirm individuals, bears will charge at herds so the slower-moving and more vulnerable individuals will be made apparent. brown bears may also ambush young animals by finding them via scent.[17] When emerging from hibernation, brown bears, whose broad paws allow them to walk over most ice and snow, may pursue large prey such as moose whose hooves cannot support them on encrusted snow.[56] Similarly, predatory attacks on large prey sometimes occur at riverbeds, when it is more difficult for the prey specimen to run away due to muddy or slippery soil.[17] On rare occasions, while confronting fully-grown, dangerous prey, bears kill them by hitting with their powerful forearms, which can break the necks and backs of large creatures such as adult moose and adult bison.[23] They also feed on carrion, and use their size to intimidate other predators, such as wolves, cougars (Puma concolor), tigers, and black bears from their kills. Carrion is especially important in the early spring (when the bears are emerging from hibernation), much of it comprised by winter-killed big game.[17] Cannibalism is not unheard of, though predation is not normally believed to be the primary motivation when brown bears attack each other.[23]

When forced to live in close proximity with humans and their domesticated animals, bears may potentially predate any type of domestic animal. Among these, domestic cattle (Bos primigenius taurus) are sometimes exploited as prey. Cattle are bitten on the neck, back or head and then the abdominal cavity is opened for eating.[17] In Norway, free-ranging domestic sheep (Ovis aries) are numerous and the local brown bears derive 65–87% of their dietary energy in late summer from sheep. Due to the vulnerability of this prey, examination of sheep remains suggest many of the sheep consumed there are adults that were killed by the bears rather than merely scavenged, and thus some local farmers received partial compensation for their stock losses. In nearby northern Sweden, free-ranging sheep are not present and the bear derive their food predominantly from natural sources.[102] Plants and fruit farmed by humans are readily consumed as well, including corn (Zea mays ), wheat (Triticum spp.), sorghum (Sorghum ssp.), melons and any form of berries.[23] They will also feed at domestic bee farms, readily consuming both honey and the contents of the honey bee (Apis mellifera) colony.[23] Human foods and trash or refuse is eaten when possible. When an open garbage dump was kept in Yellowstone, brown bears were one of the most voracious and regular scavengers. The dump was closed after both brown and American black bears came to associate humans with food and lost their natural fear of them.[23]

Enemies and competitors[edit]

Brown bear and cubs attacked by gray wolves
Taxidermy exhibit portraying a brown bear fighting a Siberian tiger, Vladivostok Museum

Adult bears are generally immune to predatory attacks except from tigers and other bears. Siberian tigers prefer preying on young bears but smaller, fully grown adult female brown bears outside their dens may also be taken.[103][104][105] Of 44 recorded encounters between the two predators, 20 resulted in confrontations; in 50% of these, the brown bears were killed, in 27% the tigers were killed, and 23% of the cases ended with both animals surviving and parting ways.[106] Some bears emerging from hibernation seek out tigers in order to steal their kills.[107] Despite the possibility of tiger predation, some large brown bears may actually benefit from the tiger's presence by appropriating tiger kills that the bears may not be able to successfully hunt themselves and follow tiger tracks.[108] Geptner et al. (1972) stated bears are generally afraid of tigers and change their path after coming across tiger trails.[109] In the winters of 1970–1973, Yudakov and Nikolaev recorded 1 case of brown bear showing no fear of the tigers and another case of brown bear changing path upon crossing tiger tracks.[110] Other researchers have observed bears following tiger tracks for various reasons.[111] Bears frequently track down tigers to usurp their kills, with fatal outcomes for the tiger. A report from 1973 describes twelve known cases of brown bears killing tigers; in all cases the tigers were eaten by the bears.[112] There are reports of brown bears specifically targeting Amur tigers and leopards to abstract their prey. In the Sikhote-Alin reserve, 35% of tiger kills were stolen by bears, with tigers either departing entirely or leaving part of the kill for the bear.[92]

Brown bears regularly intimidate gray wolves (Canis lupus) away from their kills. In Yellowstone National Park, brown bears pirate wolf kills so often, Yellowstone's Wolf Project director Doug Smith wrote, "It's not a matter of if the bears will come calling after a kill, but when." Despite the high animosity between the two species, most confrontations at kill sites or large carcasses end without bloodshed on either side. Though conflict over carcasses is common, on rare occasions the two predators tolerate each other on the same kill. To date, there is a single case of fully-grown wolves being killed by a grizzly bear.[113] Given the opportunity, however, both species will prey on the other's cubs.[114] Conclusively, the individual power of the bear against the collective strength of the wolf pack usually results in a long battle for kills or domination.

In some areas, the brown bear also regularly displaces cougars from their kills.[115] Cougars kill small bear cubs on rare occasions, but there was one report of a bear killing a cougar of unknown age and condition between 1993 and 1996.[116][117] Smaller carnivorous animals, including coyotes (Canis latrans), wolverines (Gulo gulo), lynxes (Lynx ssp.) and any other sympatric carnivores or raptorial birds, are dominated by brown bears and generally avoid direct interactions with them, unless attempting to steal scraps of food. However, wolverines have been persistent enough to fend off a grizzly bear as much as ten times their weight off a kill.[23] There is one record of a golden eagle (Aquila chrysaetos) predating a brown bear cub.[118]

Brown bears usually dominate other bear species in areas where they coexist. Due to their smaller size, American black bears are at a competitive disadvantage to brown bears in open, unforested areas. Although displacement of black bears by brown bears has been documented, actual interspecific killing of black bears by brown bears has only occasionally been reported. Confrontation is mostly avoided due to the black bear's diurnal habits and preference for heavily forested areas, as opposed to the brown bear's largely nocturnal habits and preference for open spaces.[119] Brown bears may also kill Asian black bears, though the latter species probably largely avoids conflicts with the Brown bear due to similar habits and habitat preferences to the American Black species.[120] They will eat the fruit dropped by the Asian black bear from trees, as they themselves are too large and cumbersome to climb.[121] Improbably, in the Himalayas, brown bears are reportedly intimidated by Asian black bears in confrontations.[122]

There has been a recent increase in interactions between brown bears and polar bears, theorized to be caused by climate change. Brown bears have been seen moving increasingly northward into territories formerly claimed by polar bears. Brown bears tend to dominate polar bears in disputes over carcasses,[123] and dead polar bear cubs have been found in brown bear dens.[124]

Large herbivores, such as moose, buffalo, and muskox may have an intolerance of brown bears due to their possible threat to calves; moose regularly charge grizzly bears in their calf's defense.[125][126] Bison have been known to fatally injure lone grizzly bears in battles, and even a mountain goat was observed to do so with its horns.[23]

Relationship with humans[edit]

Front paw imprint
Rear paw imprint

Conflicts between bears and humans[edit]

Bears become attracted to human-created food sources, such as garbage dumps, litter bins, and dumpsters; they venture into human dwellings or barns in search of food as humans encroach into bear habitats. In the U.S., bears sometimes kill and eat farm animals. When bears come to associate human activity with a "food reward", they are likely to continue to become emboldened; the likelihood of human-bear encounters increases, as they may return to the same location despite relocation. The saying "a fed bear is a dead bear" has come into use to popularize the idea that allowing bears to scavenge human garbage, such as trash cans and campers' backpacks, pet food, or other food sources that draw the bear into contact with humans, can result in a bear's death.

Relocation of the bear has been used to separate the bear from the human environment, but it does not address the problem of the bear's newly learned association of humans with food or the environmental situations which created the human-habituated bear. "Placing a bear in habitat used by other bears may lead to competition and social conflict, and result in the injury or death of the less dominant bear."[127]

Yellowstone National Park, a reserve located in the western United States, contains prime habitat for the grizzly bear (Ursus arctos horribilis), and due to the enormous number of visitors, human-bear encounters are common. The scenic beauty of the area has led to an influx of people moving into the area. In addition, because there are so many bear relocations to the same remote areas of Yellowstone, and because male bears tend to dominate the center of the relocation zone, female bears tend to be pushed to the boundaries of the region and beyond. As a result, a large proportion of repeat offenders, bears that are killed for public safety, are females. This creates a further depressive effect on an already endangered species. The grizzly bear is officially described as "threatened" in the U.S. Though the problem is most significant with regard to grizzlies, these issues affect the other types of brown bears as well.

In Europe, part of the problem lies with shepherds; over the past two centuries, many sheep and goat herders have gradually abandoned the more traditional practice of using dogs to guard flocks, which have concurrently grown larger. Typically, they allow the herds to graze freely over sizeable tracts of land. As bears reclaim parts of their range, they may eat livestock. In some cases, the shepherds shoot the bear, thinking their livelihood is under threat. Many are now better informed about the ample compensation available, and will make a claim when they lose livestock to a bear.

It is likely that humans caused extinction of bear populations and fragmentation of their habitats since prehistorical time. It is, for instance, shown that bear populations from the Greater Caucasus and the Lesser Caucasus Mountains, separated by the densely populated Transcaucasian Depression, have been matrilineally isolated since the early Holocene era, i.e., after permanent human settlements appeared throughout the area[69]

Relationship with Native Americans[edit]

Gorgonia, a Native American (Mescalero Apache) man. He holds a bear pelt and wears moccasin boots, a breechcloth, kilt, and vest.

Native American tribes sympatric with brown bears often view them with a mixture of awe and fear. North American brown bears have at times been so feared by the natives, that they were rarely hunted, especially alone. At traditional grizzly hunts, the expedition was conducted with the same preparation and ceremoniality as intertribal warfare, and was never done except with a company of 4–10 warriors.[clarification needed] The tribe members who dealt the killing blow were highly esteemed among their compatriots. Californian natives actively avoided prime bear habitat, and would not allow their young men to hunt alone, for fear of bear attacks. During the Spanish colonial period, some tribes, instead of hunting grizzlies themselves, would seek aid from European colonists to deal with problem bears. Many authors in the American west wrote of natives or voyageurs with lacerated faces and missing noses or eyes due to attacks from grizzlies.[11]

Sleeping Bear Dunes is named after a Native American legend, where a female bear and her cub swam across Lake Michigan. Exhausted from their journey, the bears rested on the shoreline and fell sound asleep. Over the years, the sand covered them up, creating a huge sand dune.

Many Native American tribes both respect and fear the brown bear, even thinking of it as a god.[citation needed] One tale tells of how the black bear was a creation of the Great Spirit, while the grizzly was created by the Evil Spirit.[clarification needed][128] In Kwakiutl mythology, black and brown bears became enemies when Grizzly Bear Woman killed Black Bear Woman for being lazy. Black Bear Woman's children, in turn, killed Grizzly Bear Woman's own cubs.[129]

Bear encounters[edit]

There are an average of two fatal attacks by bears per year in North America.[130] In Scandinavia, there are only four known cases since 1902 of bear encounters which have resulted in death. The two most common causes for bear attack are surprise and curiosity.[131] Some types of bears, such as polar bears, are more likely to attack humans when searching for food, while American black bears are much less likely to attack.

The Alaska Science Center ranks the following as the most likely reasons for bear attacks:[131]

  1. Surprise
  2. Curiosity
  3. Invaded personal space (this includes a mother bear protecting her young)
  4. Predatory intent
  5. Hunting wounded
  6. Carcass defense
  7. Provoked charge

Aggressive behavior in brown bears is favored by numerous selection variables. Unlike the smaller black bears, adult brown bears are too large and have improperly shaped claws to escape danger by climbing trees, so they respond to danger by standing their ground and warding off their attackers. Increased aggressiveness also assists female brown bears in better ensuring the survival of their young to reproductive age.[132] Mothers defending cubs are the most prone to attacking, being responsible for 70% of brown bear-caused human fatalities in North America.[133]

Attacks on humans[edit]

Drum or barrel trap used to safely relocate bears; currently parked adjacent to a building in Grand Teton National Park in Wyoming, United States

Brown bears seldom attack humans on sight, and usually avoid people. They are, however, unpredictable in temperament, and may attack if they are surprised or feel threatened.[134] Sows with cubs account for the majority of injuries and fatalities in North America. Habituated or food-conditioned bears can also be dangerous, as their long-term exposure to humans causes them to lose their natural shyness, and, in some cases, to associate humans with food. Small parties of one or two people are more often attacked than large groups, with only one known case of an attack on a group of six or more. In that instance, it is thought that due to surprise, the bear may not have recognized the size of the group.[135] In the majority of attacks resulting in injury, brown bears precede the attack with a growl or huffing sound,[134] In contrast to injuries caused by American black bears, which are usually minor, brown bear attacks tend to result in serious injury and, in some cases, death.[134] Brown bears seem to confront humans as they would when fighting other bears: they rise up on their hind legs, and attempt to "disarm" their victims by biting and holding on to the lower jaw to avoid being bitten in turn.[11] Due to the bears' enormous physical strength, even a single bite or swipe can be deadly, as in tigers, with some human victims having had their heads completely crushed by a bear bite.[47] Most attacks occur in the months of July, August, and September, the time when the number of outdoor recreationalists, such as hikers or hunters, is higher. People who assert their presence through noises tend to be less vulnerable, as they alert bears to their presence. In direct confrontations, people who run are statistically more likely to be attacked than those who stand their ground. Violent encounters with brown bears usually last only a few minutes, though they can be prolonged if the victims fight back.[134]

Attacks on humans are considered extremely rare in the former Soviet Union, though exceptions exist in districts where they are not pursued by hunters.[21] Siberian bears, for example, tend to be much bolder toward humans than their shyer, more persecuted European counterparts. In 2008, a platinum mining compound in the Olyotorsky district of northern Kamchatka was besieged by a group of 30 bears, who killed two guards and prevented workers from leaving their homes.[136] Ten people a year are killed by brown bears in Russia.[137] In Scandinavia, only three fatal attacks were recorded in the 20th century.[138]

In Japan, a large brown bear nicknamed "Kesagake" (袈裟懸け, "kesa-style slasher") made history for causing the worst bear attack in Japanese history at Tomamae, Hokkaidō during numerous encounters during December 1915. It killed seven people and wounded three others (with possibly another three previous fatalities to its credit) before being gunned down after a large-scale beast-hunt. Today, there is still a shrine at Rokusensawa (六線沢), where the event took place, in memory of the victims of the incident.

Within Yellowstone National Park, injuries caused by grizzly attacks in developed areas averaged approximately one per year during the 1930s through to the 1950s, though it increased to four per year during the 1960s. They then decreased to one injury every two years during the 1970s. Between 1980 and 2002, there have been only two human injuries caused by grizzly bears in a developed area. Though grizzly attacks were rare in the backcountry before 1970, the number of attacks increased to an average of approximately one per year during the 1970s, 1980s, and 1990s.[139]

History of defense from bears[edit]

A study by US and Canadian researchers has found pepper spray to be more effective at stopping aggressive bear behavior than guns, working in 92% of studied incidents versus 67% for guns.[140] Carrying pepper spray is highly recommended by many authorities when traveling in bear country; however, carrying two means of deterrent, one of which is a large caliber gun, is also advised. Solid shotgun slugs, or three buckshot rounds, or a pistol of .44 caliber or more is suggested if a heavy hunting rifle is not available. Guns remain a viable, last resort option to be used in defense of life from aggressive bears.[141] Too often, people do not carry a proper caliber weapon to neutralize the bear. According to the Alaska Science Center, a 12 gauge shotgun with slugs has been the most effective weapon. There have been fewer injuries as a result of only carrying lethal loads in the shotgun, as opposed to deterrent rounds. State of Alaska Defense of Life or Property (DLP) laws require one to report the kill to authorities, and salvage the hide, skull, and claws.[142]

Campers are often told to wear bright colored red ribbons and bells, and carry whistles to ward off bears. They are told to look for grizzly scat in camping areas, and be careful to carry the bells and whistles in those areas. Grizzly scat is difficult to differentiate from black bear scat, as diet is in a constant state of flux depending on the availability of seasonal food items. If a bear is killed near camp, the bear's carcass must be adequately disposed of, including entrails and blood, if possible. Failure to move the carcass has often resulted in it attracting other bears and further exacerbating a bad situation. Moving camps immediately is another recommended method.

A page at the State of Alaska Department of Natural Resources website offers information about how to "select a gun that will stop a bear (12-gauge shotgun or .300 mag rifle)."[143] This information is helpful for people venturing into "bear country", regardless of state or country.

Culture[edit]

"The Story of the Three Bears", illustration from Childhood's Favorites and Fairy Stories

Brown bears often figure into the literature of Europe and North America, in particular that which is written for children. "The Brown Bear of Norway" is a Scottish fairy tale telling the adventures of a girl who married a prince magically turned into a bear, and who managed to get him back into a human form by the force of her love and after many trials and difficulties. With "Goldilocks and the Three Bears", a story from England, the three bears are usually depicted as brown bears. In German speaking countries, children are often told the fairytale of Snow White and Rose Red; the handsome prince in this tale has been transfigured into a brown bear. In the United States, parents often read their preschool age children the book Brown Bear, Brown Bear, What Do You See? to teach them their colors and how they are associated with different animals.

The Russian bear is a common national personification for Russia (as well as the former Soviet Union). The brown bear is also Finland's national animal.

Berni is a brown bear mascot of German football club Bayern Munich

The Bundesliga club Bayern Munich has a brown bear mascot named Berni.

The National Football League (NFL) franchise in Chicago, Illinois, is named the Bears. In this context, no differentiation between black and brown bears is needed.

The school mascot for George Fox University, Brown University, the University of California, Los Angeles, the University of California, Berkeley, the University of California, Riverside, and the University of Alberta is the brown bear.

The coat of arms of Madrid depicts a bear reaching up into a madroño or strawberry tree (Arbutus unedo) to eat some of its fruit, whereas the Swiss city of Bern's coat of arms also depicts a bear and the city's name is popularly thought to derive from the German word for bear.

In the town of Prats de Molló, in Vallespir, southern France, a "bear festival" (festa de l'ós) is celebrated annually at the beginning of spring, in which the locals dress up as bears, cover themselves with soot or coal and oil, and "attack" the onlookers, attempting to get everyone dirty. The festival ends with the ball de l'os (bear dance).

The grizzly bear is the state animal of Montana. The California golden bear is the state animal of California. Both animals are sub-species of the brown bear.

Legal status[edit]

  • The grizzly bear, sometimes called the silvertip bear, is listed as threatened in the contiguous United States. It is slowly repopulating in areas where it was previously extirpated, though it is still vulnerable.
  • The California golden bear (Ursus arctos californicus[144]) disappeared from the state of California in 1922, when the last one was shot in Tulare County. It is the official state animal.[145]
  • The Mexican grizzly bear is listed as an endangered species, but it may be extinct.
  • In Canada, it is listed as vulnerable in Alberta, British Columbia, Northwest Territories, and Yukon Territory. Prairie populations of grizzly bear are listed as extirpated in Alberta, Manitoba, and Saskatchewan.
  • The brown bear is a European Protected Species, given protection throughout the European Union.
  • The brown bear is the national animal of Finland.
  • The brown bear is depicted on the reverse of the Croatian 5 kuna coin, minted since 1993.

See also[edit]

References[edit]

  1. ^ a b c d e McLellan, B. N., Servheen, C. & Huber, D. (2008). "Ursus arctos". IUCN Red List of Threatened Species. Version 2013.1. International Union for Conservation of Nature. Retrieved 2013-10-05. 
  2. ^ Boddington, Craig; illustrations by Carlson, Ken (2004). Fair Chase in North America. Boone and Crockett Club. p. 45. ISBN 0-940864-47-9. 
  3. ^ a b Himalayan brown bears now critically endangered. Euronews.com. 6 January 2014
  4. ^ a b White, Paul. "Brown Bear". Transylvania Wildlife Project. Archived from the original on 19 May 2013. Retrieved 13 January 2014. 
  5. ^ American Heritage Dictionary of the English Language: Fourth Edition. 2000.
  6. ^ Hunting the Grisly and other Sketches. FullTextArchive.com. Retrieved 15 September 2011.
  7. ^ Liddell, Henry George and Scott, Robert. "Ursus." A Latin Dictionary. Perseus Digital Library.
  8. ^ Liddell, Henry George and Scott, Robert."Arktos." A Greek-English Lexicon. Perseus Digital Library.
  9. ^ a b c d e f g h i McLellan, Bruce and Reiner, David C. (1994). A Review of bear evolution. Int. Conf. Bear Res. and Manage. 9(1):85–96
  10. ^ Kurten., Bjorn. The Cave Bear Story 1976. New York.: Columbia University Press. 
  11. ^ a b c Geist, Valerius (1989) "Did Large Predators keep Humans out of North America?", pp. 282–294 in Juliet Clutton-Brock (ed.) The Walking larder: patterns of domestication, pastoralism, and predation, Unwin Hyman, ISBN 0044450133.
  12. ^ Harris, Arthur H. (2013). "Pleistocene Vertebrates of Arizona, New Mexico, and Trans-Pecos Texas". UTEP Biodiversity Collections, University of Texas at El Paso. 
  13. ^ Population Genomics Reveal Recent Speciation and Rapid Evolutionary Adaptation in Polar Bears
  14. ^ a b Calvignac, S.; Hughes, S.; Tougard, C.; Michaux, J.; Thevenot, M.; Philippe, M.; Hamdine, W.; Hanni, C. (2008). "Ancient DNA evidence for the loss of a highly divergent brown bear clade during historical times". Mol. Ecol. 17 (8): 1962–1970. doi:10.1111/j.1365-294x.2008.03631.x. PMID 18363668. 
  15. ^ a b c Storer, T.I.; Tevis, L.P. (1996-12-27). California Grizzly. Berkeley, CA: University of California Press. p. 335. ISBN 0520205200. 
  16. ^ U.S. Fish and Wildlife Service (17 November 2006). "Endangered and Threatened Wildlife and Plants; Designating the Greater Yellowstone Ecosystem Population of Grizzly Bears as a Distinct Population Segment; Removing the Yellowstone Distinct Population Segment of Grizzly Bears From the Federal List of Endangered and Threatened Wildlife" (PDF). Federal Register / Vol. 70, No. 221. pp. 69854–69884. Archived from the original (PDF) on 25 August 2006. Retrieved 1 August 2006. 
  17. ^ a b c d e f g h i j k l m n o p q "Mammalian Species- Ursus arctos" (PDF). American Society of Mammalogists, Smith College. Retrieved 14 June 2012. 
  18. ^ a b c Waits, L.P.; et al. (1998). "Mitochondrial DNA Phylogeography of the North American Brown Bear and Implications for Conservation". Conservation Biology 12 (2): 408–417. doi:10.1111/j.1523-1739.1998.96351.x. 
  19. ^ a b Wozencraft, W.C. (2005). "Ursus arctos". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 588–589. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  20. ^ Ursus arctos, ITIS
  21. ^ a b c d e f g h i j k l m n o p Mammals of the Soviet Union Vol.II Part 1a, SIRENIA AND CARNIVORA (Sea cows; Wolves and Bears), V.G Heptner and N.P Naumov editors, Science Publishers, Inc. USA. 1998. ISBN 1-886106-81-9
  22. ^ Baryshnikov, Gennady; Mano, Tsutmano; Masuda, Ryuchi (2004). "Taxonomic Differentiation of Ursus arctos (Carnivora, Ursidae) from the south Okhotsk Sea Islands". Russian Journal of Theriology 3 (2): 77–88. 
  23. ^ a b c d e f g h i j k l m n o p q r s t Bear Anatomy and Physiology from Gary Brown's The Great Bear Almanac, Lyons & Burford, Publishers, 1993
  24. ^ Ciucci, Paolo; et al. (2014). "Seasonal and annual variation in the food habits of Apennine brown bears, central Italy". Journal of Mammalog 95 (3): 572–586. doi:10.1644/13-mamm-a-218. Retrieved 18 March 2015. 
  25. ^ 野生動物調査痕跡学図鑑』 p.356
  26. ^ Kamchatkan Bear. iza-yoi.net
  27. ^ 体重400キロのヒグマ捕獲 なぜ巨大化? news24.jp (12 October 2015)
  28. ^ Ursus arctos alascensis. Mammal Species of the World. Bucknell University.
  29. ^ "Ursus arctos alascensis Merriam, 1896". Integrated Taxonomic Information System (ITIS). 
  30. ^ Burt. Henry W. (1952) A Field Guide to the Mammals. p. 42.
  31. ^ a b Bear Conservation UK. "Stickeen Brown Bear". Retrieved 15 June 2013. 
  32. ^ SPECIES VARIATION with literature reports for the Brown bear – Ursus arctos. twycrosszoo.org
  33. ^ Miller, CR; Waits, L.P. (2006). "Phylogeography and mitochondrial diversity of extirpated brown bear (Ursus arctos) populations in the contiguous United States and Mexico". Mol Ecol. 15 (14): 4477–85. doi:10.1111/j.1365-294x.2006.03097.x. PMID 17107477. 
  34. ^ "Wild find: Half grizzly, half polar bear: Hunter bags what expert 'never thought would happen' in wild". Associated Press. 11 May 2006. Retrieved 14 May 2006. 
  35. ^ James Mallet (2008). "Hybridization, ecological races and the nature of species: empirical evidence for the ease of speciation" (PDF). Philosophical Transactions of the Royal Society B 363 (1506): 2971–2986. doi:10.1098/rstb.2008.0081. PMC 2607318. PMID 18579473. 
  36. ^ Barnosky, Anthony D. (2009). Heatstroke: Nature in an Age of Global Warming. Washington, DC: Island Press/Shearwater Books. ISBN 1-59726-197-1. 
  37. ^ Hailer, F.; Kutschera, V. E.; Hallstrom, B. M.; Klassert, D.; Fain, S. R.; Leonard, J. A.; Arnason, U.; Janke, A. (2012). "Nuclear Genomic Sequences Reveal that Polar Bears Are an Old and Distinct Bear Lineage". Science 336 (6079): 344–7. Bibcode:2012Sci...336..344H. doi:10.1126/science.1216424. PMID 22517859. 
  38. ^ Miller, W.; Schuster, S. C.; Welch, A. J.; Ratan, A.; Bedoya-Reina, O. C.; Zhao, F.; Kim, H. L.; Burhans, R. C.; Drautz, D. I.; Wittekindt, N. E.; Tomsho, L. P.; Ibarra-Laclette, E.; Herrera-Estrella, L.; Peacock, E.; Farley, S.; Sage, G. K.; Rode, K.; Obbard, M.; Montiel, R.; Bachmann, L.; Ingolfsson, O.; Aars, J.; Mailund, T.; Wiig, O.; Talbot, S. L.; Lindqvist, C. (2012). "Polar and brown bear genomes reveal ancient admixture and demographic footprints of past climate change". Proceedings of the National Academy of Sciences 109 (36): E2382. Bibcode:2012PNAS..109E2382M. doi:10.1073/pnas.1210506109. PMC 3437856. PMID 22826254. 
  39. ^ a b "La recuperació del rei dels boscos" (in Basque, Catalan, Galician, and Spanish). Retrieved 19 November 2010. 
  40. ^ a b Valdiosera, C. E.; García, N.; Anderung, C.; Dalén, L.; Crégut-Bonnoure, E.; Kahlke, R. D.; Stiller, M.; Brandström, M.; Thomas, M. G. (2007). "Staying out in the cold: Glacial refugia and mitochondrial DNA phylogeography in ancient European brown bears". Molecular Ecology 16 (24): 5140–5148. doi:10.1111/j.1365-294X.2007.03590.x. PMID 18031475. 
  41. ^ Cuántos osos hay y dónde viven. Fundación Oso Pardo – distribution maps and population from 2008 (in Spanish)
  42. ^ DEPANA – detailed distribution maps and census information from 2009 (in Catalan).
  43. ^ The Cantabrian Brown Bear Ursus arctos arctos. picos-accommodation.co.uk.
  44. ^ Heptner, "Sirenia and carnivora (Sea cows; Wolves and Bears)"
  45. ^ a b c d e f g h i j k l m n o p q r s t Carnivores of the World by Dr. Luke Hunter. Princeton University Press (2011), ISBN 9780691152288
  46. ^ a b c Wood, Gerald (1983). The Guinness Book of Animal Facts and Feats. ISBN 978-0-85112-235-9. 
  47. ^ a b ''The living animals of the world; a popular natural history with one thousand illustrations'' Volume 1: Mammals, by Cornish, C. J., 1858–1906; Selous, Frederick Courteney, 1851–1917; Johnston, Harry Hamilton, Sir, 1858–1927; Maxwell, Herbert, Sir, published by New York, Dodd, Mead and Company]. Archive.org. Retrieved 15 September 2011.
  48. ^ "Brown Grizzly or Brown Bear (Ursus arctos)". Mammals of Texas- Online Edition. Retrieved 14 June 2012. 
  49. ^ "Ursus arctos- Brown Bear species". Wildpro. Retrieved 14 June 2012. 
  50. ^ "Polar bear,(Ursus maritimus)" (PDF). U.S. Fish and Wildlife Service. Retrieved 22 March 2008. Appearance.(Overview page) 
  51. ^ a b c "Kodiak Brown Bear- Kodiak- U.S. Fish & Wildlife Service". U.S. Fish & Wildlife Service. Retrieved 2012-06-14. 
  52. ^ (Overview page)
  53. ^ a b LaFee, Scott (2008-05-29). "Seeds of doubt". San Diego Union Tribune. Retrieved 24 May 2015. 
  54. ^ Brown bear zoo.org
  55. ^ "Kodiak Bear Fact Sheet". Alaska Department of Fish and Game, Division of Wildlife Conservation. 2008. Archived from the original on 10 March 2010. Retrieved 27 October 2008. 
  56. ^ a b c d Macdonald, D.W.; Barrett, P. (1993). Mammals of Europe. New Jersey: Princeton University Press. p. 107. ISBN 0-691-09160-9. 
  57. ^ "Ancient bear made early migration". BBC News. 12 November 2004. Retrieved 2 January 2010. 
  58. ^ "Brown Bear. Species". WWF. 
  59. ^ "Brown Bear". Rolling Hills Wildlife Adventure. 
  60. ^ "Neixen quatre cries d'ós bru als Pirineus". elperiodico.cat (in Catalan and Spanish). 2 August 2010. Retrieved 19 November 2010. 
  61. ^ "Situació preocupant de l'ós bru als Pirineus" (in Catalan). ecologistasenaccion.org. May 2010. Retrieved 19 November 2010. 
  62. ^ "Osos, el desafío de una población creciente". ABC (in Spanish). Spain. 
  63. ^ "Proact Local Campaigns: Carpathian Brown Bear". Archived from the original on 24 March 2004. Retrieved 19 November 2010. 
  64. ^ "Carpathian Brown Bear Management Plan [in Poland]". carpathianbear.pl. Archived from the original on 13 August 2011. Retrieved 19 November 2010. 
  65. ^ "Brown Bear – Population & Distribution: A Truly International Species". wwf.panda.org. Retrieved 19 November 2010. 
  66. ^ "Karhu" (in Finnish). suurpedot.fi. Archived from the original on 7 November 2014. Retrieved 22 October 2010. 
  67. ^ Bear Online Information System for Europe. Kora.ch. Retrieved 15 September 2011.
  68. ^ "Brown bears declared extinct in Austria – Again". Wildlife Extra. Retrieved 20 March 2015. 
  69. ^ a b Murtskhvaladze, M.; Gavashelishvili, A.; Tarkhnishvili, D. (2010). "Geographic and genetic boundaries of brown bear (Ursus arctos) population in the Caucasus". Molecular Ecology 19 (9): 1829–1841. doi:10.1111/j.1365-294X.2010.04610.x. PMID 20345670. 
  70. ^ "Brown Bear Hunting in Russia". Russianbearhunt.com. Retrieved 10 October 2009. 
  71. ^ a b Bears in the Mountain National Parks. Grizzly Bears. Parks Canada
  72. ^ "Brown Bear Research in Alaska". Alaska Department of Fish and Game. Retrieved April 2012. 
  73. ^ a b c McLellan, B. (1995). "Maintaining Viability of Brown Bears along the Southern Fringe of Their Distribution.". Proceedings of the 10th international conference on bear research and management 10: 607–611. 
  74. ^ Mattson, David J.; Merrill, Troy (2002). "Extirpations of Grizzly Bears in the Contiguous United States, 1850 –2000". Conservation Biology 16 (4): 1123–1136. doi:10.1046/j.1523-1739.2002.00414.x. 
  75. ^ Gallo-Reynoso, Juan-Pablo; Van Devender, Thomas; Reina-Guerrero, Ana Lilia; Egido-Villarreal, Janitzio; Pfeiler, Edward (2008). "Probable Occurrence of a Brown Bear (Ursus arctos) in Sonora, Mexico,". The Southwestern Naturalist, 2008. 53 (2): 256–260. doi:10.1894/0038-4909(2008)53[256:pooabb]2.0.co;2. Retrieved 19 March 2015. 
  76. ^ Nomura, Fuyuki; Higashi, Seigo (2000). "Effects of food distribution on the habitat usage of a female brown bear Ursus arctos yesoensis in a beech‐forest zone of northernmost Japan". Ecological Research 15 (2): 209–217. doi:10.1046/j.1440-1703.2000.00342.x. 
  77. ^ Hirata, Daisuke; et al. (2013). "Molecular Phylogeography of the Brown Bear (Ursus arctos) in Northeastern Asia Based on Analyses of Complete Mitochondrial DNA Sequences". Mol Biol Evol. 30 (7): 1644–1652. doi:10.1093/molbev/mst077. PMID 23619144. 
  78. ^ Higuma Population Estimates. pref.hokkaido.lg.jp (2 December 2015)
  79. ^ 2015 ANNUAL POPULATION STATUS REPORT FOR BROWN BEARS IN NORTHERN DINARIC MOUNTAINS AND EASTERN ALPS. dinalpbear.eu (May 2015).
  80. ^ Boitani, L.; Cowling, R.M.; Dublin, H.T. (2008). "Change the IUCN Protected Area Categories to Reflect Biodiversity Outcomes.". PLoS Biology 6 (3): 436–438. doi:10.1371/journal.pbio.0060066. 
  81. ^ Farhadinia, Mohammad Sadegh; Valizadegan, Negin (2015). "A preliminary baseline status of the Syrian Brown Bear Ursus arctos syriacus (Mammalia: Carnivora: Ursidae) in Golestanak, northern Iran.". Journal of Threatened Taxa 7 (1): 6796–6799. doi:10.11609/jott.o3708.6796-9. 
  82. ^ Calvignac, Sebastien; Hughes, Sandrine; Hanni, Catherine (2009). "Genetic diversity of endangered brown bear (Ursus arctos) populations at the crossroads of Europe, Asia and Africa". Diversity and Distributions 15 (5): 742–750. doi:10.1111/j.1472-4642.2009.00586.x. 
  83. ^ "Brown / Grizzly Bear Facts". North American Bear Center. 
  84. ^ Egbert, Allan L.; Stokes, Allen W.; Egbert, A. L. (1974). "The social behaviour of brown bears on an Alaskan salmon stream" (PDF). International Conference Bear Res. and Manage 3: 41–56. doi:10.2307/3872753. JSTOR 3872753. 
  85. ^ Stonorov, Derek and Stokes, Allen W. (1972) "Social Behavior of the Alaska Brown Bear" Panel 4: Bear Behaviour
  86. ^ 288回「知床ヒグマ親子 番屋に大集合!」 ダーウィンが来た!生きもの新伝説
  87. ^ Bidon, T.; Janke, A.; Fain, S. R.; Eiken, H. G.; Hagen, S. B.; Saarma, U.; Hallstrom, B. M.; Lecomte, N.; Hailer, F. (2014). "Brown and Polar Bear Y Chromosomes Reveal Extensive Male-Biased Gene Flow within Brother Lineages". Molecular Biology and Evolution 31 (6): 1353–63. doi:10.1093/molbev/msu109. PMID 24667925. 
  88. ^ Types of Bears – Information on Specific Bear Species (2009)
  89. ^ "Brown Bear Reproduction". Shadowofthebear.com. Archived from the original on 1 May 2008. Retrieved 10 October 2009. 
  90. ^ Bellemain, Eva; Swenson, Jon E.; Taberlet, Pierre (2006). "Mating Strategies in Relation to Sexually Selected Infanticide in a Non-Social Carnivore: The Brown Bear". Ethology 112 (3): 238–246. doi:10.1111/j.1439-0310.2006.01152.x. 
  91. ^ "Brown/Grizzly Bear Hunting in Alaska". Alaska Department of Fish and Game. 
  92. ^ a b Seryodkin, I.V. (2007). Роль Бурого Медведя в Экосистемах Дальнего Востока России [Role of the Brown Bear in the Ecosystem of Far East Russia] (in Russian). International Conference in Dnepropetrovsk. pp. 502–503. 
  93. ^ "Alaska is bear territory!". Alaska Office of Economic Development. Dced.state.ak.us. Archived from the original on 15 April 2009. Retrieved 10 October 2009. 
  94. ^ "Yellowstone Grizzly Bears Eat 40,000 Moths a Day In August". Yellowstonepark.com. 21 June 2011. 
  95. ^ a b Gau, R. J., Case, R., Penner, D. F., & McLoughlin, P. D. (2002). "Feeding patterns of barren-ground grizzly bears in the central Canadian Arctic" (PDF). Arctic 55 (4): 339–344. JSTOR 40512491. 
  96. ^ McGrady, M. J., Potapov, E., & Utekhina, I. (1999). "Brown bear (Ursus arctos) feeds on Steller's Sea Eagle (Haliaeetus pelagicus) nestling". Journal of Raptor Research 33 (4): 342–343. 
  97. ^ Brown, Susan, A. "Inherited behavior traits of the domesticated ferret". weaselwords.com. Retrieved 29 January 2010. 
  98. ^ French, S. P.; French, M. G. (1990). "Predatory behavior of grizzly bears feeding on elk calves in Yellowstone National Park, 1986–1988" (PDF). International Conf. Bear Res. and Manage 8: 335–341. doi:10.2307/3872937. JSTOR 3872937. 
  99. ^ Persson, I. L.; Wikan, S.; Swenson, J. E.; Mysterud, I. (2001). "The diet of the brown bear Ursus arctos in the Pasvik Valley, northeastern Norway". Wildlife Biology 7 (1): 27–37. 
  100. ^ Mattson, D. J. (1997). "Use of ungulates by Yellowstone grizzly bears Ursus arctos". Biological Conservation 81 (1): 161–177. doi:10.1016/s0006-3207(96)00142-5. 
  101. ^ Wyman, T. (2002). Grizzly bear predation on a bull bison in Yellowstone National Park. Ursus, 375–377.
  102. ^ Dahle, B.; Sørensen, O. J.; Wedul, E. H.; Swenson, J. E.; Sandegren, F. (1998). "The diet of brown bears Ursus arctos in central Scandinavia: effect of access to free-ranging domestic sheep Ovis aries". Wildlife biology 4 (3): 147–158. 
  103. ^ Prynn, David (2004). Amur tiger. Russian Nature Press. p. 115. 
  104. ^ Frasef, A. (2012). Feline Behaviour and Welfare. CABI. pp. 72–77. ISBN 978-1-84593-926-7. 
  105. ^ Seryodkin; et al. (2003). "Denning ecology of brown bears and Asiatic black bears in the Russian Far East". Ursus 14 (2): 159. 
  106. ^ Seryodkin, I. V.; Goodrich, J. M.; Kostyria, A. V.; Smirnov, E. N.; Miquelle, D. G. (2011). "Intraspecific relationships between brown bears, Asiatic black bears and the Amur tiger". 20th International Conference on Bear Research & Management (PDF). International Association for Bear Research and Management. p. 64. 
  107. ^ Matthiessen, Peter; Hornocker, Maurice (2001). Tigers In The Snow. North Point Press. ISBN 0-86547-596-2. 
  108. ^ Miquelle, D.G., Smirnov, E.N., Goodrich, J.M. (2005). "1". Tigers of Sikhote-Alin Zapovednik: ecology and conservation. Vladivostok, Russia: PSP. 
  109. ^ Geptner, V. G., Sludskii, A. A. (1972). Mlekopitaiuščie Sovetskogo Soiuza. Vysšaia Škola, Moskva. (In Russian; English translation: Heptner, V. G.; Sludskii, A. A.; Bannikov, A. G.; (1992). Mammals of the Soviet Union. Volume II, Part 2: Carnivora (Hyaenas and Cats). Smithsonian Institution and the National Science Foundation, Washington DC). pp. 95–202.
  110. ^ Yudakov, A. G., Nikolaev, I. G. (2004). "Hunting Behavior and Success of the Tigers' Hunts". The Ecology of the Amur Tiger based on Long-Term Winter Observations in 1970–1973 in the Western Sector of the Central Sikhote-Alin Mountains (english translation ed.). Institute of Biology and Soil Science, Far-Eastern Scientific Center, Academy of Sciences of the USSR. 
  111. ^ Seryodkin, I. V., Goodrich, J. M., Kostyrya, A. V., Schleyer, B. O., Smirnov, E. N., Kerley, L. L., & Miquelle, D. G. (2005). "Глава 19. Взаимоотношения амурского тигра с бурым и гималайским медведями [Chapter 19. Relationship of Amur tigers with brown and Himalayan black bear]". In Miquelle, D. G.; Smirnov, E. N.; Goodrich, J. M. Tigers of Sikhote-Alin Zapovednik: Ecology and Conservation (in Russian). Vladivostok, Russia: PSP. pp. 156–163. 
  112. ^ "Brown Bear predation of Amur Tiger 1973 account". International Wildlife Magazine. 
  113. ^ Jimenez, Michael D.; Asher, Valpa J.; Bergman, Carita; Bangs, Edward E. & Woodruff, Susannah P. (2008). "Gray Wolves, Canis lupus, Killed by Cougars, Puma concolor, and a Grizzly Bear, Ursus arctos, in Montana, Alberta, and Wyoming". The Canadian Field-Naturalist 122 (1): 76.  PDF.
  114. ^ Downey, Betsy (2006). "Personal Encounter. Wolf-Grizzly interaction in Yellowstone National Park" (PDF). International Wolf Center. Archived from the original on 27 February 2008. 
  115. ^ Murphy, Kerry M.; Felzien, Gregory S.; Hornocker, Maurice G. & Ruth, Toni K. (1998). "Encounter Competition between Bears and Cougars: Some Ecological Implications". Ursus 10: 55–60. JSTOR 3873109. 
  116. ^ ADW: Ursus arctos: Information. Arlis.org. Retrieved 9 August 2012.
  117. ^ Hornocker, M., and S. Negri (eds.) (2009). Cougar: ecology and conservation. University of Chicago Press. Chicago, IL, ISBN 0226353443.
  118. ^ Sørensen, Ole; Mogens Totsås; Tore Solstad; Robin Rigg (2008). "Predation by a Golden Eagle on a Brown Bear Cub" (PDF). Ursus 19 (2): 190–193. doi:10.2192/08SC008.1. 
  119. ^ Gunther, Kerry A.; Biel, Mark J.; Anderson, Neil & Watts, Lisette (2002). "Probable grizzly bear predation on an American black bear in Yellowstone National Park" (PDF). Ursus 13: 372–374. 
  120. ^ Seryodkin, Ivan V.; Kostyria, A. V.; Goodrich, J. M.; Miquelle, D. G.; Smirnov, E. N.; Kerley, L. L.; Quigley, H. B.; Hornocker, M. G. (2003). "Denning ecology of brown bears and Asiatic black bears in the Russian Far East" (PDF). Ursus 14 (2): 153–161. JSTOR 3873015. 
  121. ^ The Intellectual observer: review of natural history, microscopic research, and recreative science, Groombridge, 1865
  122. ^ Adams, Andrew Leith (1867) Wanderings of a naturalist in India: the western Himalayas, and Cashmere, Edmonston and Douglas
  123. ^ O'Hara, Dough (24 April 2005) Polar bears, grizzlies increasingly gather on North Slope. Anchorage Daily News.
  124. ^ "ABC News: Grizzlies Encroaching on Polar Bear Country". ABC News. Retrieved 10 October 2009. 
  125. ^ "Our Alaska: Grizzly vs. moose". Alaska Dispatch. 17 July 2011. Archived from the original on 3 December 2013. 
  126. ^ "Moose Moms Prefer Traffic to Grizzly Bears, Study Says". National Geographic. Retrieved 12 October 2007. 
  127. ^ "Relocation". Get Bear Smart Society. [dead link]
  128. ^ Folklore and Legends of the North American Indian compiled by Joshua B Lippincott, published by Abela Publishing Ltd, 2009, ISBN 0-9560584-6-9
  129. ^ Averkieva, Julia P. and Sherman, Mark A. (1992) Kwakiutl String Figures UBC Press, ISBN 0-7748-0432-7
  130. ^ Herrero, Stephen (2002) Bear Attacks: Their Causes and Avoidance, revised edition, Lyons Press, ISBN 158574557X.
  131. ^ a b Smith, Tom S. & Herrero, Steven. "Ursus arctos californicus". Alaska Science Center – Biological Science Office. Archived from the original on 14 August 2009. Retrieved 12 April 2008. 
  132. ^ Why are grizzly bears more aggressive than our black bears?. Digital Collegian. 27 April 2004
  133. ^ Rogers, Lynn L. How Dangerous are Black Bears. bear.org
  134. ^ a b c d Herrero, Stepehen (1985) Bear Attacks: Their Causes and Avoidance, Hurtig Publishers Ltd./ Edmonton
  135. ^ Medred, Craig (26 July 2011) Alaska bear attack: NOLS kids did a 'phenomenal job', Alaska Dispatch
  136. ^ Bears besiege Russian mine after killing guards. The Times. 24 July 2008
  137. ^ Dinets, Vladimir. "Brown Bears of Russia". Archived from the original on 15 December 2012. Retrieved 10 January 2016. 
  138. ^ "Brown Bear (Ursus arctos))". Tooth & Claw. Archived from the original on 16 May 2008. Retrieved 5 January 2008. 
  139. ^ Bear Caused Human Injuries and Deaths In Yellowstone National Park. Yellowstone-bearman.com (1 January 2000). Retrieved 2011-09-15.
  140. ^ Smith, Herrero, DeBruyn, Wilde (2008). "Spray more effective than guns against bears: study". North American Bear Center. Archived from the original on 1 January 2011. Retrieved 28 March 2009. 
  141. ^ Smith, Tom S. "Brown Bear Projects at the Alaska Science Center". Alaska Science Center – Biological Science Office. Archived from the original on 14 August 2009. Retrieved 2 June 2008. 
  142. ^ "Alaska State Troopers Press Release of Monday, November 19, 2007" (Case Number: 07-96958). Alaska Department of Public Safety. 19 November 2007. Retrieved 2 June 2008. [dead link]
  143. ^ "Safety: bears and you". State of Alaska Department of Natural Resources. 
  144. ^ "Ursus arctos californicus Merriam, 1896". itis.gov. Retrieved 1 April 2008. 
  145. ^ "History and Culture – State Symbols". California State Library. Retrieved 23 September 2011. 

Bibliography[edit]

  • Heptner V.G. & Sludskii, A.A. (1992). Mammals of the Soviet Union, Volume II, Part 2. Leiden u.a.: Brill. ISBN 90-04-08876-8. 

External links[edit]