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Wikipedia

Brown bear

January 01, 1970

This article is about the animal. For the athletics teams at Brown University, see Brown Bears. For the research ship, see MV Brown Bear.

The brown bear (Ursus arctos) is a large bear species found across Eurasia and North America. It is one of the largest living terrestrial members of the order Carnivora, rivaled in size only by its closest relative, the polar bear, which is much less variable in size and slightly bigger on average. Adults of different subspecies range in weight from 80 to 600 kg (180 to 1,320 lb), with males being heavier than females. Despite its name, brown bears aren't entirely brown; the pelage can be reddish to yellowish-brown, and dark brown to cream in color. During winter, brown bears in some populations hibernate and emerge during spring to regain up to 180 kg (400 lb) of weight. They have well developed dentition and claws, ideal for their lifestyle.

Brown bear
Temporal range: 0.5–0 Ma
↓
Middle Pleistocene-Holocene
Kodiak bear on Kodiak Island
CITES Appendix II (CITES)[2][note 1]
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Ursidae
Genus: Ursus
Species:
U. arctos
Binomial name
Ursus arctos
Linnaeus, 1758
Subspecies

15, see text and article

Brown bear range map

The brown bear is mostly found in forested habitats in elevations of up to 5,000 m (16,000 ft). It is omnivorous, and consumes a variety of plant and animal species; with the former comprising 90% of its diet. The bear hunts animals as small as rodents, to animals as large as moose or muskoxen. In parts of coastal Alaska, brown bears predominately feed on spawning salmon that come ashore to lay their eggs. The brown bear is a solitary animal, except in the breeding season. Females protect their young for an average of 1.5 to 4.5 years. Brown bears have one of the largest skulls of any land-based carnivore, and are able to make use of tools. They are long lived animals, with an average lifespan of 25 years in the wild. Attacks on humans, though reported, are generally rare.

While the brown bear's range has shrunk, and it has faced local extinctions across its wide range, it remains listed as a least concern species by the International Union for Conservation of Nature (IUCN) with a total estimated population in 2017 of 110,000. Populations that were hunted to extinction in the 19th and 20th centuries are the Atlas bear of North Africa and the Californian, Ungavan and Mexican populations of the grizzly bear of North America. Many of the populations in the southern parts of Eurasia are highly endangered as well. One of the smaller-bodied forms, the Himalayan brown bear, is critically endangered, occupying only 2% of its former range and threatened by uncontrolled poaching for its body parts. The Marsican brown bear of central Italy is one of several currently isolated populations of the Eurasian brown bear and is believed to have a population of just c. 50 bears.

Etymology

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).[3][4] In the mid-19th century United States, the brown bear was termed "Old Ephraim" and sometimes as "Moccasin Joe".[5]

The scientific name of the brown bear, Ursus arctos, comes from the Latin ursus, meaning "bear",[6] and the Greek ἄρκτος/arktos, also meaning "bear".[7]

Evolution and taxonomy

Evolution

The brown bear is one of eight extant species in the bear family Ursidae and of six extant species in the subfamily Ursinae.

A possible phylogeny based on complete mitochondrial DNA sequences from Yu et al. (2007).[8]
Ursidae

Giant panda (Ailuropoda melanoleuca)  

Spectacled bear (Tremarctos ornatus)  

Ursinae

Sloth bear (Melursus ursinus)  

Sun bear (Helarctos malayanus)  

Asian black bear (Ursus thibetanus)  

American black bear (Ursus americanus)  

Polar bear (Ursus maritimus)  

Brown bear (Ursus arctos)  

The polar bear and the brown bear form a close grouping, while the relationships of the other species are not very well resolved.[9]
A more recent phylogeny based on the genetic study of Kumar et al. (2017).[10]
Ursidae

Giant panda (Ailuropoda melanoleuca)  

Spectacled bear (Tremarctos ornatus)  

Ursinae

Sloth bear (Melursus ursinus)  

Sun bear (Helarctos malayanus)  

Asian black bear (Ursus thibetanus)  

American black bear (Ursus americanus)  

Polar bear (Ursus maritimus)  

Brown bear (Ursus arctos)  

The study concludes that Ursine bears originated around 5 million years ago and show extensive hybridization of species in their lineage.[10]


During the Pleistocene, brown bears were sympatric with extinct Ursinae species in North America.[11][12] Brown bears are thought to have evolved from Ursus etruscus in Asia.[13][14] The brown bear, per Kurten (1976), has been stated as "clearly derived from the Asian population of Ursus savini about 800,000 years ago; spread into Europe, to the New World."[15] A genetic analysis indicated that the brown bear lineage diverged from the cave bear species complex approximately 1.2–1.4 million years ago, but did not clarify if U. savini persisted as a paraspecies for the brown bear before perishing.[16] The oldest fossils positively identified as from this species occur in China from about 0.5 million years ago.[13][17] In Yakutia, brown bears are known to have been present from at least 400,000 to 300,000 years ago.[18] Brown bears entered Europe about 250,000 years ago and North Africa shortly after.[13] Brown bears were present in the Zagros Mountains by the Late Pleistocene.[19] Brown bear remains from the Pleistocene period are common in the British Isles, where amongst other factors, they may have contributed to the extinction of cave bears (Ursus spelaeus).[20]

Brown bears (along with lions, bison and red foxes) first emigrated to North America from Eurasia via Beringia during the Illinoian Glaciation.[21] Although Björn Kurtén hypothesized skull shape as an indicator of evolutionary history,[20] genetic evidence suggests several different populations of brown bear migrated into North America, aligning with the glacial cycles of the Pleistocene. The founding population of most North American brown bears arrived first, with the genetic lineage developing around ~177,000 BP, with fossils being first recovered ~111,000 BP from East Beringia. Genetic divergences suggest brown bears first migrated south during MIS-5 (~92,000–83,000 BP) upon the opening of the ice-free corridor,[21][22] with the first fossils being near Edmonton (26,000 BP).[12] The Alexander and Haida Gwaii archipelagoes have a separate lineage, which first appears around 20,000 BP. After a local extinction in Beringia ~33,000 BP, two new but closely related lineages repopulated Alaska and northern Canada from Eurasia after the Last Glacial Maximum (>25,000 BP).[21]

Brown bear fossils discovered in Ontario, Ohio, Kentucky and Labrador show that the species occurred farther east than indicated in historic records.[13] In North America, two types of the subspecies Ursus arctos horribilis are generally recognized—the coastal brown bear and the inland grizzly bear.[23]

Subspecies

Main article: Subspecies of brown bear
 
Adult female Eurasian brown bear, the nominate subspecies

Brown bear taxonomy and subspecies classification has been described as "formidable and confusing," with few authorities listing the same specific set of subspecies.[24] There are hundreds of obsolete brown bear subspecies, each with its own name, so this can become confusing. As many as 90 subspecies have been proposed.[25][26] However, recent DNA analysis has identified as few as five main clades which contain all extant brown bears,[27][28] while a 2017 phylogenetic study revealed nine clades, including one representing polar bears.[29] As of 2005, 15 extant or recently extinct subspecies were recognized by the general scientific community.[30][31]

As well as the exact number of overall brown bear subspecies, its precise relationship to the polar bear also remains in debate. The polar bear is a recent offshoot of the brown bear. The point at which the polar bear diverged from the brown bear is unclear, with estimations based on genetics and fossils ranging from 400,000 to 70,000 years ago, but most recent analysis has indicated that the polar bear split somewhere between 275,000 and 150,000 years ago.[32] Under some definitions, the brown bear can be construed as the paraspecies for the polar bear.[33]

DNA analysis shows that, apart from recent human-caused population fragmentation,[34] brown bears in North America are generally part of a single interconnected population system, with the exception of the population (or subspecies) in the Kodiak Archipelago, which has probably been isolated since the end of the last Ice Age.[35][36] These data demonstrate that U. a. gyas, U. a. horribilis, U. a. sitkensis and U. a. stikeenensis are not distinct or cohesive groups, and would more accurately be described as ecotypes. For example, brown bears in any particular region of the Alaska coast are more closely related to adjacent grizzly bears than to distant populations of brown bears,[37] the morphological distinction seemingly driven by brown bears having access to a rich salmon food source, while grizzly bears live at higher elevation, or further from the coast, where plant material is the base of the diet. The history of the bears of the Alexander Archipelago is unusual in that these island populations carry polar bear DNA, presumably originating from a population of polar bears that was left behind at the end of the Pleistocene, but have since been connected with adjacent mainland populations through movement of males, to the point where their nuclear genomes are now more than 90% of brown bear ancestry.[38] MtDNA analysis revealed that brown bears are apparently divided into five different clades, some of which coexist or co-occur in different regions.[39]

Hybrids

See also: Grizzly–black bear hybrid and Grizzly–polar bear hybrid
 
Possible grizzly-black bear hybrid in the Yukon Territory, Canada

A grizzly–polar bear hybrid is a rare ursid hybrid resulting from a crossbreeding 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, and seven more hybrids have since been confirmed in the same region, all descended from a single female polar bear.[40] 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).[41]

Analyses of the genomes of bears have shown that introgression between species was widespread during the evolution of the genus Ursus,[42] including the introgression of polar bear DNA introduced to brown bears during the Pleistocene.[43]

Description

 
Brown bears are highly variable in size. Eurasian brown bears often fall around the middle to low sizes for the species.

The brown bear is the most variable in size of modern bears. The typical size depends upon which population it is from, and most accepted subtypes vary widely in size. This is in part due to sexual dimorphism, as male brown bears average at least 30% larger in most subtypes. Individual bears vary in size seasonally, weighing the least in spring due to lack of foraging during hibernation, and the most in late fall, after a period of hyperphagia to put on additional weight to prepare for hibernation. Therefore, a bear may need to be weighed in both spring and fall to get an idea of its mean annual weight.[44][45]

Generally, brown bears weigh anywhere from 80 to 600 kg (180 to 1,320 lb), with males outweighing females.[46] The normal range of physical dimensions for a brown bear is a head-and-body length of 1.4 to 2.8 m (4 ft 7 in to 9 ft 2 in) and a shoulder height of 70 to 153 cm (2 ft 4 in to 5 ft 0 in). The tail is relatively short, as in all bears, ranging from 6 to 22 cm (2.4 to 8.7 in) in length.[47][48] The smallest brown bears, females during spring among barren-ground populations, can weigh so little as to roughly match the body mass of males of the smallest living bear species, the sun bear (Helarctos malayanus), while the largest coastal populations attain sizes broadly similar to those of the largest living bear species, the polar bear.[49] Interior brown bears are generally smaller than is often perceived, being around the same weight as an average lion, at an estimate average of 180 kg (400 lb) in males and 135 kg (298 lb) in females, whereas adults of the coastal populations weigh about twice as much.[50] The average weight of adult male bears from 19 populations, from around the world and various subspecies (including both large- and small-bodied subspecies), was found to be 217 kg (478 lb) while adult females from 24 populations were found to average 152 kg (335 lb).[23][51]

Coloration

 
A brown bear in Slovenia

Brown bears are often not fully brown.[52] They have long, thick fur, with a moderately long mane at the back of the neck which varies somewhat across the types.[53] In India, brown bears can be reddish with silver-tipped hairs, while in China brown bears are bicolored, with a yellowish-brown or whitish collar across the neck, chest and shoulders.[52][54] Even within well-defined subspecies, individuals may show highly variable hues of brown. North American grizzlies can be dark brown (almost black) to cream (almost white) or yellowish-brown and often have darker-colored legs. The common name "grizzly" stems from their typical coloration, with the hairs on their back usually being brownish-black at the base and whitish-cream at the tips, giving them their distinctive "grizzled" color. Apart from the cinnamon subspecies of the American black bear (U. americanus cinnamonum), the brown bear is the only modern bear species to typically appear truly brown.[55] 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, with its length and density varying geographically.[56]

Cranial morphology and size

 
Brown bear skeleton
 
Skull

Adults have massive, heavily built concave skulls, which are large in proportion to the body.[55] The projections of the skull are well developed.[56] Skull lengths of Russian brown 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.[56] 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.[56] The teeth of brown bears reflect their dietary plasticity and are broadly similar to other bears.[57][58] The teeth are reliably larger than American black bears, but average smaller in molar length than polar bears.[59] Brown bears have the broadest skull of any extant ursine bear.[50]

Claws and feet

 
Front paws

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 may measure 7 to 10 centimetres (2.8 to 3.9 in) along the curve.[56] They are generally dark with a light tip, with some forms having completely light claws.[56] Brown bear claws are longer and straighter than those of American black bears (Ursus americanus).[55] 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 typically climb trees as well as black bears, although in rare cases adult female brown bears have been seen scaling trees.[60] The claws of a polar bear are quite different, being notably shorter but broader with a strong curve and sharper point.[17][61] The paws of the brown bear are quite large. The rear feet of adult bears measure 21 to 36 cm (8.3 to 14.2 in) long, while the forefeet tend to measure about 40% less in length.[62] Brown bears are the only extant bears with a hump at the top of their shoulder, which is made entirely of muscle, this feature having developed presumably for imparting more force in digging, which is habitual during foraging for the bear and also used heavily in den construction prior to hibernation.[55] The brown bear's strength has been roughly estimated as 2.5 to 5 times that of a human.[63]

Distribution and habitat

Main article: Distribution of brown bears
 
Brown bear at Brooks Falls

Brown bears were once native to Europe, much of Asia, the Atlas Mountains of Africa, and North America,[64] but are now extirpated in some areas, and their populations have greatly decreased in other areas. There are approximately 200,000 brown bears left in the world.[65] The largest populations are in Russia with 130,000,[66] the United States with 32,500, and Canada with around 25,000. Brown bears live in Alaska, east through the Yukon and Northwest Territories, south through British Columbia and through the western half of Alberta. The Alaskan population is estimated at a healthy 30,000 individuals.[67] In the lower 48 states, they are repopulating slowly, but steadily along the Rockies and the western Great Plains.[68]

In Europe, in 2010, there were 14,000 brown bears in ten fragmented populations, from Spain (estimated at only 20–25 animals in the Pyrenees in 2010,[69] in a range shared between Spain, France and Andorra, and some 210 animals in Asturias, Cantabria, Galicia and León, in the Picos de Europa and adjacent areas in 2013[70]) in the west, to Russia in the east, and from Sweden and Finland in the north to Romania (5000–6000), Bulgaria (900–1200), Slovakia (with about 600–800 animals), Slovenia (500–700 animals) and Greece (with Karamanlidis et al. 2015 estimating >450 animals) in the south.[71][72]

In Asia, brown bears are found primarily throughout Russia, thence more spottily southwest to parts of the Middle East, including almost all parts of Kurdistan, to as far south as southwestern Iran, and to the southeast in Northeast China. Brown bears are also found in Western China, Kyrgyzstan, North Korea, Pakistan, Afghanistan and India. A population of brown bear can be found on the Japanese island of Hokkaidō, which holds the largest number of non-Russian brown bears in eastern Asia with about 2,000–3,000 animals.[62]

This species inhabits the broadest range of habitats of any living bear species.[62] They seem to have no altitudinal preferences and have been recorded from sea level to an elevation of 5,000 m (16,000 ft) in the Himalayas.[62] In most of their range, brown bears generally seem 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.[62]

Conservation status

 
A Marsican brown bear, with a range restricted to the Abruzzo, Lazio and Molise National Park, Italy

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.[1][39] However, the California grizzly bear, Ungava brown bear, Atlas bear and Mexican grizzly bear, as well as brown bear populations in the Pacific Northwest, were hunted to extinction in the 19th and early 20th centuries and many of the southern Asian subspecies are highly endangered.[73] The Syrian brown bear (U. a. syriacus) is very rare and it has been extirpated from more than half of its historic range.[74] One of the smallest-bodied subspecies, the Himalayan brown bear (U. a. isabellinus), is critically endangered, occupying only 2% of its former range and threatened by uncontrolled poaching for its body parts.[75] The Marsican brown bear in central Italy is believed to have a population of just 50 bears.[76]

Behavior and life history

 
Like all bears, brown bears can stand on their hindlegs and walk for a few steps in this position, usually motivated to do so by curiosity, hunger or alarm

The brown bear is often described as nocturnal. However, it frequently seems to peak in activity in the morning and early evening hours.[77] Activity can occur at nearly any time of night or day, with bears who dwell in areas with more extensive human contact being more likely to be fully nocturnal. Furthermore, yearling and newly independent bears are more likely to be active diurnally and many adult bears in low-disturbance areas are largely crepuscular.[78][79][80] In summer through autumn, a brown bear can double its weight from the spring, gaining up to 180 kg (400 lb) of fat, on which it relies to make it through winter, when it becomes very lethargic.[81][82] Although they are not full hibernators and can be woken easily, both sexes like to den in a protected spot during the winter months. Hibernation dens may consist of any spot that provides cover from the elements and that can accommodate their bodies, such as a cave, crevice, cavernous tree roots, or hollow logs.[83][84]

Brown bears have one of the largest brains of any extant carnivoran relative to their body size and have been shown to engage in tool use, which requires advanced cognitive abilities.[85] This species is mostly solitary, although bears may gather in large numbers at major food sources (e.g., open garbage dumps or rivers holding spawning salmon) and form social hierarchies based on age and size.[86][87] Adult male bears are particularly aggressive and are avoided by adolescent and subadult males, both at concentrated feeding opportunities and chance encounters. Female bears with cubs rival adult males in aggression and are much 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.[88][89] 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.[86] During combat, bears use their paws to strike their opponents in the chest or shoulders and bite the head or neck.[90]

Communication

 
Brown bear sounds
Brown bear sounds including huffing, jaw popping and growling
Brown bear roars
Brown bears roaring over a carcass
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Several different facial expressions have been documented in brown bears. The "relaxed-face" is made in everyday activities and has the ears pointed to the sides and the mouth closed or slackly open. During social play, bears make "relaxed open-mouth face" in which the mouth is open, with a curled upper lip and hanging lower lip, and the ears alert and shifting. When looking at another animal at a distance, the bear makes an "alert face" as the ears are cocked and alert, the eyes wide open but the mouth is closed or only open slightly. The "tense closed mouth face" is made with the ears laid back and the mouth closed and occurs when the bear feels threatened. When approached by another individual, the animal makes a "puckered-lip face" with a protruding upper lip and ears which go from cocked and alert when at a certain distance to laid back when closer or when retreating. The "jaw gape face" consists of an open mouth with visible lower canines and hanging lips while the "biting face" is similar to the "relaxed open-mouth face" except the ears are flattened and the eyes are wide enough to expose the sclera. Both the "jaw gape face" and the "biting face" are made in aggression and bears switch between them.[90]

Brown bears also produce various vocalizations. Huffing occurs when the animal is tense while woofing is made when alarmed. Both sounds are produced by exhalations though huffing is harsher and is made continuously (two per second). Growls and roars are made in aggression. Growling is "harsh" and "guttural" and can range from a simple grrr to a rumble. A rumbling growl can escalate into a roar when the bear is charging. Roaring is described as "thunderous" and can travel 2 km (1.2 mi). Mothers and cubs wanting physical contact will bawl, which is heard as waugh!, waugh!.[90]

Home ranges

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.[23][89] Males always cover more area than females each year. Despite their lack of traditional territorial behavior, adult males can seem to have a "personal zone" in which other bears are not tolerated if they are seen.[91] Males always wander further than females, due to both increasing access to females and food sources, while females are advantaged by smaller territories in part since it decreases the likelihood of encounters with male bears who may endanger their cubs.[23][92] 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 and 318 km2 (44 and 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 Romania, the largest home range was recorded for adult males (3,143 km2; 1,214 sq mi).[93] 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.[94][89]

Reproduction

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

The mating season is from mid-May to early July, shifting later the further north the bears are found.[95] Being serially monogamous, brown bears remain with the same mate from a couple of days to a couple of weeks.[23] Outside of this narrow time frame, adult male and female brown bears show no sexual interest in each other.[23] Females mature sexually between the age of four and eight years of age, with an average age at sexual maturity of 5.2–5.5 years old,[96] 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.[91][97] Males will try to mate with as many females as they can; usually a successful one mates with two females in a span of one to three weeks.[48][97] The adult female brown bear is similarly promiscuous, mating with up to four, rarely even eight, males while in heat and potentially breeding with two males in a single day.[98] Females come into oestrus on average every three to four years, with a full range of 2.4 to 5.7 years. The urine markings of a female in oestrus can attract several males via scent.[99] Dominant males may try to sequester a female for her entire oestrus period of approximately two weeks, but usually are unable to retain her for the entire time.[50][92] Copulation is vigorous and prolonged and can last up to an hour, although the mean time is about 23–24 minutes.[50][100]

 
Grizzly bear cubs often imitate their mothers closely.

Males take no part in raising their cubs – parenting is left entirely to the females.[89][101] 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 while gestating, the embryo does not implant and is reabsorbed into the body.[96][102][103] There have been cases of brown bears with as many as six cubs, although the average litter size is one to three, with more than four being considered uncommon.[104] There are records of females sometimes adopting stray cubs or even trading or kidnapping cubs when they emerge from hibernation (a larger female may claim cubs away from a smaller one).[62][105][106] Older and larger females within a population tend to give birth to larger litters.[107] The size of a litter also depends on factors such as geographic location and food supply.[108] At birth, the cubs are blind, toothless and hairless and may weigh from 350 to 510 g (0.77 to 1.12 lb), again reportedly based on the age and condition of the mother. 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 over long distances and begin to forage for solid food.[23][109]

 
Kodiak bear cubs play-fighting

The cubs are fully dependent on the mother and a close bond is formed. During the dependency stage, the cubs learn (rather than inherit as instincts from birth) 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.[94] Increased brain size in large carnivores has been positively linked to whether a given species is solitary, as is the brown bear, or raises their offspring communally, thus female brown bears have relatively large, well-developed brains, presumably key in teaching behavior.[110] The cubs learn by following and imitating their mother's actions during the period they are with her.[62] Cubs remain with their mother for an average of 2.5 years in North America, uncommonly being independent as early as 1.5 years of age or as late as 4.5 years of age.[50] The stage at which independence is attained may generally be earlier in some parts of Eurasia, as the latest date which mother and cubs were together was 2.3 years, most families separated in under two years in a study from Hokkaido and in Sweden most cubs on their own were still yearlings.[111][112] Brown bears practice infanticide, as an adult male bear may kill the cubs of another.[92] When an adult male brown bear kills a cub, it is usually because he is trying to bring the female into oestrus, as she will enter that state within two to four days after the death of her cubs.[92] Cubs may flee up a tree when they see a strange male bear approaching. The mother often successfully defends them, even though the male may be twice as heavy as her. However, females have been known to die in such confrontations.[92][113][114]

Dietary habits

Main article: Dietary biology of the brown bear
 
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.[62] Despite their reputation, most brown bears are not highly carnivorous, as they derive up to 90% of their dietary food energy from vegetable matter.[115] They often feed on a variety of plant life, including berries, grasses, flowers, acorns and pine cones, as well as fungi such as mushrooms.[23] Among all bears, brown bears are uniquely equipped to dig for tough foods such as roots, bulbs[116] and shoots. They use their long, strong claws to dig out earth to reach the roots and their powerful jaws to bite through them.[23] In spring, winter-provided carrion, grasses, shoots, sedges, moss[116] and forbs are the dietary mainstays for brown bears internationally.[62] 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.[62] 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 in a single day, and may derive up to half of their annual food energy from these insects.[117] 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.[55] This species may eat birds and their eggs, including almost entirely ground- or rock-nesting species.[55] The diet may be supplemented by rodents or similar smallish mammals, including marmots, ground squirrels, mice, rats, lemmings and voles.[23] With particular regularity, bears in Denali National Park will wait at burrows of Arctic ground squirrels hoping to pick off a few of the 1 kg (2.2 lb) rodents.[118]

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 nutrious 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.[55]

 
A cow moose with calves being approached by an inland brown bear, Denali National Park, Alaska

Beyond the regular predation of salmon, most brown bears are not particularly active predators.[55] 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.[55] 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.[55] Large mammals preyed on can include various ungulate species such as elk, moose, caribou, muskoxen and wild boar.[23] 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.[119] It will also bite or swipe some prey to stun it enough to knock it over for consumption.[120] 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 ambush young animals by finding them via scent.[23] 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.[120] 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.[23] 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.[55] They feed on carrion, and use their size to intimidate other predators, such as wolves, cougars, tigers, and American black bears from their kills. Carrion is especially important in the early spring (when the bears are emerging from hibernation), much of it comprising winter-killed big game.[23] Cannibalism is not unheard of, though predation is not normally believed to be the primary motivation when brown bears attack each other.[55]

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 are sometimes exploited as prey. Cattle are bitten on the neck, back or head and then the abdominal cavity is opened for eating.[23] Plants and fruit farmed by humans are readily consumed as well, including corn, wheat, sorghum, melons and any form of berries.[55] They may feed on domestic bee yards, readily consuming both honey and the brood (grubs and pupae) of the honey bee colony.[55] Human foods and trash are 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.[55]

Interspecific predatory relationships

 
Brown bear being followed by a wolf

Adult bears are generally immune to predatory attacks except from large Siberian (Amur) tigers and other bears. Following a decrease of ungulate populations from 1944 to 1959, 32 cases of Siberian tigers attacking both Ussuri brown bears (Ursus arctos lasiotus) and Ussuri black bears (U. thibetanus ussuricus) were recorded in the Russian Far East, and hair of bears were found in several tiger scat samples. Tigers attack black bears less often than brown bears, since the black bears live in more open habitats and are not able to climb trees. In the same time period, four cases of brown bears killing female tigers and young cubs were reported, both in disputes over prey and in self-defense. Tigers mainly feed on the bear's fat deposits, such as the back, hams and groin.[56]

In rare cases, when Amur tigers prey on brown bears, they usually target young and sub-adult bears, besides small female adults taken outside their dens, generally when lethargic from hibernation.[121] Predation by tigers on denned brown bears was not detected during a study carried between 1993 and 2002.[122] Ussuri brown bears, along with the smaller black bears constitute 2.1% of the Siberian tiger's annual diet, of which 1.4% are brown bears.[123][124]

The effect the presence of tigers have on a brown bears behavior seems to vary. In the winters of 1970–1973, Yudakov and Nikolaev recorded two cases of bears showing no fear of tigers and another case of a brown bear changing path upon crossing tiger tracks.[125] Other researchers have observed bears following tiger tracks to scavenge tiger kills and to potentially prey on tigers.[56][123] Despite the threat of predation, some brown bears actually benefit from the presence of tigers by appropriating tiger kills that the bears may not be able to successfully hunt themselves.[123] During telemetry research in the Sikhote-Alin Nature Reserve, 44 direct confrontations between bears and tigers were observed, in which bears (not just brown bears) in general were killed in 22 cases, and tigers in 12 cases.[126] There are reports of brown bears specifically targeting Amur leopards and tigers 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.[127] Some studies show that bears frequently track down tigers to usurp their kills, with occasional fatal outcomes for the tiger. A report from 1973 describes twelve known cases of brown bears killing tigers, including adult males. In all cases, the tigers were subsequently eaten by the bears.[128][129]

Brown bears regularly intimidate wolves to drive them away from their kills. In Yellowstone National Park, 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 recorded case of fully-grown wolves being killed by a grizzly bear.[130] Given the opportunity, however, both species will prey on the other's cubs.[131]

In some areas, grizzly bears regularly displace cougars from their kills.[132] 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.[133][134] Smaller carnivorous animals, including coyotes, wolverines, lynxes, and any other sympatric carnivores or raptorial birds, are dominated by grizzly 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.[55] There is one record of a golden eagle preying on a brown bear cub.[135]

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 grizzly bears in open, unforested areas. Although displacement of black bears by grizzly bears has been documented, actual interspecific killing of black bears by grizzlies 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 grizzly's largely nocturnal habits and preference for open spaces.[136] 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.[137]

As of the 21st century, there has been an increase in interactions between brown bears and polar bears, theorized to be caused by climate change. Brown and grizzly bears have been seen moving increasingly northward into territories formerly claimed by polar bears. They tend to dominate polar bears in disputes over carcasses,[138] and dead polar bear cubs have been found in brown bear dens.[139]

Longevity and mortality

 
Front paw imprint
 
Rear paw imprint

The brown bear has a naturally long life. Wild females have been observed reproducing up to 28 years of age, which is the oldest known age for reproduction of any ursid in the wild. The peak reproductive age for females ranges from four to 20 years old.[62][140] The lifespan of brown bears of both sexes within minimally hunted populations is estimated at an average of 25 years. The oldest wild brown bear on record was nearly 37 years old.[141] The oldest recorded female in captivity was nearly 40 years old, while males in captivity have been verified to live up to 47 years, with one captive male possibly attaining 50 years of age.[23][48]

While male bears potentially live longer in captivity, female grizzly bears have a greater annual survival rate than males within wild populations per a study done in the Greater Yellowstone Ecosystem.[142] Annual mortality for bears of any age is estimated at 10% in most protected areas.[62] Around 13% to 44% of cubs die within their first year even in well-protected areas.[23] Beyond predation by large predators including wolves, Siberian tigers and other brown bears, starvation and accidents also claim the lives of cubs. Studies have indicated that the most prevalent source of mortality for first-year cubs is malnutrition.[23]

Brown bears are susceptible to parasites such as flukes, ticks, tapeworms, roundworms and biting lices.[143][144] It is thought that brown bears may catch canine distemper virus (CDV) from other Caniforms like stray dogs and wolves.[145] A captive individual allegedly succumbed to Aujeszky's disease.[146]

Hibernation physiology

Scientists are interested in understanding the hibernation physiology of brown bears, specifically in how they stay healthy after six months of inactivity.[147] It has been suggested by scientists that understanding how brown bears stay healthy during hibernation may potentially help lead to cures for human diseases such as diabetes or cardiovascular disease.[147] A study conducted through the Scandinavian Brown Bear Research Project did a proteomic analysis of the brown bear's blood, organs, and tissues to pinpoint proteins and peptides that either increased or decreased in expression in the winter and summer months.[147] One major finding was that the plasma protein sex hormone-binding globulin (SHBG) increased during the brown bear's hibernation period by 45 times.[147] Although scientists do not yet understand the role of SHBG in maintaining the brown bear's health, some scientists believe these findings could potentially be a factor in preventing human diseases that come from a sedentary life style.[147]

Relationship with humans

Attacks on humans

Main article: Bear attack
 
A statue of the Ussuri brown bear from Hokkaido, Japan

Brown bears usually avoid areas where extensive development or urbanization has occurred.[148] They rarely attack humans on sight and usually avoid people.[149] They are, however, unpredictable in temperament, and may attack if they are surprised or feel threatened.[150] The two most common causes for bear attacks are surprise and curiosity.[151] Mothers defending cubs are the most prone to attacking, being responsible for 70% of brown bear-caused human fatalities in North America.[152] Brown bear attacks tend to result in serious injury and, in some cases, death.[150] Due to the bears' enormous physical strength, even a single bite or swipe can be deadly.[153] Violent encounters with brown bears usually last only a few minutes, though they can be prolonged if the victims fight back.[150] A study conducted in 2019 found that 664 bear attacks were reported during a 15-year period (2000–2015) throughout North America and Eurasia. There were 568 injuries and 95 casualties.[154] Around 10 people a year are killed by brown bears in Russia, more than all the other parts of the brown bear's range combined.[155] In Japan, a large brown bear nicknamed "Kesagake" ("kesa-style slasher") caused the worst brown bear attack in Japanese history at Tomamae, Hokkaidō during numerous encounters during December 1915. It killed seven people and wounded three others before being gunned down after a large-scale beast-hunt.[156] A study by U.S. and Canadian researchers has found bear spray to be more effective at stopping aggressive bear behavior than guns, working in 92% of studied incidents versus 67% for guns.[157]

Bear hunting

Main article: Bear hunting

Hunting of brown bears has occurred since ancient times, mainly for sports and royal showings of power. Humans have been recorded hunting brown bears for over 10,300–9,300 years.[158] In Europe, between the 17th and 18th centuries, humans sought to control brown bear numbers by awarding those who managed to kill one. This bounty scheme pushed the brown bear population to the brink of extinction before comprehensive protection was offered in the 1900s. In northern Europe, hunting still persists in the 21st century, contributing significantly to the drop in brown bear numbers.[159] In some places in Europe, bears are tricked into entering traps with the help of a bait.[154] Brown bears are legally hunted in some American states, such as Alaska. However, a hunting license is required and killings of females with cubs will result in a prison sentence.[160]

Their meat is sometimes consumed and used in recipes such as dumplings, hams and stews. The Indigenous people of James Bay (Eastern) Cree use their flesh in traditional dishes. In Asia and Romania, the paws are consumed as exotic delectables; it has been a prevalent component of traditional Chinese food since 500 BC. The total weight of commercially sold brown bear meat is estimated at 17 tons annually.[161]

In captivity

There are more than 700 brown bears in zoos and wildlife parks worldwide. Captive bears are largely lethargic and spend a considerable amount of time doing nothing. When active, captive bears may be subject to repetitive back and forth motion, known widely as pacing. This behavior is most prevalent in bears kept in small, cramped cages often with no natural setting. Pacing is a way of coping with stress that comes with being trapped in unnaturally small spaces.[162] Stereotyped behavior has decreased due to better and larger enclosures being built, and more sustainable management from zoo staff.[163]

They are also exploited as dancing bears. The process begins at an early age. Cubs, for example, are positioned on hot metal plates, causing them to exhibit dancing movements, and at the same time, violin music is running in the background. The process is repeated, resulting in bears being trained to dance when a violin is played. Similarly, brown bears are also displayed in tiny enclosures near a restaurant, mainly for the purpose of luring customers. Privately owned bears are often placed in insufficient environments and often suffer from malnutrition and obesity.[164] According to a 2009 analysis, the brown bear was the second most exploited circus animal after the tiger.[165]

Culture

Further information: Cultural depictions of bears
 
"The Story of the Three Bears", illustration from Childhood's Favorites and Fairy Stories
 
Cover of an American humor magazine Puck at 20 January 1904

Brown bears often figure into the literature of Europe and North America as "cute and cuddly", 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.[166]

In Ancient Greek mythology, bears have been compared to humans as similar, mainly due to their ability of standing upright. In many western stories and older fables the portrayed attributes of bears are sluggishness, foolishness and gullibility, which contradicts actual species. For example, bears have been reported tricking hunters by backtracking in the snow.[167] The Romans built small carved figures of bears that were used during burials of infants.[168] The earliest cave paintings of bears occurred in the Paleolithic, with over 100 recorded paintings.[169]

The Russian bear is a common national personification for Russia (as well as the former Soviet Union), despite the country having no officially-designated national animal. The brown bear is Finland's national animal.[170][171] The grizzly bear is the state animal of Montana.[172] The California golden bear is the state animal of California, despite being extinct.[173] In North America, the brown bear is considered a charismatic megafauna and has long piqued people's interest. The death of Bear 148 at the hands of a trophy hunter in 2017, sparked media outrage and the continued disapproval of trophy hunting.[174]

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.[175][176] The brown bear is depicted on the reverse of the Croatian 5 kuna coin, minted since 1993.[177]

References

  1. ^ a b McLellan, B.N.; Proctor, M.F.; Huber, D.; Michel, S. (2017). "Ursus arctos". IUCN Red List of Threatened Species. 2017: e.T41688A121229971. doi:10.2305/IUCN.UK.2017-3.RLTS.T41688A121229971.en. Retrieved 19 November 2021.
  2. ^ "Appendices | CITES". cites.org. from the original on 5 December 2017. Retrieved 14 January 2022.
  3. ^ Harper, Douglas. "bruin". Online Etymology Dictionary.
  4. ^ Bruin 20 September 2022 at the Wayback Machine, Encyclopaedia Britannica
  5. ^ Hunting the Grisly and other Sketches 6 September 2017 at the Wayback Machine. FullTextArchive.com. Retrieved 15 September 2011.
  6. ^ Liddell, Henry George and Scott, Robert. "Ursus." 1 March 2021 at the Wayback Machine A Latin Dictionary. Perseus Digital Library.
  7. ^ Liddell, Henry George and Scott, Robert."Arktos." 30 June 2023 at the Wayback Machine A Greek-English Lexicon. Perseus Digital Library.
  8. ^ Yu, Li; Li, Yi-Wei; Ryder, Oliver A.; Zhang, Ya-Ping (2007). "Analysis of complete mitochondrial genome sequences increases phylogenetic resolution of bears (Ursidae), a mammalian family that experienced rapid speciation". BMC Evolutionary Biology. 7 (198): 198. Bibcode:2007BMCEE...7..198Y. doi:10.1186/1471-2148-7-198. PMC 2151078. PMID 17956639.
  9. ^ Servheen, C.; Herrero, S.; Peyton, B. (1999). Bears: Status Survey and Conservation Action Plan (PDF). IUCN. pp. 26–30. ISBN 978-2-8317-0462-3. Archived (PDF) from the original on 9 October 2022.
  10. ^ a b Kumar, V.; Lammers, F.; Bidon, T.; Pfenninger, M.; Kolter, L.; Nilsson, M. A.; Janke, A. (2017). "The evolutionary history of bears is characterized by gene flow across species". Scientific Reports. 7: 46487. Bibcode:2017NatSR...746487K. doi:10.1038/srep46487. PMC 5395953. PMID 28422140.
  11. ^ Kumar, V.; Lammers, F.; Bidon, T.; Pfenninger, M.; Kolter, L.; Nilsson, M. A.; Janke, A. (2017). "The evolutionary history of bears is characterized by gene flow across species". Scientific Reports. 7: 46487. Bibcode:2017NatSR...746487K. doi:10.1038/srep46487. PMC 5395953. PMID 28422140.
  12. ^ a b Steffen, Martina L.; Fulton, Tara L. (1 February 2018). "On the association of giant short-faced bear (Arctodus simus) and brown bear (Ursus arctos) in late Pleistocene North America". Geobios. 51 (1): 61–74. Bibcode:2018Geobi..51...61S. doi:10.1016/j.geobios.2017.12.001.
  13. ^ a b c d McLellan, Bruce; Reiner, David C. (1994). "A Review of bear evolution" (PDF). Int. Conf. Bear Res. And Manage. 9 (1): 85–96. doi:10.2307/3872687. JSTOR 3872687. Archived (PDF) from the original on 9 October 2022.
  14. ^ Pérez-Hidalgo, T. (1992). "The European descendants of Ursus etruscus C. Cuvier (Mammalia, Carnivora, Ursidae)" (PDF). Boletín del Instituto Geológico y Minero de España. 103 (4): 632–642. Archived (PDF) from the original on 9 October 2022.
  15. ^ Kurten, Bjorn (1976). The Cave Bear Story. New York: Columbia University Press. p. 5.
  16. ^ Loreille, O.; Orlando, L.; Patou-Mathis, M.; Philippe, M.; Taberlet, P.; Hänni, C. (2001). "Ancient DNA analysis reveals divergence of the cave bear, Ursus spelaeus, and brown bear, Ursus arctos, lineages". Current Biology. 11 (3): 200–203. Bibcode:2001CBio...11..200L. doi:10.1016/s0960-9822(01)00046-x. PMID 11231157. S2CID 14645603.
  17. ^ a b Herrero, S. (1972). "Aspects of evolution and adaptation in American black bears (Ursus americanus Pallas) and brown and grizzly bears (U. arctos Linne.) of North America" (PDF). Bears: Their Biology and Management. 2: 221–231. doi:10.2307/3872586. JSTOR 3872586. Archived (PDF) from the original on 9 October 2022.
  18. ^ Kosintsev, Pavel A.; Bocherens, Hervé; Kirillova, Irina V.; Levchenko, Vladimir A.; Zazovskaya, Elya P.; Trofimova, Svetlana S.; Lan, Tianying; Lindqvist, Charlotte (5 December 2024). "Palaeoecological and genetic analyses of Late Pleistocene bears in Asiatic Russia". Boreas. 51 (2): 465–480. doi:10.1111/bor.12570. ISSN 0300-9483. Retrieved 4 May 2024 – via Wiley Online Library.
  19. ^ Monchot, Hervé; Mashkour, Marjan; Biglari, Fereidoun; Abdi, Kamyar (April–June 2020). "The Upper Pleistocene brown bear (Carnivora, Ursidae) in the Zagros: Evidence from Wezmeh Cave, Kermanshah, Iran". Annales de Paléontologie. 106 (2): 102381. doi:10.1016/j.annpal.2019.102381. Retrieved 9 May 2024 – via Elsevier Science Direct.
  20. ^ a b McLellan, Bruce; Reiner, David C. (1994). "A Review of bear evolution" (PDF). Int. Conf. Bear Res. And Manage. 9 (1): 85–96. doi:10.2307/3872687. JSTOR 3872687. Archived (PDF) from the original on 9 October 2022.
  21. ^ a b c Salis, Alexander T.; Bray, Sarah C. E.; Lee, Michael S. Y.; Heiniger, Holly; Barnett, Ross; Burns, James A.; Doronichev, Vladimir; Fedje, Daryl; Golovanova, Liubov; Harington, C. Richard; Hockett, Bryan; Kosintsev, Pavel; Lai, Xulong; Mackie, Quentin; Vasiliev, Sergei (December 2022). "Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge". Molecular Ecology. 31 (24): 6407–6421. Bibcode:2022MolEc..31.6407S. doi:10.1111/mec.16267. hdl:11343/299180. ISSN 0962-1083. PMID 34748674.
  22. ^ Kubiak, Cara; Grimes, Vaughan; Van Biesen, Geert; Keddie, Grant; Buckley, Mike; Macdonald, Reba; Richards, M. P. (27 June 2022). "Dietary niche separation of three Late Pleistocene bear species from Vancouver Island, on the Pacific Northwest Coast of North America". Journal of Quaternary Science. 38: 8–20. doi:10.1002/jqs.3451. ISSN 0267-8179. S2CID 250134103. from the original on 31 October 2023. Retrieved 2 January 2024.
  23. ^ a b c d e f g h i j k l m n o p q r (PDF). American Society of Mammalogists, Smith College. 23 April 1993. Archived from the original (PDF) on 31 March 2017.
  24. ^ Wilson, D. E.; Ruff, S. (1999), The Smithsonian Book of North American Mammals, Washington, D.C.: Smithsonian Institution Press, pp. 160–201
  25. ^ Harris, Arthur H. (2013). "Pleistocene Vertebrates of Arizona, New Mexico, and Trans-Pecos Texas". UTEP Biodiversity Collections, University of Texas at El Paso.
  26. ^ Storer, T.I.; Tevis, L.P. (1996). California Grizzly. Berkeley, CA: University of California Press. pp. 335, 42–187. ISBN 978-0-520-20520-8. from the original on 2 February 2024. Retrieved 18 November 2019. Alt URL
  27. ^ U.S. Fish and Wildlife Service (17 November 2006). (PDF). Federal Register. 70 (221): 69854–69884. Archived from the original (PDF) on 25 August 2006.
  28. ^ 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" (PDF). Molecular Ecology. 17 (8): 1962–1970. Bibcode:2008MolEc..17.1962C. doi:10.1111/j.1365-294x.2008.03631.x. PMID 18363668. S2CID 23361337. Archived (PDF) from the original on 9 October 2022.
  29. ^ Lan, T.; Gill, S.; Bellemain, E.; Bischof, R.; Zawaz, M. A.; Lindqvist, C. (2017). "Evolutionary history of enigmatic bears in the Tibetan Plateau–Himalaya region and the identity of the yeti". Proceedings of the Royal Society B: Biological Sciences. 284 (1, 868): 20,171,804. doi:10.1098/rspb.2017.1804. PMC 5740279. PMID 29187630.
  30. ^ Wozencraft, W. C. (2005). "Ursus arctos". In Wilson, D. E.; Reeder, D. M. (eds.). 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.
  31. ^ Ursus arctos 22 January 2019 at the Wayback Machine, ITIS
  32. ^ Lindqvist, C.; Schuster, S. C.; Sun, Y.; et al. (2010). "Complete mitochondrial genome of a Pleistocene jawbone unveils the origin of polar bear". Proceedings of the National Academy of Sciences. 107 (11): 5053–5057. Bibcode:2010PNAS..107.5053L. doi:10.1073/pnas.0914266107. PMC 2841953. PMID 20194737.
  33. ^ Liu, Shiping; Lorenzen, Eline D.; Fumagalli, Matteo; Li, Bo; Harris, Kelley; Xiong, Zijun; Zhou, Long; Korneliussen, Thorfinn Sand; Somel, Mehmet; Babbitt, Courtney; Wray, Greg; Li, Jianwen; He, Weiming; Wang, Zhuo; Fu, Wenjing; Xiang, Xueyan; Morgan, Claire C.; Doherty, Aoife; o'Connell, Mary J.; McInerney, James O.; Born, Erik W.; Dalén, Love; Dietz, Rune; Orlando, Ludovic; Sonne, Christian; Zhang, Guojie; Nielsen, Rasmus; Willerslev, Eske; Wang, Jun (2014). "Population Genomics Reveal Recent Speciation and Rapid Evolutionary Adaptation in Polar Bears". Cell. 157 (4): 785–794. doi:10.1016/j.cell.2014.03.054. PMC 4089990. PMID 24813606.
  34. ^ Proctor, Michael F.; Paetkau, David; Mclellan, Bruce N.; Stenhouse, Gordon B.; Kendall, Katherine C.; Mace, Richard D.; Kasworm, Wayne F.; Servheen, Christopher; Lausen, Cori L. (2012). "Population fragmentation and inter-ecosystem movements of grizzly bears in western Canada and the northern United States". Wildlife Monographs. 180 (1): 1–46. Bibcode:2012WildM.180....1P. doi:10.1002/wmon.6. ISSN 1938-5455. S2CID 16790669.
  35. ^ Strobeck, Curtis; Craighead, Lance; Clarkson, Peter L.; Waits, Lisette P.; Paetkau, David (1 December 1997). "An Empirical Evaluation of Genetic Distance Statistics Using Microsatellite Data From Bear (Ursidae) Populations". Genetics. 147 (4): 1943–1957. doi:10.1093/genetics/147.4.1943. ISSN 0016-6731. PMC 1208359. PMID 9409849. from the original on 29 January 2019. Retrieved 29 January 2019.
  36. ^ Waits, L. P.; et al. (1998). "Mitochondrial DNA Phylogeography of the North American Brown Bear and Implications for Conservation". Conservation. 12 (2): 408–417. Bibcode:1998ConBi..12..408W. doi:10.1111/j.1523-1739.1998.96351.x. S2CID 86172292.
  37. ^ Paetkau, D.; Shields, G. F.; Strobeck, C. (1998). "Gene flow between insular, coastal and interior populations of brown bears in Alaska". Molecular Ecology. 7 (#10): 1283–1292. Bibcode:1998MolEc...7.1283P. doi:10.1046/j.1365-294x.1998.00440.x. ISSN 0962-1083. PMID 9787441. S2CID 21848010.
  38. ^ Shapiro, Beth; Slatkin, Montgomery; Stirling, Ian; John, John St.; Salamzade, Rauf; Ovsyanikov, Nikita; Jay, Flora; Stiller, Mathias; Fulton, Tara L. (14 March 2013). "Genomic Evidence for Island Population Conversion Resolves Conflicting Theories of Polar Bear Evolution". PLOS Genetics. 9 (3): e1003345. doi:10.1371/journal.pgen.1003345. ISSN 1553-7404. PMC 3597504. PMID 23516372.
  39. ^ a b Servheen, C.; Herrero, S.; Peyton, B.; Pelletier, K.; Moll, K.; Moll, J., eds. (1999), (PDF), vol. 44, Gland, Switzerland: IUCN, archived from the original (PDF) on 23 September 2015, retrieved 18 November 2019
  40. ^ Richardson, Evan; Branigan, Marsha; Paetkau, David; Pongracz, Jodie D. (31 May 2017). "Recent Hybridization between a Polar Bear and Grizzly Bears in the Canadian Arctic". Arctic. 70 (#2): 151–160. doi:10.14430/arctic4643. ISSN 1923-1245.
  41. ^ Janke, Axel; Nilsson, Maria A.; Kolter, Lydia; Pfenninger, Markus; Bidon, Tobias; Lammers, Fritjof; Kumar, Vikas (19 April 2017). "The evolutionary history of bears is characterized by gene flow across species". Scientific Reports. 7: 46,487. Bibcode:2017NatSR...746487K. doi:10.1038/srep46487. ISSN 2045-2322. PMC 5395953. PMID 28422140.
  42. ^ 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 (6, 079): 344–247. Bibcode:2012Sci...336..344H. doi:10.1126/science.1216424. hdl:10261/58578. PMID 22517859. S2CID 12671275.
  43. ^ 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): E2382–2390. Bibcode:2012PNAS..109E2382M. doi:10.1073/pnas.1210506109. PMC 3437856. PMID 22826254.
  44. ^ Hissa, R.; Siekkinen, J.; Hohtola, E.; Saarela, S.; Hakala, A.; Pudas, J. (1994). "Seasonal patterns in the physiology of the European brown bear (Ursus arctos arctos) in Finland". Comparative Biochemistry and Physiology Part A: Physiology. 109 (#3): 781–791. doi:10.1016/0300-9629(94)90222-4. PMID 8529017. from the original on 15 June 2020. Retrieved 12 December 2016.
  45. ^ McLellan, B. N. (2011). "Implications of a high-energy and low-protein diet on the body composition, fitness, and competitive abilities of black (Ursus americanus) and grizzly (Ursus arctos) bears". Canadian Journal of Zoology. 89 (#6): 546–558. doi:10.1139/z11-026.
  46. ^ Maria Pasitschniak-Arts (23 April 1993). "Ursus arctos" (PDF). Mammalian Species (439): 1–10. doi:10.2307/3504138. JSTOR 3504138. (PDF) from the original on 5 March 2023. Retrieved 5 March 2023.
  47. ^ Grzimek, Bernhard (1990). Grzimek's Encyclopedia of Mammals. McGraw-Hill. pp. 52–53. ISBN 978-0-07-909508-4.
  48. ^ a b c Nowak, Ronald M. (1991). Walker's Mammals of the World. Johns Hopkins University Press. pp. 1083–1093. ISBN 978-0-8018-3970-2.
  49. ^ Christiansen, P. (1999). "What size were Arctodus simus and Ursus spelaeus (Carnivora: Ursidae)?". Annales Zoologici Fennici. 36 (2): 93–102. JSTOR 23735739.
  50. ^ a b c d e Feldhamer, G. A.; Thompson, B. C.; Chapman, J. A. (2003). Wild mammals of North America: biology, management, and conservation. JHU Press. ISBN 9780801874161. from the original on 2 February 2024. Retrieved 9 November 2020.
  51. ^ Swenson, Jon E.; Adamič, Miha; Huber, Djuro; Stokke, Sigbjørn (2007). "Brown bear body mass and growth in northern and southern Europe" (PDF). Oecologia. 153 (#1): 37–47. Bibcode:2007Oecol.153...37S. doi:10.1007/s00442-007-0715-1. PMID 17415593. S2CID 5836340. Archived (PDF) from the original on 9 October 2022.
  52. ^ a b Jones, S. V. (1923). "Color variations in wild animals". Journal of Mammalogy. 4 (#3): 172–177. doi:10.2307/1373567. JSTOR 1373567.
  53. ^ Swenson, J. E. (2000). Action plan for the conservation of the brown bear in Europe (Ursus arctos) 20 December 2016 at the Wayback Machine (No. 18-114). Council of Europe.
  54. ^ Sahajpal, V; Goyal, S. P.; Jayapal, R; Yoganand, K; Thakar, M. K. (2008). "Hair characteristics of four Indian bear species". Science & Justice. 48 (#1): 8–15. doi:10.1016/j.scijus.2007.05.001. PMID 18450212.
  55. ^ a b c d e f g h i j k l m n o p 'Brown, Gary (1993). "Bear Anatomy and Physiology". The great bear almanac. Internet Archive. New York : Lyons & Burford. ISBN 978-1-55821-210-7.
  56. ^ a b c d e f g h V. G. Heptner; N. P. Naumov, eds. (1998). Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola [Mammals of the Soviet Union, Volume II, Part 1a, Sirenia and Carnivora (Sea cows; Wolves and Bears)]. Vol. II, Part 1a. Washington, D.C., US: Science Publishers, Inc. pp. 601–676. ISBN 1-886106-81-9.
  57. ^ Christiansen, Per (2008). "Feeding ecology and morphology of the upper canines in bears (carnivora: Ursidae)". Journal of Morphology. 269 (#7): 896–908. doi:10.1002/jmor.10643. PMID 18488989. S2CID 33532210.
  58. ^ Kurtén, B. (1966). "Pleistocene bears of North America: Genus Tremarctos, spectacled bears". Acta Zoologica Fennica. 115: 1–96.
  59. ^ Sacco, Tyson; Van Valkenburgh, Blaire (2004). "Ecomorphological indicators of feeding behaviour in the bears (Carnivora: Ursidae)". Journal of Zoology. 263: 41. doi:10.1017/S0952836904004856.
  60. ^ Seryodkin, Ivan V. (1 June 2016). "Behavior of Brown Bears During Feeding in the Sikhote-Alin". Achievements in the Life Sciences. 10 (1): 38–47. doi:10.1016/j.als.2016.05.003. ISSN 2078-1520.
  61. ^ Stirling, Ian (1988). Polar Bears. University of Michigan Press. pp. 20–25. ISBN 978-0-472-10100-9.
  62. ^ a b c d e f g h i j k l Hunter, Luke (2011). Carnivores of the world. Internet Archive. Princeton, NJ : Princeton University Press. pp. 122–133. ISBN 978-0-691-15228-8.
  63. ^ Ellig, Tracy (3 July 2006). "MSU researcher tests grizzly bear strength for National Geographic documentary". Montana State University. from the original on 1 June 2013. Retrieved 11 June 2021.
  64. ^ "Ancient bear made early migration". BBC News. 12 November 2004. from the original on 2 December 2005. Retrieved 2 January 2010.
  65. ^ "Brown Bear | Species | WWF". WWF. from the original on 29 September 2018. Retrieved 2 December 2013.
  66. ^ Chestin, Igor E.; Gubar, Yuliy P.; Sokolov, Vladimir E.; Lobachev, Vladimir S. (1992). "The brown bear (Ursus arctos L.) in the USSR: numbers, hunting and systematics". Annales Zoologici Fennici. 29 (2): 57–68. ISSN 0003-455X. JSTOR 23735466.
  67. ^ "Brown Bear Research in Alaska". Alaska Department of Fish and Game. from the original on 3 May 2012. Retrieved 4 April 2012.
  68. ^ "Neixen quatre cries d'ós bru als Pirineus". elperiodico.cat (in Catalan and Spanish). 2 August 2010. from the original on 24 July 2011. Retrieved 19 November 2010.
  69. ^ "Situació preocupant de l'ós bru als Pirineus" (in Catalan). ecologistasenaccion.org. May 2010. from the original on 13 November 2012. Retrieved 19 November 2010.
  70. ^ "Osos, el desafío de una población creciente". ABC (in Spanish). Spain. 28 September 2013. from the original on 26 October 2013. Retrieved 14 October 2013.
  71. ^ Martínez-Abraín, A.; Jiménez, J.; Oro, D. (2018). "Pax Romana: 'refuge abandonment' and spread of fearless behavior in a reconciling world". Animal Conservation. 22 (1). John Wiley & Sons, Inc (Zoological Society of London (ZSL)): 3–13. doi:10.1111/acv.12429. ISSN 1367-9430. S2CID 89938098.
  72. ^ "Bear". ΑΡΚΤΟΥΡΟΣ. from the original on 2 February 2024. Retrieved 13 October 2022.
  73. ^ 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. Bibcode:2006MolEc..15.4477M. doi:10.1111/j.1365-294x.2006.03097.x. PMID 17107477. S2CID 7336900.
  74. ^ 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. Bibcode:2009DivDi..15..742C. doi:10.1111/j.1472-4642.2009.00586.x. S2CID 21666120.
  75. ^ Himalayan brown bears now critically endangered 16 October 2018 at the Wayback Machine. Euronews.com. 6 January 2014
  76. ^ Gervasi, V.; Ciucci, P. (1 January 2018). "Demographic projections of the Apennine brown bear population Ursus arctos marsicanus (Mammalia: Ursidae) under alternative management scenarios". The European Zoological Journal. 85 (1): 242–252. doi:10.1080/24750263.2018.1478003. hdl:11573/1114254. ISSN 2475-0263.
  77. ^ Ordiz, Andrés; Kindberg, Jonas; Sæbø, Solve; Swenson, Jon E.; Støen, Ole-Gunnar (1 May 2014). "Brown bear circadian behavior reveals human environmental encroachment". Biological Conservation. 173: 1–9. Bibcode:2014BCons.173....1O. doi:10.1016/j.biocon.2014.03.006. ISSN 0006-3207.
  78. ^ Klinka, D. R.; Reimchen, T. E. (2002). "Nocturnal and diurnal foraging behaviour of brown bears (Ursus arctos) on a salmon stream in coastal British Columbia" (PDF). Canadian Journal of Zoology. 80 (#8): 1317−1322. doi:10.1139/z02-123. Archived (PDF) from the original on 9 October 2022.
  79. ^ Moe, T. F.; Kindberg, J.; Jansson, I.; Swenson, J. E. (2007). "Importance of diel behaviour when studying habitat selection: examples from female Scandinavian brown bears (Ursus arctos)". Canadian Journal of Zoology. 85 (#4): 518−525. doi:10.1139/Z07-034. from the original on 15 June 2020. Retrieved 12 December 2016.
  80. ^ Kaczensky, P.; Huber, D.; Knauer, F.; Roth, H.; Wagner, A.; Kusak, J. (2006). "Activity patterns of brown bears (Ursus arctos) in Slovenia and Croatia". Journal of Zoology. 269 (#4): 474−485. doi:10.1111/j.1469-7998.2006.00114.x.
  81. ^ Kingsley, M. C. S.; Nagy, J. A.; Russell, R. H. (1983). "Patterns of weight gain and loss for grizzly bears in northern Canada". Bears: Their Biology and Management. 5: 174–178. doi:10.2307/3872535. JSTOR 3872535. S2CID 90555276.
  82. ^ Hissa, R.; Hohtola, E.; Tuomala-Saramäki, T.; Laine, T. (1998). "Seasonal changes in fatty acids and leptin contents in the plasma of the European brown bear (Ursus arctos arctos)". Annales Zoologici Fennici. 35 (#4): 215–224. JSTOR 23735612.
  83. ^ Farley, S. D.; Robbins, C. T. (1995). "Lactation, hibernation, and mass dynamics of American black bears and grizzly bears". Canadian Journal of Zoology. 73 (#12): 2216−2222. doi:10.1139/z95-262.
  84. ^ Evans, A. L.; Sahlén, V.; Støen, O. G.; Fahlman, Å.; Brunberg, S.; Madslien, K.; Forbert, O.; Swenson, J.E.; Arnemo, J. M. (2012). "Capture, anesthesia, and disturbance of free-ranging brown bears (Ursus arctos) during hibernation". PLOS ONE. 7 (#7): e40520. Bibcode:2012PLoSO...740520E. doi:10.1371/journal.pone.0040520. PMC 3398017. PMID 22815757.
  85. ^ Deecke, V. B. (2012). "Tool-use in the brown bear (Ursus arctos)" (PDF). Animal Cognition. 15 (#4): 725–730. doi:10.1007/s10071-012-0475-0. PMID 22367156. S2CID 9076161. Archived (PDF) from the original on 9 October 2022.
  86. ^ a b Egbert, Allan L.; Stokes, Allen W.; Egbert, A. L. (16 September 1976). (PDF). Bears: Their Biology and Management. 3: 41–56. doi:10.2307/3872753. JSTOR 3872753. Archived from the original (PDF) on 17 July 2015. Retrieved 29 April 2011.
  87. ^ Sandell, M. (1989). "The mating tactics and spacing patterns of solitary carnivores". Carnivore behavior, ecology, and evolution. Springer. pp. 164–182. doi:10.1007/978-1-4613-0855-3_7. ISBN 978-1-4613-0855-3.
  88. ^ Gau, R. J.; McLoughlin, P. D.; Case, R.; Cluff, H. D.; Mulders, R.; Messier, F. (2004). "Movements of subadult male grizzly bears, Ursus arctos, in the central Canadian arctic". The Canadian Field-Naturalist. 118 (#2): 239–242. doi:10.22621/cfn.v118i2.920.
  89. ^ a b c d Dahle, B.; Swenson, J. E. (2003). "Seasonal range size in relation to reproductive strategies in brown bears Ursus arctos". Journal of Animal Ecology. 72 (#4): 660–667. Bibcode:2003JAnEc..72..660D. doi:10.1046/j.1365-2656.2003.00737.x. JSTOR 3505643. PMID 30893970. S2CID 67818528.
  90. ^ a b c Egbert, A. L. (1978). The Social Behavior of Brown Bears at McNeil River, Alaska (Ph.D. thesis). Utah State University. pp. 30–35. from the original on 2 February 2024. Retrieved 25 April 2021.
  91. ^ a b Pearson, A. M. (1975). The northern interior grizzly bear Ursus arctos L. Information Canada. p. 86
  92. ^ a b c d e Bellemain, Eva; Swenson, Jon E.; Taberlet, Pierre (2006). "Mating Strategies in Relation to Sexually Selected Infanticide in a Non-Social Carnivore: The Brown Bear" (PDF). Ethology. 112 (#3): 238–246. Bibcode:2006Ethol.112..238B. doi:10.1111/j.1439-0310.2006.01152.x. Archived (PDF) from the original on 9 October 2022.
  93. ^ Pop, Ioan Mihai; Bereczky, Leonardo; Chiriac, Silviu; Iosif, Ruben; Nita, Andreea; Popescu, Viorel Dan; Rozylowicz, Laurențiu (2018). "Movement ecology of brown bears (Ursus arctos) in the Romanian Eastern Carpathians". Nature Conservation. 26: 15–31. doi:10.3897/natureconservation.26.22955. ISSN 1314-3301.
  94. ^ a b Novak, M., Baker, J.A., Obbard, M.E. & Malloch, B. (1987). Wild furbearer management and conservation in North America. Ontario Ministry of Natural Resources.
  95. ^ Herrero, S.; Hamer, D. (1977). "Courtship and copulation of a pair of grizzly bears, with comments on reproductive plasticity and strategy". Journal of Mammalogy. 58 (#3): 441–444. doi:10.2307/1379352. JSTOR 1379352.
  96. ^ a b Craighead, J. J., Sumner, J. S., & Mitchell, J. A. (1995). "The grizzly bears of Yellowstone: their ecology in the Yellowstone ecosystem, 1959–1992". Island Press. pp. 21–56
  97. ^ a b White, D.J.; Berardinelli, J.G.; Aune, K.E. (1998). "Reproductive characteristics of the male grizzly bear in the continental United States". Ursus. 10: 497–501. JSTOR 3873161.
  98. ^ Ambarlı, H. (2016). "Litter size and basic diet of brown bears (Ursus arctos, Carnivora) in northeastern Turkey". Mammalia. 80 (2). doi:10.1515/mammalia-2014-0111. S2CID 87968464.
  99. ^ Halloran, D. W.; Pearson, A. M. (1972). "Blood chemistry of the brown bear (Ursus arctos) from southwestern Yukon Territory, Canada". Canadian Journal of Zoology. 50 (#6): 827–833. doi:10.1139/z72-112. PMID 5038730.
  100. ^ Craighead, J. J.; Hornocker, M. G.; Craighead Jr, F. C. (1969). "Reproductive biology of young female grizzly bears". J. Reprod. Fertil. Suppl. 6: 447–475.
  101. ^ Corbet, Gordon Barclay (1966). The Terrestrial Mammals of Western Europe. London, UK: Foulis. pp. 52–55.
  102. ^ Tsubota, T. & Kanagawa, H. (1993). "Morphological Characteristics of the Ovary, Uterus and Embryo during the Delayed Implantation Period in the Hokkaido Brown Bear (Ursus arctos yesoensis)". Journal of Reproduction and Development. 39 (#4): 325–331. doi:10.1262/jrd.39.325. S2CID 84359252.
  103. ^ Hensel, R. J.; Troyer, W. A. & Erickson, A. W. (1969). "Reproduction in the female brown bear". The Journal of Wildlife Management. 33 (#2): 357–365. doi:10.2307/3799836. JSTOR 3799836.
  104. ^ Pazetnov, V.A. & Pazetnov, S.V. (2005). "Female brown bear with six cubs" (PDF). International Bear News. 14 (#2): 17. Archived (PDF) from the original on 9 October 2022.
  105. ^ Erickson, A. W.; Miller, L. H. (1963). "Cub adoption in the brown bear". Journal of Mammalogy. 44 (#4): 584–585. doi:10.2307/1377153. JSTOR 1377153.
  106. ^ Barnes Jr, V.; Smith, R. (1993). "Cub adoption by brown bears (Ursus arctos middendorffi) on Kodiak Island, Alaska". Canadian Field-Naturalist. 107 (3): 365–367. doi:10.5962/p.357155. from the original on 20 December 2016. Retrieved 12 December 2016.
  107. ^ Stringham, S. F. (1990). "Grizzly bear reproductive rate relative to body size" (PDF). Bears: Their Biology and Management. 8: 433–443. doi:10.2307/3872948. JSTOR 3872948. Archived (PDF) from the original on 9 October 2022.
  108. ^ Steyaert, S. M.; Endrestol, A.; Hacklaender, K.; Swenson, J. E.; Zedrosser, A. (2012). "The mating system of the brown bear Ursus arctos". Mammal Review. 42 (#1): 12–34. doi:10.1111/j.1365-2907.2011.00184.x.
  109. ^ Dahle, B.; Zedrosser, A.; Swenson, J. E. (2006). "Correlates with body size and mass in yearling brown bears (Ursus arctos)". Journal of Zoology. 269 (#3): 273–283. doi:10.1111/j.1469-7998.2006.00127.x.
  110. ^ Gittleman, J. L. (1994). "Female brain size and parental care in carnivores". Proceedings of the National Academy of Sciences. 91 (#12): 5495–5497. Bibcode:1994PNAS...91.5495G. doi:10.1073/pnas.91.12.5495. PMC 44022. PMID 8202515.
  111. ^ Mano, T.; Tsubota, T. (2002). "Reproductive characteristics of brown bears on the Oshima peninsula, Hokkaido, Japan". Journal of Mammalogy. 83 (#4): 1026–1034. doi:10.1644/1545-1542(2002)083<1026:RCOBBO>2.0.CO;2. JSTOR 1383508.
  112. ^ Dahle, B.; Swenson, J. E. (2003). "Family breakup in brown bears: are young forced to leave?". Journal of Mammalogy. 84 (#2): 536–540. doi:10.1644/1545-1542(2003)084<0536:FBIBBA>2.0.CO;2.
  113. ^ Swenson, J. E.; Dahle, B.; Sandegren, F. (2001). "Intraspecific predation in Scandinavian brown bears older than cubs-of-the-year". Ursus. 12: 81–91. JSTOR 3873233. from the original on 16 November 2017. Retrieved 12 December 2016.
  114. ^ Mörner, T.; Eriksson, H.; Bröjer, C.; Nilsson, K.; Uhlhorn, H.; Ågren, E.; Segerstad, C.H.; Jansson, D.S.; Gavier-Widén, D. (2005). "Diseases and mortality in free-ranging brown bear (Ursus arctos), gray wolf (Canis lupus), and wolverine (Gulo gulo) in Sweden". Journal of Wildlife Diseases. 41 (#2): 298–303. doi:10.7589/0090-3558-41.2.298. PMID 16107663. S2CID 43774546.
  115. ^ . Alaska Office of Economic Development. Dced.state.ak.us. Archived from the original on 15 April 2009. Retrieved 10 October 2009.
  116. ^ a b "Ursus arctos (Brown bear)". Animal Diversity Web. from the original on 13 May 2021. Retrieved 28 May 2021.
  117. ^ . Yellowstonepark.com. 21 June 2011. Archived from the original on 15 July 2010.
  118. ^ Brown, Susan, A (17 January 2010). "Inherited behavior traits of the domesticated ferret". weaselwords.com. from the original on 4 October 2009. Retrieved 29 January 2010.{{cite web}}: CS1 maint: multiple names: authors list (link)
  119. ^ 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. Archived (PDF) from the original on 9 October 2022.
  120. ^ a b Macdonald, D.W.; Barrett, P. (1993). Mammals of Europe. New Jersey: Princeton University Press. p. 107. ISBN 978-0-691-09160-0.
  121. ^ Fraser, A. F. (2012). Feline Behaviour and Welfare. CABI. pp. 72–77. ISBN 978-1-84593-926-7.
  122. ^ Seryodkin, I. V.; Kostyria, A. V.; Goodrich, J. M.; Miquelle, D. G.; Smirnov, E. N.; Kerley, L. L. & Hornocker, M. G. (2003). . Ursus. 14 (2): 159. Archived from the original on 17 August 2011. Retrieved 15 September 2014.
  123. ^ a b c 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. (eds.). Tigers of Sikhote-Alin Zapovednik: Ecology and Conservation (in Russian). Vladivostok, Russia: PSP. pp. 156–163. from the original on 18 October 2020. Retrieved 19 March 2023.
  124. ^ Seryodkin, I. (2006). (PhD) (in Russian). Vladivostok, Russia: Far Eastern National University. pp. 61–73. Archived from the original on 24 December 2013.
  125. ^ Yudakov, A. G. & Nikolaev, I. G. (2004). . 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. Institute of Biology and Soil Science, Far-Eastern Scientific Center, Academy of Sciences of the USSR. Archived from the original on 19 February 2020. Retrieved 17 September 2014.
  126. ^ 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" (PDF). 20th International Conference on Bear Research & Management. International Association for Bear Research and Management. p. 64. (PDF) from the original on 13 November 2018. Retrieved 11 September 2014.
  127. ^ Seryodkin, I. V. (2007). . Биоразнообразие и роль животных в экосистемах: Материалы IV Международной научной конференции. Denpropetrovsk: Oles Honchar Dnipro National University. pp. 502–503. Archived from the original on 17 August 2011. Retrieved 15 July 2015.
  128. ^ "Brown Bear predation of Amur Tiger 1973 account". International Wildlife Magazine. 20 October 2009. from the original on 22 June 2016. Retrieved 14 July 2015.
  129. ^ Goodrich, J. M.; Kerley, L. L.; Smirnov, E. N.; Miquelle, D. G.; McDonald, L.; Quigley, H. B.; Hornocker, M. G. & McDonald, T. (2008). "Survival rates and causes of mortality of Amur tigers on and near the Sikhote-Alin Biosphere Zapovednik". Journal of Zoology. 276 (4): 323. doi:10.1111/j.1469-7998.2008.00458.x.
  130. ^ 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. doi:10.22621/cfn.v122i1.550. PDF 19 January 2021 at the Wayback Machine.
  131. ^ Downey, Betsy. (PDF). International Wolf Center. Archived from the original (PDF) on 27 February 2008.
  132. ^ 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.
  133. ^ ADW: Ursus arctos: Information 12 May 2013 at the Wayback Machine. Arlis.org. Retrieved 9 August 2012.
  134. ^ Hornocker, Maurice; Negri, Sharon (15 December 2009). Cougar: Ecology and Conservation. University of Chicago Press. pp. 163–174. ISBN 978-0-226-35347-0.
  135. ^ 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. S2CID 55281438. Archived (PDF) from the original on 9 October 2022.
  136. ^ 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. Archived (PDF) from the original on 9 October 2022.
  137. ^ 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. Archived (PDF) from the original on 9 October 2022.
  138. ^ Dough O'Hara . Anchorage Daily News. 24 April 2005
  139. ^ "ABC News: Grizzlies Encroaching on Polar Bear Country". ABC News. from the original on 5 August 2017. Retrieved 10 October 2009.
  140. ^ Schwartz, C. C.; Keating, K. A.; Reynolds III, H. V.; Barnes Jr, V. G.; Sellers, R. A.; Swenson, J. E.; Miller, S.D.; McLellan, B.; Keay, J.; McCann, R.; Gibeau, M.; Wakkinen, W.F.; Mace, R.D.; Kasworm, W.; Smith, R.; Herrero, S. (2003). "Reproductive maturation and senescence in the female brown bear". Ursus. 14 (#2): 109–119. JSTOR 3873012.
  141. ^ Macdonald, David David Whyte (2001). "Bear family". The New Encyclopedia of Mammals. Oxford University Press, Incorporated. ISBN 978-0-19-850823-6.
  142. ^ Schwartz, C. C.; Haroldson, M. A.; White, G. C.; Harris, R. B.; Cherry, S.; Keating, K. A.; Moody, D.; Servheen, C. (2006). "Temporal, Spatial, and Environmental Influences on the Demographics of Grizzly Bears in the Greater Yellowstone Ecosystem". Wildlife Monographs. 161: 1–68. doi:10.2193/0084-0173(2006)161[1:TSAEIO]2.0.CO;2. S2CID 85600938.
  143. ^ Rogers, Lynn L.; Rogers, Susanne M. (1976). "Parasites of Bears: A Review". Bears: Their Biology and Management. 3: 411–430. doi:10.2307/3872791. ISSN 1936-0614. JSTOR 3872791.
  144. ^ J. Brglez, S. Valentinclc (1968). "Parasites of brown bear, Ursus arctos L." Acta Veterinaria (Beograd). 18 (6): 379–384.
  145. ^ Di Francesco, Cristina Esmeralda; Gentile, Leonardo; Di Pirro, Vincenza; Ladiana, Lara; Tagliabue, Silvia; Marsilio, Fulvio (1 January 2015). "Serologic Evidence for Selected Infectious Diseases in Marsican Brown Bears (Ursus arctos marsicanus) in Italy (2004–09)". Journal of Wildlife Diseases. 51 (1): 209–213. doi:10.7589/2014-01-021. ISSN 0090-3558. PMID 25375945.
  146. ^ Mörner, Torsten; Eriksson, Hanna; Bröjer, Caroline; Nilsson, Kristina; Uhlhorn, Henrik; Ågren, Erik; Segerstad, Carl Hård af; Jansson, Désirée S.; Gavier-Widén, Dolores (April 2005). "Diseases and mortality in free-ranging brown bear (Ursus arctos), gray wolf (Canis lupus), and wolverine (Gulo gulo) in Sweden". Journal of Wildlife Diseases. 41 (2): 298–303. doi:10.7589/0090-3558-41.2.298. ISSN 0090-3558. PMID 16107663.
  147. ^ a b c d e Fröbert, O.; Frøbert, A. M.; Kindberg, J.; Arnemo, J. M.; Overgaard, M. T. (March 2020). "The brown bear as a translational model for sedentary lifestyle-related diseases". Journal of Internal Medicine. 287 (3): 263–270. doi:10.1111/joim.12983. hdl:11250/2642746. ISSN 0954-6820. PMID 31595572.
  148. ^ Woodroffe, R. (2000). "Predators and people: Using human densities to interpret declines of large carnivores" (PDF). Animal Conservation. 3 (2): 165. Bibcode:2000AnCon...3..165W. doi:10.1111/j.1469-1795.2000.tb00241.x. S2CID 84430290. Archived (PDF) from the original on 9 October 2022.
  149. ^ Kistchinski, A. A. (1972). "'Life history of the brown bear (Ursus arctos L.) in north-east Siberia". Bears: Their Biology and Management. 2: 67–73. doi:10.2307/3872570. JSTOR 3872570.
  150. ^ a b c Herrero, Stephen (2002). Bear Attacks: Their Causes and Avoidance. Globe Pequot Press. pp. 10–25. ISBN 978-1-58574-557-9.
  151. ^ Smith, Tom S.; Herrero, Steven. . Alaska Science Center – Biological Science Office. Archived from the original on 14 August 2009.
  152. ^ Rogers, Lynn L., , Bear.org, archived from the original on 16 October 2002
  153. ^ Cardall, T. Y.; Rosen, P. (2003). "Grizzly bear attack". The Journal of Emergency Medicine. 24 (3): 331–333. doi:10.1016/s0736-4679(03)00004-0. PMID 12676309.
  154. ^ a b Bombieri, G.; Naves, J.; Penteriani, V.; Selva, N.; Fernández-Gil, A.; López-Bao, J. V.; Ambarli, H.; Bautista, C.; Bespalova, T.; Bobrov, V.; Bolshakov, V.; Bondarchuk, S.; Camarra, J. J.; Chiriac, S.; Ciucci, P. (12 June 2019). "Brown bear attacks on humans: a worldwide perspective". Scientific Reports. 9 (1): 8573. Bibcode:2019NatSR...9.8573B. doi:10.1038/s41598-019-44341-w. ISSN 2045-2322. PMC 6562097. PMID 31189927.
  155. ^ Dinets, Vladimir. . Archived from the original on 15 December 2012.
  156. ^ Dickman, A. J.; Hazzah, L. (2016). "Money, Myths and Man-Eaters: Complexities of Human–Wildlife Conflict". Problematic Wildlife. Springer. pp. 339–356. ISBN 978-3-319-22246-2.
  157. ^ Smith, Herrero; DeBruyn, Wilde (2008). . North American Bear Center. Archived from the original on 1 January 2011.
  158. ^ McLaren, Duncan & Wigen, Rebecca & Mackie, Quentin & Fedje, Daryl. (2005). Bear Hunting at the Pleistocene/Holocene Transition on the Northern Northwest Coast of North America. pp. 4–23
  159. ^ Bischof, Richard; Bonenfant, Christophe; Rivrud, Inger Maren; Zedrosser, Andreas; Friebe, Andrea; Coulson, Tim; Mysterud, Atle; Swenson, Jon E. (January 2018). "Regulated hunting re-shapes the life history of brown bears". Nature Ecology & Evolution. 2 (1): 116–123. doi:10.1038/s41559-017-0400-7. hdl:10852/67379. ISSN 2397-334X. PMID 29230025. S2CID 3288368.
  160. ^ Albert, David M.; Bowyer, R. Terry; Miller, Sterling D. (2001). "Effort and Success of Brown Bear Hunters in Alaska". Wildlife Society Bulletin. 29 (2): 501–508. ISSN 0091-7648. JSTOR 3784174.
  161. ^ Kelava Ugarković, Nikolina; Konjačić, Miljenko; Malnar, Josip; Tomljanović, Kristijan; Šprem, Nikica; Ugarković, Damir (January 2021). "Proximate Chemical Composition, Fatty Acid Profile, and Lipid Qualitative Indices of Brown Bear Meat". Foods. 10 (1): 36. doi:10.3390/foods10010036. ISSN 2304-8158. PMC 7824718. PMID 33374425.
  162. ^ Pastorino, Giovanni Quintavalle; Christodoulides, Yiannis; Curone, Giulio; Pearce-Kelly, Paul; Faustini, Massimo; Albertini, Mariangela; Preziosi, Richard; Mazzola, Silvia Michela (May 2017). "Behavioural Profiles of Brown and Sloth Bears in Captivity". Animals. 7 (5): 39. doi:10.3390/ani7050039. ISSN 2076-2615. PMC 5447921. PMID 28505095.
  163. ^ Montaudouin, S.; Pape, G. Le (1 July 2005). "Comparison between 28 zoological parks: stereotypic and social behaviours of captive brown bears (Ursus arctos)". Applied Animal Behaviour Science. 92 (1): 129–141. doi:10.1016/j.applanim.2004.10.015. ISSN 0168-1591.
  164. ^ Stagni, Elena; Sequeira, Sara; Brscic, Marta; Redtenbacher, Irene; Hartmann, Sabine (13 December 2023). "A retrospective study on the prevalence of main clinical findings in brown bears (Ursus arctos) rescued from substandard husbandry conditions". Frontiers in Veterinary Science. 10: 1299029. doi:10.3389/fvets.2023.1299029. ISSN 2297-1769. PMC 10773888. PMID 38192718.
  165. ^ Iossa, G.; Soulsbury, C. D.; Harris, S. (May 2009). "Are wild animals suited to a travelling circus life?". Animal Welfare. 18 (2): 129–140. doi:10.1017/S0962728600000270. ISSN 0962-7286. S2CID 32259865.
  166. ^ Newman, A. R. (1987). "Images of the bear in Children's Literature". Children's Literature in Education. 18 (#3): 131–138. doi:10.1007/bf01130991. S2CID 143882256.
  167. ^ Bieder, Robert E. (18 August 2005). "Survivors". Bear. Reaktion Books. ISBN 978-1-86189-482-3.
  168. ^ O'Regan, Hannah (4 September 2023). "Brown bears in burials and entertainment in later prehistoric to modern Britain (c. 2400 BC – AD 1900s)". The Archaeology of Northern Europe: 187–208. doi:10.1484/M.TANE-EB.5.134334. ISBN 978-2-503-60611-8.
  169. ^ De Swart, Herman (2004). "Cave bears in prehistoric art; a survey from the literature". Publications du musée des Confluences. 2 (1): 121–128.
  170. ^ "Karhu on Suomen kansalliseläin". yle.fi. 16 September 2010. from the original on 11 January 2016. Retrieved 18 August 2016.
  171. ^ Riabov, Oleg (2020). "The Birth of the Russian Bear? The Bear Symbol in the Satirical Journals of the Russian Revolution of 1905". Region. 9 (1): 139–168. ISSN 2166-4307. JSTOR 27007706.
  172. ^ . Montana Historical Society. Archived from the original on 18 December 2009. Retrieved 18 August 2016.
  173. ^ . California State Library. Archived from the original on 5 January 2019. Retrieved 24 September 2011.
  174. ^ Hughes, Courtney; Foote, Lee; Yarmey, Nicholas T.; Hwang, Christina; Thorlakson, Jessica; Nielsen, Scott (April 2020). "From human invaders to problem bears: A media content analysis of grizzly bear conservation". Conservation Science and Practice. 2 (4). Bibcode:2020ConSP...2E.176H. doi:10.1111/csp2.176. ISSN 2578-4854.
  175. ^ Deyermond, A. D. (1997). Festschrift. Tamesis. pp. 177–187. ISBN 978-1-85566-051-9.
  176. ^ Swenson, Jon E.; Europe, Council of (1 January 2000). Action Plan for the Conservation of the Brown Bear in Europe (Ursus Arctos). Council of Europe. p. 69. ISBN 978-92-871-4426-3.
  177. ^ Majić, Aleksandra; Marino Taussig de Bodonia, Agnese; Huber, Đuro; Bunnefeld, Nils (December 2011). "Dynamics of public attitudes toward bears and the role of bear hunting in Croatia". Biological Conservation. 144 (12): 3018–3027. Bibcode:2011BCons.144.3018M. doi:10.1016/j.biocon.2011.09.005. ISSN 0006-3207.

Notes

  1. ^ Populations of Bhutan, China, Mexico and Mongolia are included in Appendix I. All other populations are included in Appendix II.

Bibliography

  • Vaisfeld, M.A. and Chestin I. E., ed. (1993). Bears: Brown Bear, Polar Bear, Asian Black Bear. Distribution, ecology, use and protection (in Russian and English). Moscow: Nauka. ISBN 978-5020035676.

External links

 
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  • from National Geographic
  • Bear Hunting Altered Genetics More Than Ice Age Isolation
  • Ancient Fossil Offers New Clues To Brown Bears Past

January 01, 1970
brown, bear, this, article, about, animal, athletics, teams, brown, university, brown, bears, research, ship, brown, bear, brown, bear, ursus, arctos, large, bear, species, found, across, eurasia, north, america, largest, living, terrestrial, members, order, c. This article is about the animal For the athletics teams at Brown University see Brown Bears For the research ship see MV Brown Bear The brown bear Ursus arctos is a large bear species found across Eurasia and North America It is one of the largest living terrestrial members of the order Carnivora rivaled in size only by its closest relative the polar bear which is much less variable in size and slightly bigger on average Adults of different subspecies range in weight from 80 to 600 kg 180 to 1 320 lb with males being heavier than females Despite its name brown bears aren t entirely brown the pelage can be reddish to yellowish brown and dark brown to cream in color During winter brown bears in some populations hibernate and emerge during spring to regain up to 180 kg 400 lb of weight They have well developed dentition and claws ideal for their lifestyle Brown bearTemporal range 0 5 0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Middle Pleistocene Holocene Kodiak bear on Kodiak Island Conservation status Least Concern IUCN 3 1 1 CITES Appendix II CITES 2 note 1 Scientific classification Domain Eukaryota Kingdom Animalia Phylum Chordata Class Mammalia Order Carnivora Family Ursidae Genus Ursus Species U arctos Binomial name Ursus arctosLinnaeus 1758 Subspecies 15 see text and article Brown bear range map The brown bear is mostly found in forested habitats in elevations of up to 5 000 m 16 000 ft It is omnivorous and consumes a variety of plant and animal species with the former comprising 90 of its diet The bear hunts animals as small as rodents to animals as large as moose or muskoxen In parts of coastal Alaska brown bears predominately feed on spawning salmon that come ashore to lay their eggs The brown bear is a solitary animal except in the breeding season Females protect their young for an average of 1 5 to 4 5 years Brown bears have one of the largest skulls of any land based carnivore and are able to make use of tools They are long lived animals with an average lifespan of 25 years in the wild Attacks on humans though reported are generally rare While the brown bear s range has shrunk and it has faced local extinctions across its wide range it remains listed as a least concern species by the International Union for Conservation of Nature IUCN with a total estimated population in 2017 of 110 000 Populations that were hunted to extinction in the 19th and 20th centuries are the Atlas bear of North Africa and the Californian Ungavan and Mexican populations of the grizzly bear of North America Many of the populations in the southern parts of Eurasia are highly endangered as well One of the smaller bodied forms the Himalayan brown bear is critically endangered occupying only 2 of its former range and threatened by uncontrolled poaching for its body parts The Marsican brown bear of central Italy is one of several currently isolated populations of the Eurasian brown bear and is believed to have a population of just c 50 bears Contents 1 Etymology 2 Evolution and taxonomy 2 1 Evolution 2 2 Subspecies 2 3 Hybrids 3 Description 3 1 Coloration 3 2 Cranial morphology and size 3 3 Claws and feet 4 Distribution and habitat 4 1 Conservation status 5 Behavior and life history 5 1 Communication 5 2 Home ranges 5 3 Reproduction 5 4 Dietary habits 5 5 Interspecific predatory relationships 5 6 Longevity and mortality 5 7 Hibernation physiology 6 Relationship with humans 6 1 Attacks on humans 6 2 Bear hunting 6 3 In captivity 6 4 Culture 7 References 8 Notes 9 Bibliography 10 External linksEtymologyThe 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 3 4 In the mid 19th century United States the brown bear was termed Old Ephraim and sometimes as Moccasin Joe 5 The scientific name of the brown bear Ursus arctos comes from the Latin ursus meaning bear 6 and the Greek ἄrktos arktos also meaning bear 7 Evolution and taxonomyEvolution The brown bear is one of eight extant species in the bear family Ursidae and of six extant species in the subfamily Ursinae A possible phylogeny based on complete mitochondrial DNA sequences from Yu et al 2007 8 Ursidae Giant panda Ailuropoda melanoleuca nbsp Spectacled bear Tremarctos ornatus nbsp Ursinae Sloth bear Melursus ursinus nbsp Sun bear Helarctos malayanus nbsp Asian black bear Ursus thibetanus nbsp American black bear Ursus americanus nbsp Polar bear Ursus maritimus nbsp Brown bear Ursus arctos nbsp The polar bear and the brown bear form a close grouping while the relationships of the other species are not very well resolved 9 A more recent phylogeny based on the genetic study of Kumar et al 2017 10 Ursidae Giant panda Ailuropoda melanoleuca nbsp Spectacled bear Tremarctos ornatus nbsp Ursinae Sloth bear Melursus ursinus nbsp Sun bear Helarctos malayanus nbsp Asian black bear Ursus thibetanus nbsp American black bear Ursus americanus nbsp Polar bear Ursus maritimus nbsp Brown bear Ursus arctos nbsp The study concludes that Ursine bears originated around 5 million years ago and show extensive hybridization of species in their lineage 10 During the Pleistocene brown bears were sympatric with extinct Ursinae species in North America 11 12 Brown bears are thought to have evolved from Ursus etruscus in Asia 13 14 The brown bear per Kurten 1976 has been stated as clearly derived from the Asian population of Ursus savini about 800 000 years ago spread into Europe to the New World 15 A genetic analysis indicated that the brown bear lineage diverged from the cave bear species complex approximately 1 2 1 4 million years ago but did not clarify if U savini persisted as a paraspecies for the brown bear before perishing 16 The oldest fossils positively identified as from this species occur in China from about 0 5 million years ago 13 17 In Yakutia brown bears are known to have been present from at least 400 000 to 300 000 years ago 18 Brown bears entered Europe about 250 000 years ago and North Africa shortly after 13 Brown bears were present in the Zagros Mountains by the Late Pleistocene 19 Brown bear remains from the Pleistocene period are common in the British Isles where amongst other factors they may have contributed to the extinction of cave bears Ursus spelaeus 20 Brown bears along with lions bison and red foxes first emigrated to North America from Eurasia via Beringia during the Illinoian Glaciation 21 Although Bjorn Kurten hypothesized skull shape as an indicator of evolutionary history 20 genetic evidence suggests several different populations of brown bear migrated into North America aligning with the glacial cycles of the Pleistocene The founding population of most North American brown bears arrived first with the genetic lineage developing around 177 000 BP with fossils being first recovered 111 000 BP from East Beringia Genetic divergences suggest brown bears first migrated south during MIS 5 92 000 83 000 BP upon the opening of the ice free corridor 21 22 with the first fossils being near Edmonton 26 000 BP 12 The Alexander and Haida Gwaii archipelagoes have a separate lineage which first appears around 20 000 BP After a local extinction in Beringia 33 000 BP two new but closely related lineages repopulated Alaska and northern Canada from Eurasia after the Last Glacial Maximum gt 25 000 BP 21 Brown bear fossils discovered in Ontario Ohio Kentucky and Labrador show that the species occurred farther east than indicated in historic records 13 In North America two types of the subspecies Ursus arctos horribilis are generally recognized the coastal brown bear and the inland grizzly bear 23 Subspecies Main article Subspecies of brown bear nbsp Adult female Eurasian brown bear the nominate subspecies Brown bear taxonomy and subspecies classification has been described as formidable and confusing with few authorities listing the same specific set of subspecies 24 There are hundreds of obsolete brown bear subspecies each with its own name so this can become confusing As many as 90 subspecies have been proposed 25 26 However recent DNA analysis has identified as few as five main clades which contain all extant brown bears 27 28 while a 2017 phylogenetic study revealed nine clades including one representing polar bears 29 As of 2005 update 15 extant or recently extinct subspecies were recognized by the general scientific community 30 31 As well as the exact number of overall brown bear subspecies its precise relationship to the polar bear also remains in debate The polar bear is a recent offshoot of the brown bear The point at which the polar bear diverged from the brown bear is unclear with estimations based on genetics and fossils ranging from 400 000 to 70 000 years ago but most recent analysis has indicated that the polar bear split somewhere between 275 000 and 150 000 years ago 32 Under some definitions the brown bear can be construed as the paraspecies for the polar bear 33 DNA analysis shows that apart from recent human caused population fragmentation 34 brown bears in North America are generally part of a single interconnected population system with the exception of the population or subspecies in the Kodiak Archipelago which has probably been isolated since the end of the last Ice Age 35 36 These data demonstrate that U a gyas U a horribilis U a sitkensis and U a stikeenensis are not distinct or cohesive groups and would more accurately be described as ecotypes For example brown bears in any particular region of the Alaska coast are more closely related to adjacent grizzly bears than to distant populations of brown bears 37 the morphological distinction seemingly driven by brown bears having access to a rich salmon food source while grizzly bears live at higher elevation or further from the coast where plant material is the base of the diet The history of the bears of the Alexander Archipelago is unusual in that these island populations carry polar bear DNA presumably originating from a population of polar bears that was left behind at the end of the Pleistocene but have since been connected with adjacent mainland populations through movement of males to the point where their nuclear genomes are now more than 90 of brown bear ancestry 38 MtDNA analysis revealed that brown bears are apparently divided into five different clades some of which coexist or co occur in different regions 39 Hybrids See also Grizzly black bear hybrid and Grizzly polar bear hybrid nbsp Possible grizzly black bear hybrid in the Yukon Territory Canada A grizzly polar bear hybrid is a rare ursid hybrid resulting from a crossbreeding 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 and seven more hybrids have since been confirmed in the same region all descended from a single female polar bear 40 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 41 Analyses of the genomes of bears have shown that introgression between species was widespread during the evolution of the genus Ursus 42 including the introgression of polar bear DNA introduced to brown bears during the Pleistocene 43 Description nbsp Brown bears are highly variable in size Eurasian brown bears often fall around the middle to low sizes for the species The brown bear is the most variable in size of modern bears The typical size depends upon which population it is from and most accepted subtypes vary widely in size This is in part due to sexual dimorphism as male brown bears average at least 30 larger in most subtypes Individual bears vary in size seasonally weighing the least in spring due to lack of foraging during hibernation and the most in late fall after a period of hyperphagia to put on additional weight to prepare for hibernation Therefore a bear may need to be weighed in both spring and fall to get an idea of its mean annual weight 44 45 Generally brown bears weigh anywhere from 80 to 600 kg 180 to 1 320 lb with males outweighing females 46 The normal range of physical dimensions for a brown bear is a head and body length of 1 4 to 2 8 m 4 ft 7 in to 9 ft 2 in and a shoulder height of 70 to 153 cm 2 ft 4 in to 5 ft 0 in The tail is relatively short as in all bears ranging from 6 to 22 cm 2 4 to 8 7 in in length 47 48 The smallest brown bears females during spring among barren ground populations can weigh so little as to roughly match the body mass of males of the smallest living bear species the sun bear Helarctos malayanus while the largest coastal populations attain sizes broadly similar to those of the largest living bear species the polar bear 49 Interior brown bears are generally smaller than is often perceived being around the same weight as an average lion at an estimate average of 180 kg 400 lb in males and 135 kg 298 lb in females whereas adults of the coastal populations weigh about twice as much 50 The average weight of adult male bears from 19 populations from around the world and various subspecies including both large and small bodied subspecies was found to be 217 kg 478 lb while adult females from 24 populations were found to average 152 kg 335 lb 23 51 Coloration nbsp A brown bear in Slovenia Brown bears are often not fully brown 52 They have long thick fur with a moderately long mane at the back of the neck which varies somewhat across the types 53 In India brown bears can be reddish with silver tipped hairs while in China brown bears are bicolored with a yellowish brown or whitish collar across the neck chest and shoulders 52 54 Even within well defined subspecies individuals may show highly variable hues of brown North American grizzlies can be dark brown almost black to cream almost white or yellowish brown and often have darker colored legs The common name grizzly stems from their typical coloration with the hairs on their back usually being brownish black at the base and whitish cream at the tips giving them their distinctive grizzled color Apart from the cinnamon subspecies of the American black bear U americanus cinnamonum the brown bear is the only modern bear species to typically appear truly brown 55 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 with its length and density varying geographically 56 Cranial morphology and size nbsp Brown bear skeleton nbsp Skull Adults have massive heavily built concave skulls which are large in proportion to the body 55 The projections of the skull are well developed 56 Skull lengths of Russian brown 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 56 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 56 The teeth of brown bears reflect their dietary plasticity and are broadly similar to other bears 57 58 The teeth are reliably larger than American black bears but average smaller in molar length than polar bears 59 Brown bears have the broadest skull of any extant ursine bear 50 Claws and feet nbsp Front paws 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 may measure 7 to 10 centimetres 2 8 to 3 9 in along the curve 56 They are generally dark with a light tip with some forms having completely light claws 56 Brown bear claws are longer and straighter than those of American black bears Ursus americanus 55 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 typically climb trees as well as black bears although in rare cases adult female brown bears have been seen scaling trees 60 The claws of a polar bear are quite different being notably shorter but broader with a strong curve and sharper point 17 61 The paws of the brown bear are quite large The rear feet of adult bears measure 21 to 36 cm 8 3 to 14 2 in long while the forefeet tend to measure about 40 less in length 62 Brown bears are the only extant bears with a hump at the top of their shoulder which is made entirely of muscle this feature having developed presumably for imparting more force in digging which is habitual during foraging for the bear and also used heavily in den construction prior to hibernation 55 The brown bear s strength has been roughly estimated as 2 5 to 5 times that of a human 63 Distribution and habitatMain article Distribution of brown bears nbsp Brown bear at Brooks Falls Brown bears were once native to Europe much of Asia the Atlas Mountains of Africa and North America 64 but are now extirpated in some areas and their populations have greatly decreased in other areas There are approximately 200 000 brown bears left in the world 65 The largest populations are in Russia with 130 000 66 the United States with 32 500 and Canada with around 25 000 Brown bears live in Alaska east through the Yukon and Northwest Territories south through British Columbia and through the western half of Alberta The Alaskan population is estimated at a healthy 30 000 individuals 67 In the lower 48 states they are repopulating slowly but steadily along the Rockies and the western Great Plains 68 In Europe in 2010 there were 14 000 brown bears in ten fragmented populations from Spain estimated at only 20 25 animals in the Pyrenees in 2010 69 in a range shared between Spain France and Andorra and some 210 animals in Asturias Cantabria Galicia and Leon in the Picos de Europa and adjacent areas in 2013 70 in the west to Russia in the east and from Sweden and Finland in the north to Romania 5000 6000 Bulgaria 900 1200 Slovakia with about 600 800 animals Slovenia 500 700 animals and Greece with Karamanlidis et al 2015 estimating gt 450 animals in the south 71 72 In Asia brown bears are found primarily throughout Russia thence more spottily southwest to parts of the Middle East including almost all parts of Kurdistan to as far south as southwestern Iran and to the southeast in Northeast China Brown bears are also found in Western China Kyrgyzstan North Korea Pakistan Afghanistan and India A population of brown bear can be found on the Japanese island of Hokkaidō which holds the largest number of non Russian brown bears in eastern Asia with about 2 000 3 000 animals 62 This species inhabits the broadest range of habitats of any living bear species 62 They seem to have no altitudinal preferences and have been recorded from sea level to an elevation of 5 000 m 16 000 ft in the Himalayas 62 In most of their range brown bears generally seem 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 62 Conservation status nbsp A Marsican brown bear with a range restricted to the Abruzzo Lazio and Molise National Park Italy 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 update this and the American black bear are the only bear species not classified as threatened by the IUCN 1 39 However the California grizzly bear Ungava brown bear Atlas bear and Mexican grizzly bear as well as brown bear populations in the Pacific Northwest were hunted to extinction in the 19th and early 20th centuries and many of the southern Asian subspecies are highly endangered 73 The Syrian brown bear U a syriacus is very rare and it has been extirpated from more than half of its historic range 74 One of the smallest bodied subspecies the Himalayan brown bear U a isabellinus is critically endangered occupying only 2 of its former range and threatened by uncontrolled poaching for its body parts 75 The Marsican brown bear in central Italy is believed to have a population of just 50 bears 76 Behavior and life history nbsp Like all bears brown bears can stand on their hindlegs and walk for a few steps in this position usually motivated to do so by curiosity hunger or alarm The brown bear is often described as nocturnal However it frequently seems to peak in activity in the morning and early evening hours 77 Activity can occur at nearly any time of night or day with bears who dwell in areas with more extensive human contact being more likely to be fully nocturnal Furthermore yearling and newly independent bears are more likely to be active diurnally and many adult bears in low disturbance areas are largely crepuscular 78 79 80 In summer through autumn a brown bear can double its weight from the spring gaining up to 180 kg 400 lb of fat on which it relies to make it through winter when it becomes very lethargic 81 82 Although they are not full hibernators and can be woken easily both sexes like to den in a protected spot during the winter months Hibernation dens may consist of any spot that provides cover from the elements and that can accommodate their bodies such as a cave crevice cavernous tree roots or hollow logs 83 84 Brown bears have one of the largest brains of any extant carnivoran relative to their body size and have been shown to engage in tool use which requires advanced cognitive abilities 85 This species is mostly solitary although bears may gather in large numbers at major food sources e g open garbage dumps or rivers holding spawning salmon and form social hierarchies based on age and size 86 87 Adult male bears are particularly aggressive and are avoided by adolescent and subadult males both at concentrated feeding opportunities and chance encounters Female bears with cubs rival adult males in aggression and are much 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 88 89 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 86 During combat bears use their paws to strike their opponents in the chest or shoulders and bite the head or neck 90 Communication nbsp Brown bear sounds source source Brown bear sounds including huffing jaw popping and growlingBrown bear roars source source Brown bears roaring over a carcass Problems playing these files See media help Several different facial expressions have been documented in brown bears The relaxed face is made in everyday activities and has the ears pointed to the sides and the mouth closed or slackly open During social play bears make relaxed open mouth face in which the mouth is open with a curled upper lip and hanging lower lip and the ears alert and shifting When looking at another animal at a distance the bear makes an alert face as the ears are cocked and alert the eyes wide open but the mouth is closed or only open slightly The tense closed mouth face is made with the ears laid back and the mouth closed and occurs when the bear feels threatened When approached by another individual the animal makes a puckered lip face with a protruding upper lip and ears which go from cocked and alert when at a certain distance to laid back when closer or when retreating The jaw gape face consists of an open mouth with visible lower canines and hanging lips while the biting face is similar to the relaxed open mouth face except the ears are flattened and the eyes are wide enough to expose the sclera Both the jaw gape face and the biting face are made in aggression and bears switch between them 90 Brown bears also produce various vocalizations Huffing occurs when the animal is tense while woofing is made when alarmed Both sounds are produced by exhalations though huffing is harsher and is made continuously two per second Growls and roars are made in aggression Growling is harsh and guttural and can range from a simple grrr to a rumble A rumbling growl can escalate into a roar when the bear is charging Roaring is described as thunderous and can travel 2 km 1 2 mi Mothers and cubs wanting physical contact will bawl which is heard as waugh waugh 90 Home ranges 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 23 89 Males always cover more area than females each year Despite their lack of traditional territorial behavior adult males can seem to have a personal zone in which other bears are not tolerated if they are seen 91 Males always wander further than females due to both increasing access to females and food sources while females are advantaged by smaller territories in part since it decreases the likelihood of encounters with male bears who may endanger their cubs 23 92 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 and 318 km2 44 and 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 Romania the largest home range was recorded for adult males 3 143 km2 1 214 sq mi 93 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 94 89 Reproduction nbsp Pair of mating brown bears at the Ahtari Zoo in Ahtari Finland The mating season is from mid May to early July shifting later the further north the bears are found 95 Being serially monogamous brown bears remain with the same mate from a couple of days to a couple of weeks 23 Outside of this narrow time frame adult male and female brown bears show no sexual interest in each other 23 Females mature sexually between the age of four and eight years of age with an average age at sexual maturity of 5 2 5 5 years old 96 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 91 97 Males will try to mate with as many females as they can usually a successful one mates with two females in a span of one to three weeks 48 97 The adult female brown bear is similarly promiscuous mating with up to four rarely even eight males while in heat and potentially breeding with two males in a single day 98 Females come into oestrus on average every three to four years with a full range of 2 4 to 5 7 years The urine markings of a female in oestrus can attract several males via scent 99 Dominant males may try to sequester a female for her entire oestrus period of approximately two weeks but usually are unable to retain her for the entire time 50 92 Copulation is vigorous and prolonged and can last up to an hour although the mean time is about 23 24 minutes 50 100 nbsp Grizzly bear cubs often imitate their mothers closely Males take no part in raising their cubs parenting is left entirely to the females 89 101 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 while gestating the embryo does not implant and is reabsorbed into the body 96 102 103 There have been cases of brown bears with as many as six cubs although the average litter size is one to three with more than four being considered uncommon 104 There are records of females sometimes adopting stray cubs or even trading or kidnapping cubs when they emerge from hibernation a larger female may claim cubs away from a smaller one 62 105 106 Older and larger females within a population tend to give birth to larger litters 107 The size of a litter also depends on factors such as geographic location and food supply 108 At birth the cubs are blind toothless and hairless and may weigh from 350 to 510 g 0 77 to 1 12 lb again reportedly based on the age and condition of the mother 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 over long distances and begin to forage for solid food 23 109 nbsp Kodiak bear cubs play fighting The cubs are fully dependent on the mother and a close bond is formed During the dependency stage the cubs learn rather than inherit as instincts from birth 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 94 Increased brain size in large carnivores has been positively linked to whether a given species is solitary as is the brown bear or raises their offspring communally thus female brown bears have relatively large well developed brains presumably key in teaching behavior 110 The cubs learn by following and imitating their mother s actions during the period they are with her 62 Cubs remain with their mother for an average of 2 5 years in North America uncommonly being independent as early as 1 5 years of age or as late as 4 5 years of age 50 The stage at which independence is attained may generally be earlier in some parts of Eurasia as the latest date which mother and cubs were together was 2 3 years most families separated in under two years in a study from Hokkaido and in Sweden most cubs on their own were still yearlings 111 112 Brown bears practice infanticide as an adult male bear may kill the cubs of another 92 When an adult male brown bear kills a cub it is usually because he is trying to bring the female into oestrus as she will enter that state within two to four days after the death of her cubs 92 Cubs may flee up a tree when they see a strange male bear approaching The mother often successfully defends them even though the male may be twice as heavy as her However females have been known to die in such confrontations 92 113 114 Dietary habits Main article Dietary biology of the brown bear nbsp 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 62 Despite their reputation most brown bears are not highly carnivorous as they derive up to 90 of their dietary food energy from vegetable matter 115 They often feed on a variety of plant life including berries grasses flowers acorns and pine cones as well as fungi such as mushrooms 23 Among all bears brown bears are uniquely equipped to dig for tough foods such as roots bulbs 116 and shoots They use their long strong claws to dig out earth to reach the roots and their powerful jaws to bite through them 23 In spring winter provided carrion grasses shoots sedges moss 116 and forbs are the dietary mainstays for brown bears internationally 62 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 62 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 in a single day and may derive up to half of their annual food energy from these insects 117 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 55 This species may eat birds and their eggs including almost entirely ground or rock nesting species 55 The diet may be supplemented by rodents or similar smallish mammals including marmots ground squirrels mice rats lemmings and voles 23 With particular regularity bears in Denali National Park will wait at burrows of Arctic ground squirrels hoping to pick off a few of the 1 kg 2 2 lb rodents 118 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 nutrious 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 55 nbsp A cow moose with calves being approached by an inland brown bear Denali National Park Alaska Beyond the regular predation of salmon most brown bears are not particularly active predators 55 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 55 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 55 Large mammals preyed on can include various ungulate species such as elk moose caribou muskoxen and wild boar 23 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 119 It will also bite or swipe some prey to stun it enough to knock it over for consumption 120 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 ambush young animals by finding them via scent 23 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 120 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 23 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 55 They feed on carrion and use their size to intimidate other predators such as wolves cougars tigers and American black bears from their kills Carrion is especially important in the early spring when the bears are emerging from hibernation much of it comprising winter killed big game 23 Cannibalism is not unheard of though predation is not normally believed to be the primary motivation when brown bears attack each other 55 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 are sometimes exploited as prey Cattle are bitten on the neck back or head and then the abdominal cavity is opened for eating 23 Plants and fruit farmed by humans are readily consumed as well including corn wheat sorghum melons and any form of berries 55 They may feed on domestic bee yards readily consuming both honey and the brood grubs and pupae of the honey bee colony 55 Human foods and trash are 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 55 Interspecific predatory relationships nbsp Brown bear being followed by a wolf Adult bears are generally immune to predatory attacks except from large Siberian Amur tigers and other bears Following a decrease of ungulate populations from 1944 to 1959 32 cases of Siberian tigers attacking both Ussuri brown bears Ursus arctos lasiotus and Ussuri black bears U thibetanus ussuricus were recorded in the Russian Far East and hair of bears were found in several tiger scat samples Tigers attack black bears less often than brown bears since the black bears live in more open habitats and are not able to climb trees In the same time period four cases of brown bears killing female tigers and young cubs were reported both in disputes over prey and in self defense Tigers mainly feed on the bear s fat deposits such as the back hams and groin 56 In rare cases when Amur tigers prey on brown bears they usually target young and sub adult bears besides small female adults taken outside their dens generally when lethargic from hibernation 121 Predation by tigers on denned brown bears was not detected during a study carried between 1993 and 2002 122 Ussuri brown bears along with the smaller black bears constitute 2 1 of the Siberian tiger s annual diet of which 1 4 are brown bears 123 124 The effect the presence of tigers have on a brown bears behavior seems to vary In the winters of 1970 1973 Yudakov and Nikolaev recorded two cases of bears showing no fear of tigers and another case of a brown bear changing path upon crossing tiger tracks 125 Other researchers have observed bears following tiger tracks to scavenge tiger kills and to potentially prey on tigers 56 123 Despite the threat of predation some brown bears actually benefit from the presence of tigers by appropriating tiger kills that the bears may not be able to successfully hunt themselves 123 During telemetry research in the Sikhote Alin Nature Reserve 44 direct confrontations between bears and tigers were observed in which bears not just brown bears in general were killed in 22 cases and tigers in 12 cases 126 There are reports of brown bears specifically targeting Amur leopards and tigers 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 127 Some studies show that bears frequently track down tigers to usurp their kills with occasional fatal outcomes for the tiger A report from 1973 describes twelve known cases of brown bears killing tigers including adult males In all cases the tigers were subsequently eaten by the bears 128 129 Brown bears regularly intimidate wolves to drive them away from their kills In Yellowstone National Park 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 recorded case of fully grown wolves being killed by a grizzly bear 130 Given the opportunity however both species will prey on the other s cubs 131 In some areas grizzly bears regularly displace cougars from their kills 132 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 133 134 Smaller carnivorous animals including coyotes wolverines lynxes and any other sympatric carnivores or raptorial birds are dominated by grizzly 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 55 There is one record of a golden eagle preying on a brown bear cub 135 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 grizzly bears in open unforested areas Although displacement of black bears by grizzly bears has been documented actual interspecific killing of black bears by grizzlies 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 grizzly s largely nocturnal habits and preference for open spaces 136 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 137 As of the 21st century there has been an increase in interactions between brown bears and polar bears theorized to be caused by climate change Brown and grizzly bears have been seen moving increasingly northward into territories formerly claimed by polar bears They tend to dominate polar bears in disputes over carcasses 138 and dead polar bear cubs have been found in brown bear dens 139 Longevity and mortality nbsp Front paw imprint nbsp Rear paw imprint The brown bear has a naturally long life Wild females have been observed reproducing up to 28 years of age which is the oldest known age for reproduction of any ursid in the wild The peak reproductive age for females ranges from four to 20 years old 62 140 The lifespan of brown bears of both sexes within minimally hunted populations is estimated at an average of 25 years The oldest wild brown bear on record was nearly 37 years old 141 The oldest recorded female in captivity was nearly 40 years old while males in captivity have been verified to live up to 47 years with one captive male possibly attaining 50 years of age 23 48 While male bears potentially live longer in captivity female grizzly bears have a greater annual survival rate than males within wild populations per a study done in the Greater Yellowstone Ecosystem 142 Annual mortality for bears of any age is estimated at 10 in most protected areas 62 Around 13 to 44 of cubs die within their first year even in well protected areas 23 Beyond predation by large predators including wolves Siberian tigers and other brown bears starvation and accidents also claim the lives of cubs Studies have indicated that the most prevalent source of mortality for first year cubs is malnutrition 23 Brown bears are susceptible to parasites such as flukes ticks tapeworms roundworms and biting lices 143 144 It is thought that brown bears may catch canine distemper virus CDV from other Caniforms like stray dogs and wolves 145 A captive individual allegedly succumbed to Aujeszky s disease 146 Hibernation physiology Scientists are interested in understanding the hibernation physiology of brown bears specifically in how they stay healthy after six months of inactivity 147 It has been suggested by scientists that understanding how brown bears stay healthy during hibernation may potentially help lead to cures for human diseases such as diabetes or cardiovascular disease 147 A study conducted through the Scandinavian Brown Bear Research Project did a proteomic analysis of the brown bear s blood organs and tissues to pinpoint proteins and peptides that either increased or decreased in expression in the winter and summer months 147 One major finding was that the plasma protein sex hormone binding globulin SHBG increased during the brown bear s hibernation period by 45 times 147 Although scientists do not yet understand the role of SHBG in maintaining the brown bear s health some scientists believe these findings could potentially be a factor in preventing human diseases that come from a sedentary life style 147 Relationship with humansAttacks on humans Main article Bear attack nbsp A statue of the Ussuri brown bear from Hokkaido Japan Brown bears usually avoid areas where extensive development or urbanization has occurred 148 They rarely attack humans on sight and usually avoid people 149 They are however unpredictable in temperament and may attack if they are surprised or feel threatened 150 The two most common causes for bear attacks are surprise and curiosity 151 Mothers defending cubs are the most prone to attacking being responsible for 70 of brown bear caused human fatalities in North America 152 Brown bear attacks tend to result in serious injury and in some cases death 150 Due to the bears enormous physical strength even a single bite or swipe can be deadly 153 Violent encounters with brown bears usually last only a few minutes though they can be prolonged if the victims fight back 150 A study conducted in 2019 found that 664 bear attacks were reported during a 15 year period 2000 2015 throughout North America and Eurasia There were 568 injuries and 95 casualties 154 Around 10 people a year are killed by brown bears in Russia more than all the other parts of the brown bear s range combined 155 In Japan a large brown bear nicknamed Kesagake kesa style slasher caused the worst brown bear attack in Japanese history at Tomamae Hokkaidō during numerous encounters during December 1915 It killed seven people and wounded three others before being gunned down after a large scale beast hunt 156 A study by U S and Canadian researchers has found bear spray to be more effective at stopping aggressive bear behavior than guns working in 92 of studied incidents versus 67 for guns 157 Bear hunting Main article Bear hunting Hunting of brown bears has occurred since ancient times mainly for sports and royal showings of power Humans have been recorded hunting brown bears for over 10 300 9 300 years 158 In Europe between the 17th and 18th centuries humans sought to control brown bear numbers by awarding those who managed to kill one This bounty scheme pushed the brown bear population to the brink of extinction before comprehensive protection was offered in the 1900s In northern Europe hunting still persists in the 21st century contributing significantly to the drop in brown bear numbers 159 In some places in Europe bears are tricked into entering traps with the help of a bait 154 Brown bears are legally hunted in some American states such as Alaska However a hunting license is required and killings of females with cubs will result in a prison sentence 160 Their meat is sometimes consumed and used in recipes such as dumplings hams and stews The Indigenous people of James Bay Eastern Cree use their flesh in traditional dishes In Asia and Romania the paws are consumed as exotic delectables it has been a prevalent component of traditional Chinese food since 500 BC The total weight of commercially sold brown bear meat is estimated at 17 tons annually 161 In captivity There are more than 700 brown bears in zoos and wildlife parks worldwide Captive bears are largely lethargic and spend a considerable amount of time doing nothing When active captive bears may be subject to repetitive back and forth motion known widely as pacing This behavior is most prevalent in bears kept in small cramped cages often with no natural setting Pacing is a way of coping with stress that comes with being trapped in unnaturally small spaces 162 Stereotyped behavior has decreased due to better and larger enclosures being built and more sustainable management from zoo staff 163 They are also exploited as dancing bears The process begins at an early age Cubs for example are positioned on hot metal plates causing them to exhibit dancing movements and at the same time violin music is running in the background The process is repeated resulting in bears being trained to dance when a violin is played Similarly brown bears are also displayed in tiny enclosures near a restaurant mainly for the purpose of luring customers Privately owned bears are often placed in insufficient environments and often suffer from malnutrition and obesity 164 According to a 2009 analysis the brown bear was the second most exploited circus animal after the tiger 165 Culture Further information Cultural depictions of bears nbsp The Story of the Three Bears illustration from Childhood s Favorites and Fairy Stories nbsp Cover of an American humor magazine Puck at 20 January 1904 Brown bears often figure into the literature of Europe and North America as cute and cuddly 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 166 In Ancient Greek mythology bears have been compared to humans as similar mainly due to their ability of standing upright In many western stories and older fables the portrayed attributes of bears are sluggishness foolishness and gullibility which contradicts actual species For example bears have been reported tricking hunters by backtracking in the snow 167 The Romans built small carved figures of bears that were used during burials of infants 168 The earliest cave paintings of bears occurred in the Paleolithic with over 100 recorded paintings 169 The Russian bear is a common national personification for Russia as well as the former Soviet Union despite the country having no officially designated national animal The brown bear is Finland s national animal 170 171 The grizzly bear is the state animal of Montana 172 The California golden bear is the state animal of California despite being extinct 173 In North America the brown bear is considered a charismatic megafauna and has long piqued people s interest The death of Bear 148 at the hands of a trophy hunter in 2017 sparked media outrage and the continued disapproval of trophy hunting 174 The coat of arms of Madrid depicts a bear reaching up into a madrono 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 175 176 The brown bear is depicted on the reverse of the Croatian 5 kuna coin minted since 1993 177 References a b McLellan B N Proctor M F Huber D Michel S 2017 Ursus arctos IUCN Red List of Threatened Species 2017 e T41688A121229971 doi 10 2305 IUCN UK 2017 3 RLTS T41688A121229971 en Retrieved 19 November 2021 Appendices CITES cites org Archived from the original on 5 December 2017 Retrieved 14 January 2022 Harper Douglas bruin Online Etymology Dictionary Bruin Archived 20 September 2022 at the Wayback Machine Encyclopaedia Britannica Hunting the Grisly and other Sketches Archived 6 September 2017 at the Wayback Machine FullTextArchive com Retrieved 15 September 2011 Liddell Henry George and Scott Robert Ursus Archived 1 March 2021 at the Wayback Machine A Latin Dictionary Perseus Digital Library Liddell Henry George and Scott Robert Arktos Archived 30 June 2023 at the Wayback Machine A Greek English Lexicon Perseus Digital Library Yu Li Li Yi Wei Ryder Oliver A Zhang Ya Ping 2007 Analysis of complete mitochondrial genome sequences increases phylogenetic resolution of bears Ursidae a mammalian family that experienced rapid speciation BMC Evolutionary Biology 7 198 198 Bibcode 2007BMCEE 7 198Y doi 10 1186 1471 2148 7 198 PMC 2151078 PMID 17956639 Servheen C Herrero S Peyton B 1999 Bears Status Survey and Conservation Action Plan PDF IUCN pp 26 30 ISBN 978 2 8317 0462 3 Archived PDF from the original on 9 October 2022 a b Kumar V Lammers F Bidon T Pfenninger M Kolter L Nilsson M A Janke A 2017 The evolutionary history of bears is characterized by gene flow across species Scientific Reports 7 46487 Bibcode 2017NatSR 746487K doi 10 1038 srep46487 PMC 5395953 PMID 28422140 Kumar V Lammers F Bidon T Pfenninger M Kolter L Nilsson M A Janke A 2017 The evolutionary history of bears is characterized by gene flow across species Scientific Reports 7 46487 Bibcode 2017NatSR 746487K doi 10 1038 srep46487 PMC 5395953 PMID 28422140 a b Steffen Martina L Fulton Tara L 1 February 2018 On the association of giant short faced bear Arctodus simus and brown bear Ursus arctos in late Pleistocene North America Geobios 51 1 61 74 Bibcode 2018Geobi 51 61S doi 10 1016 j geobios 2017 12 001 a b c d McLellan Bruce Reiner David C 1994 A Review of bear evolution PDF Int Conf Bear Res And Manage 9 1 85 96 doi 10 2307 3872687 JSTOR 3872687 Archived PDF from the original on 9 October 2022 Perez Hidalgo T 1992 The European descendants of Ursus etruscus C Cuvier Mammalia Carnivora Ursidae PDF Boletin del Instituto Geologico y Minero de Espana 103 4 632 642 Archived PDF from the original on 9 October 2022 Kurten Bjorn 1976 The Cave Bear Story New York Columbia University Press p 5 Loreille O Orlando L Patou Mathis M Philippe M Taberlet P Hanni C 2001 Ancient DNA analysis reveals divergence of the cave bear Ursus spelaeus and brown bear Ursus arctos lineages Current Biology 11 3 200 203 Bibcode 2001CBio 11 200L doi 10 1016 s0960 9822 01 00046 x PMID 11231157 S2CID 14645603 a b Herrero S 1972 Aspects of evolution and adaptation in American black bears Ursus americanus Pallas and brown and grizzly bears U arctos Linne of North America PDF Bears Their Biology and Management 2 221 231 doi 10 2307 3872586 JSTOR 3872586 Archived PDF from the original on 9 October 2022 Kosintsev Pavel A Bocherens Herve Kirillova Irina V Levchenko Vladimir A Zazovskaya Elya P Trofimova Svetlana S Lan Tianying Lindqvist Charlotte 5 December 2024 Palaeoecological and genetic analyses of Late Pleistocene bears in Asiatic Russia Boreas 51 2 465 480 doi 10 1111 bor 12570 ISSN 0300 9483 Retrieved 4 May 2024 via Wiley Online Library Monchot Herve Mashkour Marjan Biglari Fereidoun Abdi Kamyar April June 2020 The Upper Pleistocene brown bear Carnivora Ursidae in the Zagros Evidence from Wezmeh Cave Kermanshah Iran Annales de Paleontologie 106 2 102381 doi 10 1016 j annpal 2019 102381 Retrieved 9 May 2024 via Elsevier Science Direct a b McLellan Bruce Reiner David C 1994 A Review of bear evolution PDF Int Conf Bear Res And Manage 9 1 85 96 doi 10 2307 3872687 JSTOR 3872687 Archived PDF from the original on 9 October 2022 a b c Salis Alexander T Bray Sarah C E Lee Michael S Y Heiniger Holly Barnett Ross Burns James A Doronichev Vladimir Fedje Daryl Golovanova Liubov Harington C Richard Hockett Bryan Kosintsev Pavel Lai Xulong Mackie Quentin Vasiliev Sergei December 2022 Lions and brown bears colonized North America in multiple synchronous waves of dispersal across the Bering Land Bridge Molecular Ecology 31 24 6407 6421 Bibcode 2022MolEc 31 6407S doi 10 1111 mec 16267 hdl 11343 299180 ISSN 0962 1083 PMID 34748674 Kubiak Cara Grimes Vaughan Van Biesen Geert Keddie Grant Buckley Mike Macdonald Reba Richards M P 27 June 2022 Dietary niche separation of three Late Pleistocene bear species from Vancouver Island on the Pacific Northwest Coast of North America Journal of Quaternary Science 38 8 20 doi 10 1002 jqs 3451 ISSN 0267 8179 S2CID 250134103 Archived from the original on 31 October 2023 Retrieved 2 January 2024 a b c d e f g h i j k l m n o p q r Mammalian Species Ursus arctos PDF American Society of Mammalogists Smith College 23 April 1993 Archived from the original PDF on 31 March 2017 Wilson D E Ruff S 1999 The Smithsonian Book of North American Mammals Washington D C Smithsonian Institution Press pp 160 201 Harris Arthur H 2013 Pleistocene Vertebrates of Arizona New Mexico and Trans Pecos Texas UTEP Biodiversity Collections University of Texas at El Paso Storer T I Tevis L P 1996 California Grizzly Berkeley CA University of California Press pp 335 42 187 ISBN 978 0 520 20520 8 Archived from the original on 2 February 2024 Retrieved 18 November 2019 Alt URL 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 70 221 69854 69884 Archived from the original PDF on 25 August 2006 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 PDF Molecular Ecology 17 8 1962 1970 Bibcode 2008MolEc 17 1962C doi 10 1111 j 1365 294x 2008 03631 x PMID 18363668 S2CID 23361337 Archived PDF from the original on 9 October 2022 Lan T Gill S Bellemain E Bischof R Zawaz M A Lindqvist C 2017 Evolutionary history of enigmatic bears in the Tibetan Plateau Himalaya region and the identity of the yeti Proceedings of the Royal Society B Biological Sciences 284 1 868 20 171 804 doi 10 1098 rspb 2017 1804 PMC 5740279 PMID 29187630 Wozencraft W C 2005 Ursus arctos In Wilson D E Reeder D M eds 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 Ursus arctos Archived 22 January 2019 at the Wayback Machine ITIS Lindqvist C Schuster S C Sun Y et al 2010 Complete mitochondrial genome of a Pleistocene jawbone unveils the origin of polar bear Proceedings of the National Academy of Sciences 107 11 5053 5057 Bibcode 2010PNAS 107 5053L doi 10 1073 pnas 0914266107 PMC 2841953 PMID 20194737 Liu Shiping Lorenzen Eline D Fumagalli Matteo Li Bo Harris Kelley Xiong Zijun Zhou Long Korneliussen Thorfinn Sand Somel Mehmet Babbitt Courtney Wray Greg Li Jianwen He Weiming Wang Zhuo Fu Wenjing Xiang Xueyan Morgan Claire C Doherty Aoife o Connell Mary J McInerney James O Born Erik W Dalen Love Dietz Rune Orlando Ludovic Sonne Christian Zhang Guojie Nielsen Rasmus Willerslev Eske Wang Jun 2014 Population Genomics Reveal Recent Speciation and Rapid Evolutionary Adaptation in Polar Bears Cell 157 4 785 794 doi 10 1016 j cell 2014 03 054 PMC 4089990 PMID 24813606 Proctor Michael F Paetkau David Mclellan Bruce N Stenhouse Gordon B Kendall Katherine C Mace Richard D Kasworm Wayne F Servheen Christopher Lausen Cori L 2012 Population fragmentation and inter ecosystem movements of grizzly bears in western Canada and the northern United States Wildlife Monographs 180 1 1 46 Bibcode 2012WildM 180 1P doi 10 1002 wmon 6 ISSN 1938 5455 S2CID 16790669 Strobeck Curtis Craighead Lance Clarkson Peter L Waits Lisette P Paetkau David 1 December 1997 An Empirical Evaluation of Genetic Distance Statistics Using Microsatellite Data From Bear Ursidae Populations Genetics 147 4 1943 1957 doi 10 1093 genetics 147 4 1943 ISSN 0016 6731 PMC 1208359 PMID 9409849 Archived from the original on 29 January 2019 Retrieved 29 January 2019 Waits L P et al 1998 Mitochondrial DNA Phylogeography of the North American Brown Bear and Implications for Conservation Conservation 12 2 408 417 Bibcode 1998ConBi 12 408W doi 10 1111 j 1523 1739 1998 96351 x S2CID 86172292 Paetkau D Shields G F Strobeck C 1998 Gene flow between insular coastal and interior populations of brown bears in Alaska Molecular Ecology 7 10 1283 1292 Bibcode 1998MolEc 7 1283P doi 10 1046 j 1365 294x 1998 00440 x ISSN 0962 1083 PMID 9787441 S2CID 21848010 Shapiro Beth Slatkin Montgomery Stirling Ian John John St Salamzade Rauf Ovsyanikov Nikita Jay Flora Stiller Mathias Fulton Tara L 14 March 2013 Genomic Evidence for Island Population Conversion Resolves Conflicting Theories of Polar Bear Evolution PLOS Genetics 9 3 e1003345 doi 10 1371 journal pgen 1003345 ISSN 1553 7404 PMC 3597504 PMID 23516372 a b Servheen C Herrero S Peyton B Pelletier K Moll K Moll J eds 1999 Bears status survey and conservation action plan PDF vol 44 Gland Switzerland IUCN archived from the original PDF on 23 September 2015 retrieved 18 November 2019 Richardson Evan Branigan Marsha Paetkau David Pongracz Jodie D 31 May 2017 Recent Hybridization between a Polar Bear and Grizzly Bears in the Canadian Arctic Arctic 70 2 151 160 doi 10 14430 arctic4643 ISSN 1923 1245 Janke Axel Nilsson Maria A Kolter Lydia Pfenninger Markus Bidon Tobias Lammers Fritjof Kumar Vikas 19 April 2017 The evolutionary history of bears is characterized by gene flow across species Scientific Reports 7 46 487 Bibcode 2017NatSR 746487K doi 10 1038 srep46487 ISSN 2045 2322 PMC 5395953 PMID 28422140 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 6 079 344 247 Bibcode 2012Sci 336 344H doi 10 1126 science 1216424 hdl 10261 58578 PMID 22517859 S2CID 12671275 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 2390 Bibcode 2012PNAS 109E2382M doi 10 1073 pnas 1210506109 PMC 3437856 PMID 22826254 Hissa R Siekkinen J Hohtola E Saarela S Hakala A Pudas J 1994 Seasonal patterns in the physiology of the European brown bear Ursus arctos arctos in Finland Comparative Biochemistry and Physiology Part A Physiology 109 3 781 791 doi 10 1016 0300 9629 94 90222 4 PMID 8529017 Archived from the original on 15 June 2020 Retrieved 12 December 2016 McLellan B N 2011 Implications of a high energy and low protein diet on the body composition fitness and competitive abilities of black Ursus americanus and grizzly Ursus arctos bears Canadian Journal of Zoology 89 6 546 558 doi 10 1139 z11 026 Maria Pasitschniak Arts 23 April 1993 Ursus arctos PDF Mammalian Species 439 1 10 doi 10 2307 3504138 JSTOR 3504138 Archived PDF from the original on 5 March 2023 Retrieved 5 March 2023 Grzimek Bernhard 1990 Grzimek s Encyclopedia of Mammals McGraw Hill pp 52 53 ISBN 978 0 07 909508 4 a b c Nowak Ronald M 1991 Walker s Mammals of the World Johns Hopkins University Press pp 1083 1093 ISBN 978 0 8018 3970 2 Christiansen P 1999 What size were Arctodus simus and Ursus spelaeus Carnivora Ursidae Annales Zoologici Fennici 36 2 93 102 JSTOR 23735739 a b c d e Feldhamer G A Thompson B C Chapman J A 2003 Wild mammals of North America biology management and conservation JHU Press ISBN 9780801874161 Archived from the original on 2 February 2024 Retrieved 9 November 2020 Swenson Jon E Adamic Miha Huber Djuro Stokke Sigbjorn 2007 Brown bear body mass and growth in northern and southern Europe PDF Oecologia 153 1 37 47 Bibcode 2007Oecol 153 37S doi 10 1007 s00442 007 0715 1 PMID 17415593 S2CID 5836340 Archived PDF from the original on 9 October 2022 a b Jones S V 1923 Color variations in wild animals Journal of Mammalogy 4 3 172 177 doi 10 2307 1373567 JSTOR 1373567 Swenson J E 2000 Action plan for the conservation of the brown bear in Europe Ursus arctos Archived 20 December 2016 at the Wayback Machine No 18 114 Council of Europe Sahajpal V Goyal S P Jayapal R Yoganand K Thakar M K 2008 Hair characteristics of four Indian bear species Science amp Justice 48 1 8 15 doi 10 1016 j scijus 2007 05 001 PMID 18450212 a b c d e f g h i j k l m n o p Brown Gary 1993 Bear Anatomy and Physiology The great bear almanac Internet Archive New York Lyons amp Burford ISBN 978 1 55821 210 7 a b c d e f g h V G Heptner N P Naumov eds 1998 Mlekopitajuscie Sovetskogo Soiuza Moskva Vyssaia Skola Mammals of the Soviet Union Volume II Part 1a Sirenia and Carnivora Sea cows Wolves and Bears Vol II Part 1a Washington D C US Science Publishers Inc pp 601 676 ISBN 1 886106 81 9 Christiansen Per 2008 Feeding ecology and morphology of the upper canines in bears carnivora Ursidae Journal of Morphology 269 7 896 908 doi 10 1002 jmor 10643 PMID 18488989 S2CID 33532210 Kurten B 1966 Pleistocene bears of North America Genus Tremarctos spectacled bears Acta Zoologica Fennica 115 1 96 Sacco Tyson Van Valkenburgh Blaire 2004 Ecomorphological indicators of feeding behaviour in the bears Carnivora Ursidae Journal of Zoology 263 41 doi 10 1017 S0952836904004856 Seryodkin Ivan V 1 June 2016 Behavior of Brown Bears During Feeding in the Sikhote Alin Achievements in the Life Sciences 10 1 38 47 doi 10 1016 j als 2016 05 003 ISSN 2078 1520 Stirling Ian 1988 Polar Bears University of Michigan Press pp 20 25 ISBN 978 0 472 10100 9 a b c d e f g h i j k l Hunter Luke 2011 Carnivores of the world Internet Archive Princeton NJ Princeton University Press pp 122 133 ISBN 978 0 691 15228 8 Ellig Tracy 3 July 2006 MSU researcher tests grizzly bear strength for National Geographic documentary Montana State University Archived from the original on 1 June 2013 Retrieved 11 June 2021 Ancient bear made early migration BBC News 12 November 2004 Archived from the original on 2 December 2005 Retrieved 2 January 2010 Brown Bear Species WWF WWF Archived from the original on 29 September 2018 Retrieved 2 December 2013 Chestin Igor E Gubar Yuliy P Sokolov Vladimir E Lobachev Vladimir S 1992 The brown bear Ursus arctos L in the USSR numbers hunting and systematics Annales Zoologici Fennici 29 2 57 68 ISSN 0003 455X JSTOR 23735466 Brown Bear Research in Alaska Alaska Department of Fish and Game Archived from the original on 3 May 2012 Retrieved 4 April 2012 Neixen quatre cries d os bru als Pirineus elperiodico cat in Catalan and Spanish 2 August 2010 Archived from the original on 24 July 2011 Retrieved 19 November 2010 Situacio preocupant de l os bru als Pirineus in Catalan ecologistasenaccion org May 2010 Archived from the original on 13 November 2012 Retrieved 19 November 2010 Osos el desafio de una poblacion creciente ABC in Spanish Spain 28 September 2013 Archived from the original on 26 October 2013 Retrieved 14 October 2013 Martinez Abrain A Jimenez J Oro D 2018 Pax Romana refuge abandonment and spread of fearless behavior in a reconciling world Animal Conservation 22 1 John Wiley amp Sons Inc Zoological Society of London ZSL 3 13 doi 10 1111 acv 12429 ISSN 1367 9430 S2CID 89938098 Bear ARKTOYROS Archived from the original on 2 February 2024 Retrieved 13 October 2022 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 Bibcode 2006MolEc 15 4477M doi 10 1111 j 1365 294x 2006 03097 x PMID 17107477 S2CID 7336900 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 Bibcode 2009DivDi 15 742C doi 10 1111 j 1472 4642 2009 00586 x S2CID 21666120 Himalayan brown bears now critically endangered Archived 16 October 2018 at the Wayback Machine Euronews com 6 January 2014 Gervasi V Ciucci P 1 January 2018 Demographic projections of the Apennine brown bear population Ursus arctos marsicanus Mammalia Ursidae under alternative management scenarios The European Zoological Journal 85 1 242 252 doi 10 1080 24750263 2018 1478003 hdl 11573 1114254 ISSN 2475 0263 Ordiz Andres Kindberg Jonas Saebo Solve Swenson Jon E Stoen Ole Gunnar 1 May 2014 Brown bear circadian behavior reveals human environmental encroachment Biological Conservation 173 1 9 Bibcode 2014BCons 173 1O doi 10 1016 j biocon 2014 03 006 ISSN 0006 3207 Klinka D R Reimchen T E 2002 Nocturnal and diurnal foraging behaviour of brown bears Ursus arctos on a salmon stream in coastal British Columbia PDF Canadian Journal of Zoology 80 8 1317 1322 doi 10 1139 z02 123 Archived PDF from the original on 9 October 2022 Moe T F Kindberg J Jansson I Swenson J E 2007 Importance of diel behaviour when studying habitat selection examples from female Scandinavian brown bears Ursus arctos Canadian Journal of Zoology 85 4 518 525 doi 10 1139 Z07 034 Archived from the original on 15 June 2020 Retrieved 12 December 2016 Kaczensky P Huber D Knauer F Roth H Wagner A Kusak J 2006 Activity patterns of brown bears Ursus arctos in Slovenia and Croatia Journal of Zoology 269 4 474 485 doi 10 1111 j 1469 7998 2006 00114 x Kingsley M C S Nagy J A Russell R H 1983 Patterns of weight gain and loss for grizzly bears in northern Canada Bears Their Biology and Management 5 174 178 doi 10 2307 3872535 JSTOR 3872535 S2CID 90555276 Hissa R Hohtola E Tuomala Saramaki T Laine T 1998 Seasonal changes in fatty acids and leptin contents in the plasma of the European brown bear Ursus arctos arctos Annales Zoologici Fennici 35 4 215 224 JSTOR 23735612 Farley S D Robbins C T 1995 Lactation hibernation and mass dynamics of American black bears and grizzly bears Canadian Journal of Zoology 73 12 2216 2222 doi 10 1139 z95 262 Evans A L Sahlen V Stoen O G Fahlman A Brunberg S Madslien K Forbert O Swenson J E Arnemo J M 2012 Capture anesthesia and disturbance of free ranging brown bears Ursus arctos during hibernation PLOS ONE 7 7 e40520 Bibcode 2012PLoSO 740520E doi 10 1371 journal pone 0040520 PMC 3398017 PMID 22815757 Deecke V B 2012 Tool use in the brown bear Ursus arctos PDF Animal Cognition 15 4 725 730 doi 10 1007 s10071 012 0475 0 PMID 22367156 S2CID 9076161 Archived PDF from the original on 9 October 2022 a b Egbert Allan L Stokes Allen W Egbert A L 16 September 1976 The Social Behaviour of Brown Bears on an Alaskan Salmon Stream PDF Bears Their Biology and Management 3 41 56 doi 10 2307 3872753 JSTOR 3872753 Archived from the original PDF on 17 July 2015 Retrieved 29 April 2011 Sandell M 1989 The mating tactics and spacing patterns of solitary carnivores Carnivore behavior ecology and evolution Springer pp 164 182 doi 10 1007 978 1 4613 0855 3 7 ISBN 978 1 4613 0855 3 Gau R J McLoughlin P D Case R Cluff H D Mulders R Messier F 2004 Movements of subadult male grizzly bears Ursus arctos in the central Canadian arctic The Canadian Field Naturalist 118 2 239 242 doi 10 22621 cfn v118i2 920 a b c d Dahle B Swenson J E 2003 Seasonal range size in relation to reproductive strategies in brown bears Ursus arctos Journal of Animal Ecology 72 4 660 667 Bibcode 2003JAnEc 72 660D doi 10 1046 j 1365 2656 2003 00737 x JSTOR 3505643 PMID 30893970 S2CID 67818528 a b c Egbert A L 1978 The Social Behavior of Brown Bears at McNeil River Alaska Ph D thesis Utah State University pp 30 35 Archived from the original on 2 February 2024 Retrieved 25 April 2021 a b Pearson A M 1975 The northern interior grizzly bear Ursus arctos L Information Canada p 86 a b c d e Bellemain Eva Swenson Jon E Taberlet Pierre 2006 Mating Strategies in Relation to Sexually Selected Infanticide in a Non Social Carnivore The Brown Bear PDF Ethology 112 3 238 246 Bibcode 2006Ethol 112 238B doi 10 1111 j 1439 0310 2006 01152 x Archived PDF from the original on 9 October 2022 Pop Ioan Mihai Bereczky Leonardo Chiriac Silviu Iosif Ruben Nita Andreea Popescu Viorel Dan Rozylowicz Laurențiu 2018 Movement ecology of brown bears Ursus arctos in the Romanian Eastern Carpathians Nature Conservation 26 15 31 doi 10 3897 natureconservation 26 22955 ISSN 1314 3301 a b Novak M Baker J A Obbard M E amp Malloch B 1987 Wild furbearer management and conservation in North America Ontario Ministry of Natural Resources Herrero S Hamer D 1977 Courtship and copulation of a pair of grizzly bears with comments on reproductive plasticity and strategy Journal of Mammalogy 58 3 441 444 doi 10 2307 1379352 JSTOR 1379352 a b Craighead J J Sumner J S amp Mitchell J A 1995 The grizzly bears of Yellowstone their ecology in the Yellowstone ecosystem 1959 1992 Island Press pp 21 56 a b White D J Berardinelli J G Aune K E 1998 Reproductive characteristics of the male grizzly bear in the continental United States Ursus 10 497 501 JSTOR 3873161 Ambarli H 2016 Litter size and basic diet of brown bears Ursus arctos Carnivora in northeastern Turkey Mammalia 80 2 doi 10 1515 mammalia 2014 0111 S2CID 87968464 Halloran D W Pearson A M 1972 Blood chemistry of the brown bear Ursus arctos from southwestern Yukon Territory Canada Canadian Journal of Zoology 50 6 827 833 doi 10 1139 z72 112 PMID 5038730 Craighead J J Hornocker M G Craighead Jr F C 1969 Reproductive biology of young female grizzly bears J Reprod Fertil Suppl 6 447 475 Corbet Gordon Barclay 1966 The Terrestrial Mammals of Western Europe London UK Foulis pp 52 55 Tsubota T amp Kanagawa H 1993 Morphological Characteristics of the Ovary Uterus and Embryo during the Delayed Implantation Period in the Hokkaido Brown Bear Ursus arctos yesoensis Journal of Reproduction and Development 39 4 325 331 doi 10 1262 jrd 39 325 S2CID 84359252 Hensel R J Troyer W A amp Erickson A W 1969 Reproduction in the female brown bear The Journal of Wildlife Management 33 2 357 365 doi 10 2307 3799836 JSTOR 3799836 Pazetnov V A amp Pazetnov S V 2005 Female brown bear with six cubs PDF International Bear News 14 2 17 Archived PDF from the original on 9 October 2022 Erickson A W Miller L H 1963 Cub adoption in the brown bear Journal of Mammalogy 44 4 584 585 doi 10 2307 1377153 JSTOR 1377153 Barnes Jr V Smith R 1993 Cub adoption by brown bears Ursus arctos middendorffi on Kodiak Island Alaska Canadian Field Naturalist 107 3 365 367 doi 10 5962 p 357155 Archived from the original on 20 December 2016 Retrieved 12 December 2016 Stringham S F 1990 Grizzly bear reproductive rate relative to body size PDF Bears Their Biology and Management 8 433 443 doi 10 2307 3872948 JSTOR 3872948 Archived PDF from the original on 9 October 2022 Steyaert S M Endrestol A Hacklaender K Swenson J E Zedrosser A 2012 The mating system of the brown bear Ursus arctos Mammal Review 42 1 12 34 doi 10 1111 j 1365 2907 2011 00184 x Dahle B Zedrosser A Swenson J E 2006 Correlates with body size and mass in yearling brown bears Ursus arctos Journal of Zoology 269 3 273 283 doi 10 1111 j 1469 7998 2006 00127 x Gittleman J L 1994 Female brain size and parental care in carnivores Proceedings of the National Academy of Sciences 91 12 5495 5497 Bibcode 1994PNAS 91 5495G doi 10 1073 pnas 91 12 5495 PMC 44022 PMID 8202515 Mano T Tsubota T 2002 Reproductive characteristics of brown bears on the Oshima peninsula Hokkaido Japan Journal of Mammalogy 83 4 1026 1034 doi 10 1644 1545 1542 2002 083 lt 1026 RCOBBO gt 2 0 CO 2 JSTOR 1383508 Dahle B Swenson J E 2003 Family breakup in brown bears are young forced to leave Journal of Mammalogy 84 2 536 540 doi 10 1644 1545 1542 2003 084 lt 0536 FBIBBA gt 2 0 CO 2 Swenson J E Dahle B Sandegren F 2001 Intraspecific predation in Scandinavian brown bears older than cubs of the year Ursus 12 81 91 JSTOR 3873233 Archived from the original on 16 November 2017 Retrieved 12 December 2016 Morner T Eriksson H Brojer C Nilsson K Uhlhorn H Agren E Segerstad C H Jansson D S Gavier Widen D 2005 Diseases and mortality in free ranging brown bear Ursus arctos gray wolf Canis lupus and wolverine Gulo gulo in Sweden Journal of Wildlife Diseases 41 2 298 303 doi 10 7589 0090 3558 41 2 298 PMID 16107663 S2CID 43774546 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 a b Ursus arctos Brown bear Animal Diversity Web Archived from the original on 13 May 2021 Retrieved 28 May 2021 Yellowstone Grizzly Bears Eat 40 000 Moths a Day In August Yellowstonepark com 21 June 2011 Archived from the original on 15 July 2010 Brown Susan A 17 January 2010 Inherited behavior traits of the domesticated ferret weaselwords com Archived from the original on 4 October 2009 Retrieved 29 January 2010 a href Template Cite web html title Template Cite web cite web a CS1 maint multiple names authors list link 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 Archived PDF from the original on 9 October 2022 a b Macdonald D W Barrett P 1993 Mammals of Europe New Jersey Princeton University Press p 107 ISBN 978 0 691 09160 0 Fraser A F 2012 Feline Behaviour and Welfare CABI pp 72 77 ISBN 978 1 84593 926 7 Seryodkin I V Kostyria A V Goodrich J M Miquelle D G Smirnov E N Kerley L L amp Hornocker M G 2003 Denning ecology of brown bears and Asiatic black bears in the Russian Far East Ursus 14 2 159 Archived from the original on 17 August 2011 Retrieved 15 September 2014 a b c Seryodkin I V Goodrich J M Kostyrya A V Schleyer B O Smirnov E N Kerley L L amp Miquelle D G 2005 Glava 19 Vzaimootnosheniya amurskogo tigra s burym i gimalajskim medvedyami Chapter 19 Relationship of Amur tigers with brown and Himalayan black bear In Miquelle D G Smirnov E N amp Goodrich J M eds Tigers of Sikhote Alin Zapovednik Ecology and Conservation in Russian Vladivostok Russia PSP pp 156 163 Archived from the original on 18 October 2020 Retrieved 19 March 2023 Seryodkin I 2006 The ecology behavior management and conservation status of brown bears in Sikhote Alin PhD in Russian Vladivostok Russia Far Eastern National University pp 61 73 Archived from the original on 24 December 2013 Yudakov A G amp 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 Institute of Biology and Soil Science Far Eastern Scientific Center Academy of Sciences of the USSR Archived from the original on 19 February 2020 Retrieved 17 September 2014 Seryodkin I V Goodrich J M Kostyria A V Smirnov E N amp Miquelle D G 2011 Intraspecific relationships between brown bears Asiatic black bears and the Amur tiger PDF 20th International Conference on Bear Research amp Management International Association for Bear Research and Management p 64 Archived PDF from the original on 13 November 2018 Retrieved 11 September 2014 Seryodkin I V 2007 Rol burogo medvedya v ekosistemah Dalnego Vostoka Rossii Bioraznoobrazie i rol zhivotnyh v ekosistemah Materialy IV Mezhdunarodnoj nauchnoj konferencii Denpropetrovsk Oles Honchar Dnipro National University pp 502 503 Archived from the original on 17 August 2011 Retrieved 15 July 2015 Brown Bear predation of Amur Tiger 1973 account International Wildlife Magazine 20 October 2009 Archived from the original on 22 June 2016 Retrieved 14 July 2015 Goodrich J M Kerley L L Smirnov E N Miquelle D G McDonald L Quigley H B Hornocker M G amp McDonald T 2008 Survival rates and causes of mortality of Amur tigers on and near the Sikhote Alin Biosphere Zapovednik Journal of Zoology 276 4 323 doi 10 1111 j 1469 7998 2008 00458 x 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 doi 10 22621 cfn v122i1 550 PDF Archived 19 January 2021 at the Wayback Machine Downey Betsy Personal Encounter Wolf Grizzly interaction in Yellowstone National Park PDF International Wolf Center Archived from the original PDF on 27 February 2008 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 ADW Ursus arctos Information Archived 12 May 2013 at the Wayback Machine Arlis org Retrieved 9 August 2012 Hornocker Maurice Negri Sharon 15 December 2009 Cougar Ecology and Conservation University of Chicago Press pp 163 174 ISBN 978 0 226 35347 0 Sorensen Ole Mogens Totsas 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 S2CID 55281438 Archived PDF from the original on 9 October 2022 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 Archived PDF from the original on 9 October 2022 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 Archived PDF from the original on 9 October 2022 Dough O Hara Polar bears grizzlies increasingly gather on North Slope Anchorage Daily News 24 April 2005 ABC News Grizzlies Encroaching on Polar Bear Country ABC News Archived from the original on 5 August 2017 Retrieved 10 October 2009 Schwartz C C Keating K A Reynolds III H V Barnes Jr V G Sellers R A Swenson J E Miller S D McLellan B Keay J McCann R Gibeau M Wakkinen W F Mace R D Kasworm W Smith R Herrero S 2003 Reproductive maturation and senescence in the female brown bear Ursus 14 2 109 119 JSTOR 3873012 Macdonald David David Whyte 2001 Bear family The New Encyclopedia of Mammals Oxford University Press Incorporated ISBN 978 0 19 850823 6 Schwartz C C Haroldson M A White G C Harris R B Cherry S Keating K A Moody D Servheen C 2006 Temporal Spatial and Environmental Influences on the Demographics of Grizzly Bears in the Greater Yellowstone Ecosystem Wildlife Monographs 161 1 68 doi 10 2193 0084 0173 2006 161 1 TSAEIO 2 0 CO 2 S2CID 85600938 Rogers Lynn L Rogers Susanne M 1976 Parasites of Bears A Review Bears Their Biology and Management 3 411 430 doi 10 2307 3872791 ISSN 1936 0614 JSTOR 3872791 J Brglez S Valentinclc 1968 Parasites of brown bear Ursus arctos L Acta Veterinaria Beograd 18 6 379 384 Di Francesco Cristina Esmeralda Gentile Leonardo Di Pirro Vincenza Ladiana Lara Tagliabue Silvia Marsilio Fulvio 1 January 2015 Serologic Evidence for Selected Infectious Diseases in Marsican Brown Bears Ursus arctos marsicanus in Italy 2004 09 Journal of Wildlife Diseases 51 1 209 213 doi 10 7589 2014 01 021 ISSN 0090 3558 PMID 25375945 Morner Torsten Eriksson Hanna Brojer Caroline Nilsson Kristina Uhlhorn Henrik Agren Erik Segerstad Carl Hard af Jansson Desiree S Gavier Widen Dolores April 2005 Diseases and mortality in free ranging brown bear Ursus arctos gray wolf Canis lupus and wolverine Gulo gulo in Sweden Journal of Wildlife Diseases 41 2 298 303 doi 10 7589 0090 3558 41 2 298 ISSN 0090 3558 PMID 16107663 a b c d e Frobert O Frobert A M Kindberg J Arnemo J M Overgaard M T March 2020 The brown bear as a translational model for sedentary lifestyle related diseases Journal of Internal Medicine 287 3 263 270 doi 10 1111 joim 12983 hdl 11250 2642746 ISSN 0954 6820 PMID 31595572 Woodroffe R 2000 Predators and people Using human densities to interpret declines of large carnivores PDF Animal Conservation 3 2 165 Bibcode 2000AnCon 3 165W doi 10 1111 j 1469 1795 2000 tb00241 x S2CID 84430290 Archived PDF from the original on 9 October 2022 Kistchinski A A 1972 Life history of the brown bear Ursus arctos L in north east Siberia Bears Their Biology and Management 2 67 73 doi 10 2307 3872570 JSTOR 3872570 a b c Herrero Stephen 2002 Bear Attacks Their Causes and Avoidance Globe Pequot Press pp 10 25 ISBN 978 1 58574 557 9 Smith Tom S Herrero Steven Ursus arctos californicus Alaska Science Center Biological Science Office Archived from the original on 14 August 2009 Rogers Lynn L How Dangerous are Black Bears Bear org archived from the original on 16 October 2002 Cardall T Y Rosen P 2003 Grizzly bear attack The Journal of Emergency Medicine 24 3 331 333 doi 10 1016 s0736 4679 03 00004 0 PMID 12676309 a b Bombieri G Naves J Penteriani V Selva N Fernandez Gil A Lopez Bao J V Ambarli H Bautista C Bespalova T Bobrov V Bolshakov V Bondarchuk S Camarra J J Chiriac S Ciucci P 12 June 2019 Brown bear attacks on humans a worldwide perspective Scientific Reports 9 1 8573 Bibcode 2019NatSR 9 8573B doi 10 1038 s41598 019 44341 w ISSN 2045 2322 PMC 6562097 PMID 31189927 Dinets Vladimir Brown Bears of Russia Archived from the original on 15 December 2012 Dickman A J Hazzah L 2016 Money Myths and Man Eaters Complexities of Human Wildlife Conflict Problematic Wildlife Springer pp 339 356 ISBN 978 3 319 22246 2 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 McLaren Duncan amp Wigen Rebecca amp Mackie Quentin amp Fedje Daryl 2005 Bear Hunting at the Pleistocene Holocene Transition on the Northern Northwest Coast of North America pp 4 23 Bischof Richard Bonenfant Christophe Rivrud Inger Maren Zedrosser Andreas Friebe Andrea Coulson Tim Mysterud Atle Swenson Jon E January 2018 Regulated hunting re shapes the life history of brown bears Nature Ecology amp Evolution 2 1 116 123 doi 10 1038 s41559 017 0400 7 hdl 10852 67379 ISSN 2397 334X PMID 29230025 S2CID 3288368 Albert David M Bowyer R Terry Miller Sterling D 2001 Effort and Success of Brown Bear Hunters in Alaska Wildlife Society Bulletin 29 2 501 508 ISSN 0091 7648 JSTOR 3784174 Kelava Ugarkovic Nikolina Konjacic Miljenko Malnar Josip Tomljanovic Kristijan Sprem Nikica Ugarkovic Damir January 2021 Proximate Chemical Composition Fatty Acid Profile and Lipid Qualitative Indices of Brown Bear Meat Foods 10 1 36 doi 10 3390 foods10010036 ISSN 2304 8158 PMC 7824718 PMID 33374425 Pastorino Giovanni Quintavalle Christodoulides Yiannis Curone Giulio Pearce Kelly Paul Faustini Massimo Albertini Mariangela Preziosi Richard Mazzola Silvia Michela May 2017 Behavioural Profiles of Brown and Sloth Bears in Captivity Animals 7 5 39 doi 10 3390 ani7050039 ISSN 2076 2615 PMC 5447921 PMID 28505095 Montaudouin S Pape G Le 1 July 2005 Comparison between 28 zoological parks stereotypic and social behaviours of captive brown bears Ursus arctos Applied Animal Behaviour Science 92 1 129 141 doi 10 1016 j applanim 2004 10 015 ISSN 0168 1591 Stagni Elena Sequeira Sara Brscic Marta Redtenbacher Irene Hartmann Sabine 13 December 2023 A retrospective study on the prevalence of main clinical findings in brown bears Ursus arctos rescued from substandard husbandry conditions Frontiers in Veterinary Science 10 1299029 doi 10 3389 fvets 2023 1299029 ISSN 2297 1769 PMC 10773888 PMID 38192718 Iossa G Soulsbury C D Harris S May 2009 Are wild animals suited to a travelling circus life Animal Welfare 18 2 129 140 doi 10 1017 S0962728600000270 ISSN 0962 7286 S2CID 32259865 Newman A R 1987 Images of the bear in Children s Literature Children s Literature in Education 18 3 131 138 doi 10 1007 bf01130991 S2CID 143882256 Bieder Robert E 18 August 2005 Survivors Bear Reaktion Books ISBN 978 1 86189 482 3 O Regan Hannah 4 September 2023 Brown bears in burials and entertainment in later prehistoric to modern Britain c 2400 BC AD 1900s The Archaeology of Northern Europe 187 208 doi 10 1484 M TANE EB 5 134334 ISBN 978 2 503 60611 8 De Swart Herman 2004 Cave bears in prehistoric art a survey from the literature Publications du musee des Confluences 2 1 121 128 Karhu on Suomen kansalliselain yle fi 16 September 2010 Archived from the original on 11 January 2016 Retrieved 18 August 2016 Riabov Oleg 2020 The Birth of the Russian Bear The Bear Symbol in the Satirical Journals of the Russian Revolution of 1905 Region 9 1 139 168 ISSN 2166 4307 JSTOR 27007706 Symbols of Montana Montana Historical Society Archived from the original on 18 December 2009 Retrieved 18 August 2016 History and Culture State Symbols California State Library Archived from the original on 5 January 2019 Retrieved 24 September 2011 Hughes Courtney Foote Lee Yarmey Nicholas T Hwang Christina Thorlakson Jessica Nielsen Scott April 2020 From human invaders to problem bears A media content analysis of grizzly bear conservation Conservation Science and Practice 2 4 Bibcode 2020ConSP 2E 176H doi 10 1111 csp2 176 ISSN 2578 4854 Deyermond A D 1997 Festschrift Tamesis pp 177 187 ISBN 978 1 85566 051 9 Swenson Jon E Europe Council of 1 January 2000 Action Plan for the Conservation of the Brown Bear in Europe Ursus Arctos Council of Europe p 69 ISBN 978 92 871 4426 3 Majic Aleksandra Marino Taussig de Bodonia Agnese Huber Đuro Bunnefeld Nils December 2011 Dynamics of public attitudes toward bears and the role of bear hunting in Croatia Biological Conservation 144 12 3018 3027 Bibcode 2011BCons 144 3018M doi 10 1016 j biocon 2011 09 005 ISSN 0006 3207 Notes Populations of Bhutan China Mexico and Mongolia are included in Appendix I All other populations are included in Appendix II BibliographyVaisfeld M A and Chestin I E ed 1993 Bears Brown Bear Polar Bear Asian Black Bear Distribution ecology use and protection in Russian and English Moscow Nauka ISBN 978 5020035676 External links nbsp Wikimedia Commons has media related to Ursus arctos category nbsp Wikispecies has information related to Ursus arctos Brown Bear profile from National Geographic Bear Hunting Altered Genetics More Than Ice Age Isolation Ancient Fossil Offers New Clues To Brown Bears Past Retrieved from https en wikipedia org w index php title Brown bear amp oldid 1224601678, wikipedia, wiki, book, books, library,

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