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Brown bear

The brown bear (Ursus arctos) is a large bear species found across Eurasia and North America.[1][3] In North America, the populations of brown bears are called grizzly bears, while the subspecies that inhabits the Kodiak Islands of Alaska is known as the Kodiak bear. It is one of the largest living terrestrial members of the order Carnivora, rivaled in size only by its closest relative, the polar bear (Ursus maritimus), which is much less variable in size and slightly bigger on average.[4][5][6][7][8] The brown bear's range includes parts of Russia, Central Asia, the Himalayas, China, Canada, the United States, Hokkaido, Scandinavia, Finland, the Balkans, the Picos de Europa and the Carpathian region (especially Romania), Iran, Anatolia, and the Caucasus.[1][9] The brown bear is recognized as a national and state animal in several European countries.[10]

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
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

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. As of 2012, this and the American black bear are the only bear species not classified as threatened by the IUCN, though the large sizes of both bears may be a disadvantage due to increased competition with humans.[1][3][7] Populations that were hunted to extinction in the 19th and 20th centuries are the Atlas bear of North Africa and the Californian, Ungavan[11][12] 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.[1][13] 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.[14] 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 50 to 60 bears.[10][15]

Evolution and taxonomy

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).[16][17] In the mid-19th century United States, the brown bear was termed "Old Ephraim" and sometimes as "Moccasin Joe".[18]

The scientific name of the brown bear, Ursus arctos, comes from the Latin ursus, meaning "bear",[19] and from ἄρκτος arktos, the Greek word for bear.[20]

Generalized names and evolution

Brown bears are thought to have evolved from Ursus etruscus in Asia.[21][22] 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."[23] 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.[24] The oldest fossils positively identified as from this species occur in China from about 0.5 million years ago. Brown bears entered Europe about 250,000 years ago and North Africa shortly after.[21][25] Brown bear remains from the Pleistocene period are common in the British Isles, where it is thought they might have outcompeted cave bears (Ursus spelaeus). The species entered Alaska 100,000 years ago, though they did not move south until 13,000 years ago.[21] It is speculated that brown bears were unable to migrate south until the extinction of the much larger giant short-faced bear (Arctodus simus).[26][27]

Several paleontologists suggest the possibility of two separate brown bear migrations: inland brown bears, also known as grizzlies, are thought to stem from narrow-skulled bears which migrated from northern Siberia to central Alaska and the rest of the continent, while Kodiak bears descend from broad-skulled bears from Kamchatka, which colonized the Alaskan peninsula. Brown bear fossils discovered in Ontario, Ohio, Kentucky and Labrador show that the species occurred farther east than indicated in historic records.[21] In North America, two types of the subspecies Ursus arctos horribilis are generally recognized—the coastal brown bear and the inland grizzly bear; these two types broadly define the range of sizes of all brown bear subspecies.[13]

Scientific taxonomy

There are many methods used by scientists to define bear species and subspecies, as no one method is always effective. Brown bear taxonomy and subspecies classification has been described as "formidable and confusing," with few authorities listing the same specific set of subspecies.[28] Genetic testing is now perhaps the most important way to scientifically define brown bear relationships and names. Generally, genetic testing uses the word clade rather than species because a genetic test alone cannot define a biological species. Most genetic studies report on how closely related the bears are (or their genetic distance). There are hundreds of obsolete brown bear subspecies, each with its own name, and this can become confusing; Hall (1981) lists 86 different types, and even as many as 90 have been proposed.[29][30] However, recent DNA analysis has identified as few as five main clades which contain all extant brown bears,[31][32] while a 2017 phylogenetic study revealed nine clades, including one representing polar bears.[33] As of 2005, 15 extant or recently extinct subspecies were recognized by the general scientific community.[34][35]

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.[36] Under some definitions, the brown bear can be construed as the paraspecies for the polar bear.[37][38][39][40]

DNA analysis shows that, apart from recent human-caused population fragmentation,[41] 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.[42][43] 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,[44] 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.[45]

Brown bears are apparently divided into five different clades, some of which coexist or co-occur in different regions.[3]

Hybrids

 
Possible grizzly-black bear hybrid in the Yukon Territory, Canada

A grizzly–polar bear hybrid (known either as a pizzly bear or a grolar bear) 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.[46] 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).

Analyses of the genomes of bears have shown that introgression between species was widespread during the evolution of the genus Ursus,[47][48][49] including the introgression of polar bear DNA introduced to brown bears during the Pleistocene.

A bear shot in autumn 1986 in Michigan, US, was thought by some to be a grizzly/American black bear hybrid, due to its unusually large size and its proportionately larger braincase and skull. DNA testing was unable to determine whether it was a large American black bear or a grizzly bear.[50]

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 also 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.[51][52]

 
Skeleton

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.[53][54] 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.[6] 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. 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).[13][55][56][57]

Color

 
in Slovenia

Brown bears are often not fully brown.[58] They have long, thick fur, with a moderately long mane at the back of the neck which varies somewhat across the types.[59] 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.[58][60] 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.[61] The winter fur is very thick and long, especially in northern subspecies, and can reach 11 to 12 centimetres (4 to 5 in) at the withers. The winter hairs are thin, yet rough to the touch. The summer fur is much shorter and sparser and its length and density varies geographically.[62]

Cranial morphology and size

 
Skull

Adults have massive, heavily built concave skulls, which are large in proportion to the body. The forehead is high and rises steeply.[61] The projections of the skull are well developed when compared to those of Asian black bears (Ursus thibetanus): the latter have sagittal crests not exceeding more than 19–20% of the total length of the skull, while the former have sagittal crests comprising up to 40–41% of the skull's length. Skull projections are more weakly developed in females than in males. The braincase is relatively small and elongated. There is a great deal of geographical variation in the skull, and presents itself chiefly in dimensions.[62] Grizzlies, for example, tend to have flatter profiles than European and coastal American brown bears.[63] 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.[62] 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.[62] The teeth of brown bears reflect their dietary plasticity and are broadly similar to other bears, excluding the two most herbivorous living bears, the giant panda (Ailuropoda melanoleuca) and the spectacled bear (Tremarctos ornatus), which have blunt, small premolars (ideal for grinding down fibrous plants) compared to the jagged premolars of ursid bears that at least seasonally often rely on flesh as a food source.[64][65] The teeth are reliably larger than American black bears, but average smaller in molar length than polar bears.[66][67] Brown bears have the broadest skull of any extant ursine bear; only the aforementioned most herbivorous living bears exceed them in relative breadth of the skull.[13][55][68] Another extant ursine bear, the sloth bear (Melursus ursinus), has a proportionately longer skull than the brown bear and can match the skull length of even large brown bear subtypes, presumably as an aid for foraging heavily on insect colonies for which a long muzzle is helpful as an evolved feature in several unrelated mammalian groups.[61][68]

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.[69] They are generally dark with a light tip, with some forms having completely light claws.[62] Brown bear claws are longer and straighter than those of American black bears (Ursus americanus).[61] 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 both species of black bear, although in rare cases adult female brown bears have been seen in trees.[70] The claws of a polar bear are also quite different, being notably shorter but broader with a strong curve and sharper point, presumably both as an aid to traveling over ice (sometimes nearly vertically) and procuring active prey.[25][71] The paws of the brown bear are quite large. The rear feet of adult bears have been found to typically measure 21 to 36 cm (8.3 to 14.2 in) long, while the forefeet tend to measure about 40% less in length. All four feet in average sized brown bears tend to be about 17.5 to 20 cm (6.9 to 7.9 in) in width. In large coastal or Kodiak bear males, the hindfoot may measure up to 40 cm (16 in) in length, 28.5 cm (11.2 in) in width, while outsized Kodiak bears having had confirmed measurements of up to 46 cm (18 in) along their rear foot.[72][73][74] 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 most bears of the species and also used heavily in den construction prior to hibernation.[61][75] The brown bear's strength has been roughly estimated as 2.5 to 5 times that of a human.[76]

Distribution and habitat

 
Brown bear at Brooks Falls

Brown bears were once native to Europe, much of Asia, the Atlas Mountains of Africa, and North America,[77] 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.[78] The largest populations are in Russia with 120,000,[79] 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 32,000 individuals.[80] In the lower 48 states, they are repopulating slowly, but steadily along the Rockies and the western Great Plains.

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,[81][82] 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[83]) 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)[84][85] in the south.

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. They can also 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.[72]

This species inhabits the broadest range of habitats of any living bear species.[72] They seem to have no altitudinal preferences and have been recorded from sea level to an elevation of 5,000 m (16,000 ft) (the latter in the Himalayas).[72] 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.[72]

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][3] 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.[86] The Syrian brown bear (Ursus arctos syriacus) is very rare and it has been extirpated from more than half of its historic range.[87] One of the smallest-bodied subspecies, the Himalayan brown bear (Ursus arctos isabellinus), is critically endangered, occupying only 2% of its former range and threatened by uncontrolled poaching for its body parts.[14] The Marsican brown bear in central Italy is believed to have a population of just 50–60 bears.

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.[88] Studies have shown that activity throughout the range 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.[89][90][91] 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.[92][93] 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.[94][95]

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 (e.g., using a barnacle-covered rock to scratch its neck), which requires advanced cognitive abilities.[96] 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.[97][98] 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.[99][100] 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.[101] During combat, bears use their paws to strike their opponents in the chest or shoulders and bite the head or neck.[102]

Communication

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 makes "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.[102]

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!.[102]

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.[13][100] 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.[103] 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.[13][75][55][104] In areas where food is abundant and concentrated, such as coastal Alaska, home ranges for females are up to 24 km2 (9.3 sq mi) and for males are up to 89 km2 (34 sq mi). Similarly, in British Columbia, bears of the two sexes travel relatively compact home ranges of 115 km2 (44 sq mi) and 318 km2 (123 sq mi). In Yellowstone National Park, home ranges for females are up to 281 km2 (108 sq mi) and up to 874 km2 (337 sq mi) for males. In Romania, the largest home range was recorded for adult males (3,143 km2, 1214 sq mi).[105] 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.[13][72][75][100]

A study of male-inherited Y chromosome DNA sequence found that brown bears, over the past few 10,000 years, have shown strong male-biased dispersal.[106] That study found surprisingly similar Y chromosomes in brown bear populations as far apart as Norway and coastal Alaska, indicating extensive gene flow across Eurasia and North America. Notably, this contrasts with genetic signals from female-inherited mitochondrial DNA (mtDNA), where brown bears of different geographic regions typically show strong differences in their mtDNA, a result of female philopatry.

Reproduction

 
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.[72][55][107] Being serially monogamous, brown bears remain with the same mate from a couple of days to a couple of weeks.[13][108] Outside of this narrow time frame, adult male and female brown bears show no sexual interest in each other.[13] 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, 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.[109][103][110] 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.[54][110] 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. 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.[75][55][111][112][113][114][104][115] Paternity DNA tests have shown that littermates do not share the same father in up to 29% of litters.[104] 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.[55][104] Copulation is vigorous and prolonged and can last up to an hour, although the mean time is about 23–24 minutes.[55][116]

 
Grizzly bear cubs often imitate their mothers closely.

Males take no part in raising their cubs – parenting is left entirely to the females.[100][117] 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.[109][118][119] 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.[109][120][121] 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).[72][122][123] Older and larger females within a population tend to give birth to larger litters.[124] The size of a litter also depends on factors such as geographic location and food supply.[125] 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.[126] 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.[127][128]

 
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.[75] 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.[129] The cubs learn by following and imitating their mother's actions during the period they are with her.[130] 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.[55] 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.[131][132] Brown bears practice infanticide, as an adult male bear may kill the cubs of another.[104] 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.[104] Cubs flee up a tree, if available, when they see a strange male bear and the mother often successfully defends them, even though the male may be twice as heavy as she, although females have been known to die in these confrontations.[104][133][134]

Dietary habits

 
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.[72] Throughout life, this species is regularly curious about the potential of eating virtually any organism or object that they encounter. Food that is both abundant and easily accessed or caught is preferred. Their jaw structure has evolved to fit their dietary habits. Their diet varies enormously throughout their differing areas based on opportunity.

Despite their reputation, most brown bears are not highly carnivorous, as they derive up to 90% of their dietary food energy from vegetable matter.[135] They often feed on a variety of plant life, including berries, grasses, flowers, acorns and pine cones, as well as fungi such as mushrooms.[13] Among all bears, brown bears are uniquely equipped to dig for tough foods such as roots, bulbs[136] and shoots. They use their long, strong claws to dig out earth to reach the roots and their powerful jaws to bite through them.[13] In spring, winter-provided carrion, grasses, shoots, sedges, moss[136] and forbs are the dietary mainstays for brown bears internationally.[72] 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.[72] 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.[137] 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.[61] This species may eat birds and their eggs, including almost entirely ground- or rock-nesting species.[61] The diet may be supplemented by rodents or similar smallish mammals, including marmots, ground squirrels, mice, rats, lemmings and voles.[13] 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.[138]

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.[61]

 
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.[61] 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.[61] 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.[61] Large mammals preyed on can include various ungulate species such as elk, moose, caribou, muskoxen and wild boar.[13] 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.[139] It will also bite or swipe some prey to stun it enough to knock it over for consumption.[140] To pick out young or infirm individuals, bears will charge at herds so the slower-moving and more vulnerable individuals will be made apparent. Brown bears may also ambush young animals by finding them via scent.[13] 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.[140] 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.[13] 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.[61] They also 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.[13] Cannibalism is not unheard of, though predation is not normally believed to be the primary motivation when brown bears attack each other.[61]

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.[13] Plants and fruit farmed by humans are readily consumed as well, including corn, wheat, sorghum, melons and any form of berries.[61] They will also feed at domestic bee yards, readily consuming both honey and the brood (grubs and pupae) of the honey bee colony.[61] Human foods and trash or refuse is eaten when possible. When an open garbage dump was kept in Yellowstone, brown bears were one of the most voracious and regular scavengers. The dump was closed after both brown and American black bears came to associate humans with food and lost their natural fear of them.[61]

Interspecific predatory relationships

 
Brown bear being followed by a wolf

Adult bears are generally immune to predatory attacks except from tigers and other bears. Siberian tigers prefer preying on young Ussuri brown bears, but smaller adult female brown bears outside their dens may also be taken, generally when lethargic from hibernation.[141][142][143] Of 44 recorded encounters between the two predators, 20 resulted in confrontations; in 50% of these, the bears were killed, in 27% the tigers were killed, and 23% of the cases ended with both animals surviving and parting ways.[144] Some bears emerging from hibernation seek out tigers in order to steal their kills.[145] Some large brown bears may actually benefit from the tiger's presence by appropriating tiger kills that the bears may not be able to successfully hunt themselves and follow tiger tracks.[146] Geptner et al. (1972) stated bears are generally afraid of tigers and change their path after coming across tiger trails.[147] In the winters of 1970–1973, Yudakov and Nikolaev recorded 1 case of brown bear showing no fear of the tigers and another case of brown bear changing path upon crossing tiger tracks.[148] Other researchers have observed bears following tiger tracks for various reasons.[149]

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 case of fully-grown wolves being killed by a grizzly bear.[150] Given the opportunity, however, both species will prey on the other's cubs.[151] Conclusively, the individual power of the bear against the collective strength of the wolf pack usually results in a long battle for kills or domination.

In some areas, grizzly bears also regularly displace cougars from their kills.[152] 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.[153][154] 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.[61] There is one record of a golden eagle preying on a brown bear cub.[155]

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.[156] 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.[157] They will eat the fruit dropped by the Asian black bear from trees, as they themselves are too large and cumbersome to climb.[158] Improbably, in the Himalayas Brown bears are reportedly intimidated by Asian black bears in confrontations.[159]

There has been a recent 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,[160] and dead polar bear cubs have been found in brown bear dens.[161]

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.[72][162] The lifespan of brown bears of both sexes within minimally hunted populations is estimated at an average of 25 years.[163] The oldest wild brown bear on record was nearly 37 years old.[163] 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.[13][54]

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.[164] Annual mortality for bears of any age is estimated at 10% in most protected areas;[72] however, the average annual mortality rate rises to an estimated 38% in hunted populations.[72] Around 13% to 44% of cubs die within their first year even in well-protected areas.[13] Mortality rates of 75–100% among the cubs of any given year are not uncommon.[165] 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.[13][75][134][166] By the second and third years of their lives, the annual mortality rate among cubs in the care of their mothers drops to 10-15%.[75]

Even in populations living in protected areas, humans are still the leading cause of mortality for brown bears.[3][163][167] The largest amount of legalized brown bear hunting occurs in Canada, Finland, Russia, Slovakia and Alaska.[72] Hunting is unregulated in many areas within the range of the brown bear. Even where hunting is legally permitted, most biologists feel that the numbers hunted are excessive considering the low reproduction rate and sparse distribution of the species.[3][127][165] Brown bears are also killed in collisions with automobiles, which is a significant cause of mortality in the United States and Europe.[168][169]

Relationship with humans

Conflicts between bears and humans

 
Ancient depiction of a brown bear in the arena (Papyrus 3053)

Brown bears usually avoid areas where extensive development or urbanization has occurred, unlike the smaller, more inoffensive American black bear which can adapt to peri-urban regions.[170][171][172][173] Under many circumstances, extensive human development may cause brown bears to alter their home ranges.[174][175] High road densities (both paved and gravel roads) are often associated with higher mortality, habitat avoidance and lower bear density.[176] However, brown bears can easily lose their natural cautiousness upon being attracted to human-created food sources, such as garbage dumps, litter bins and dumpsters. Brown bears may even venture into human dwellings or barns in search of food as humans encroach on bear habitats.[177] In other areas, such as Alaska, dumps may continue to be an attractant for brown bears.[178][179] In different parts of their distribution, brown bears sometimes kill and eat domesticated animals.[180][181][182] The saying "A fed bear is a dead bear" has come into use to popularize the idea that allowing a bear to scavenge human garbage, such as trash cans and campers' backpacks, pet food, or other food sources that draw the bear into contact with humans, can result in the bear's death.[183] Results of a 2016 study performed in a southeastern British Columbian valley indicate that areas where attractive bear food and concentrated human settlements overlap, human-bear conflict can create an ecological trap resulting in a lower apparent survival rate for brown bears, as well as attracting additional bears and thereby causing overall population declines.[184]

When brown bears come to associate human activity with a "food reward", they are likely to continue to become emboldened; the likelihood of human-bear encounters increases, as they may return to the same location despite relocation. Relocation of the bear has been used to separate the bear from the human environment, but it does not address the problem of the bear's newly learned association of humans with food or the environmental situations which created the human-habituated bear. "Placing a bear in habitat used by other bears may lead to competition and social conflict, and result in the injury or death of the less dominant bear."[185] Yellowstone National Park, a reserve located in the western United States, contains prime habitat for the grizzly bear (Ursus arctos horribilis) and due to the enormous number of visitors, human-bear encounters are common. The scenic beauty of the area has led to an influx of people moving into the area. In addition, because there are so many bear relocations to the same remote areas of Yellowstone, and because male bears tend to dominate the center of the relocation zone, female bears tend to be pushed to the boundaries of the region and beyond. As a result, a large proportion of repeat offenders, bears that are killed for public safety, are females. This creates a further depressive effect on an already-endangered subspecies. The grizzly bear is officially described as "Threatened" in the U.S.. Although the problem is most significant with regard to grizzlies, these issues affect the other types of brown bears as well.[186][187][188]

In Europe, part of the problem lies with shepherds; over the past two centuries, many sheep and goat herders have gradually abandoned the more traditional practice of using dogs to guard flocks, which have concurrently grown larger. Typically, they allow the herds to graze freely over sizeable tracts of land. As brown bears reclaim parts of their range, they may eat livestock as sheep and goats are relatively easy for a bear to kill. In some cases, the shepherds shoot the bear, thinking their livelihood is under threat. Many are now better informed about the ample compensation available and will make a claim when they lose livestock to a bear.[189][190][191] Another issue in several parts of their range in Europe is supplemental feeding stations where various kind of animal carrion is offered, which are set up mainly in Scandinavia and eastern Europe both to support the locally threatened species and so humans can enjoy watching bears that may otherwise prove evasive. Despite that most stations were cautiously set in remote areas far from human habitations, some brown bears in such areas have become conditioned to associate humans with food and become excessively bold "problem bears". Also, supplemental feeding appears to cause no decrease in livestock predation.[192][193]

Bear encounters and attacks

 
A statue of the Ussuri brown bear from Hokkaido which perpetrated the worst brown bear attack in Japanese history, killing seven people

Brown bears seldom attack humans on sight and usually avoid people. In Russia, it is estimated that 1 in 1,000 on-foot encounters with brown bears results in an attack.[194] They are, however, unpredictable in temperament, and may attack if they are surprised or feel threatened.[195] There are an average of two fatal attacks by bears per year in North America.[196] In Scandinavia, there are only four known cases since 1902 of bear encounters which have resulted in death. The two most common causes for bear attack are surprise and curiosity.[197] Some types of bears, such as polar bears, are more likely to attack humans when searching for food, while American black bears are much less likely to attack. Despite their boldness and potential for predation if the bear is hungry, polar bears rarely attack humans, because they are infrequently encountered in the Arctic sea.[196] Aggressive behavior in brown bears is favored by numerous selection variables. Increased aggressiveness also assists female brown bears in better ensuring the survival of their young to reproductive age.[198] Mothers defending cubs are the most prone to attacking, being responsible for 70% of brown bear-caused human fatalities in North America.[199]

Sows with cubs account for many attacks on humans by brown bears in North America. Habituated or food-conditioned bears can also be dangerous, as their long-term exposure to humans causes them to lose their natural shyness and, in some cases, to associate humans with food. Small parties of one or two people are more often attacked by brown bears than large groups, with only one known case of an attack on a group of six or more. In that instance, it is thought that due to surprise, the grizzly bear may not have recognized the size of the group.[200] In the majority of attacks resulting in injury, brown bears precede the attack with a growl or huffing sound.[195] In contrast to injuries caused by American black bears, which are usually minor, brown bear attacks more often tend to result in serious injury and, in some cases, death.[195] Brown bears seem to confront humans as they would when fighting other bears: they rise up on their hind legs, and attempt to "disarm" their victims by biting and holding on to the lower jaw to avoid being bitten in turn.[26] Due to the bears' enormous physical strength, even a single bite or swipe can be deadly as in tigers, with some human victims having had their heads completely crushed by a bear bite.[63][201][failed verification] Most attacks occur in the months of July, August and September, the time when the number of outdoor recreationalists, such as hikers or hunters, is higher. People who assert their presence through noises tend to be less vulnerable, as they alert bears to their presence. In direct confrontations, people who run are statistically more likely to be attacked than those who stand their ground. Violent encounters with brown bears usually last only a few minutes, though they can be prolonged if the victims fight back.[195] In Alberta, two common behaviors by human hunters, imitating the calls of deer to attract them and carrying ungulate carcasses, seem to court aggressive behavior and lead to a higher rate of attack from grizzly bears.[202]

Attacks on humans are considered extremely rare in the former Soviet Union, though exceptions exist in districts where they are not as often pursued by hunters.[62] East Siberian brown bears, for example, tend to be much bolder toward humans than their shyer, more persecuted European counterparts. The delineation in Eurasia between areas where aggressiveness of brown bears tends to increase is the Ural Mountains, although the brown bears of eastern Europe are somewhat more aggressive than those of western Europe.[163][203] In 2008, a platinum mining compound in the Olyotorsky district of northern Kamchatka was besieged by a group of 30 bears, who killed two guards and prevented workers from leaving their homes.[204] 10 people a year on average are killed by brown bears in Russia, more than all the other parts of the brown bear's international range combined, although Russia also holds more brown bears than all other parts of the world combined.[205] In Scandinavia, only three fatal attacks were recorded in the 20th century.[206]

In Japan, a large brown bear nicknamed "Kesagake" (袈裟懸け, "kesa-style slasher") made history for causing 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 (with possibly another three previous fatalities to its credit) before being gunned down after a large-scale beast-hunt. Today, there is still a shrine at Rokusensawa (六線沢), where the event took place in memory of the victims of the incident.[207]

Within Yellowstone National Park, injuries caused by grizzly attacks in developed areas averaged approximately one per year during the 1930s through to the 1950s, though it increased to four per year during the 1960s. They then decreased to one injury every two years during the 1970s. Between 1980 and 2002, there have been only two human injuries caused by grizzly bears in a developed area. Although grizzly attacks were rare in the backcountry before 1970, the number of attacks increased to an average of approximately one per year during the 1970s, 1980s and 1990s.[208] In Alberta, from 1960 to 1998, the number of attacks by grizzly bears ending in injury were nearly three times more common than attacks ending in injury by American black bears, despite the American black bear being an estimated 38 times more numerous in the province than the grizzly bear.[209]

History of defense from bears

 
can of bear spray

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.[210] Carrying pepper spray is highly recommended by many authorities when traveling in bear country; however, carrying two means of deterrent, one of which is a large caliber gun, is also advised. Solid shotgun slugs, or three buckshot rounds, or a pistol of .44 caliber or more is suggested if a heavy hunting rifle is not available. Guns remain a viable, last resort option to be used in defense of life from aggressive brown bears.[211] Too often, people do not carry a proper caliber weapon to neutralize the bear. According to the Alaska Science Center, a 12-gauge shotgun with slugs has been the most effective weapon. There have been fewer injuries as a result of only carrying lethal loads in the shotgun, as opposed to deterrent rounds. State of Alaska Defense of Life or Property (DLP) laws require one to report the kill to the authorities and salvage the hide, skull and claws.[212] A page at the State of Alaska Department of Natural Resources website offers information about how to "select a gun that will stop a bear (12-gauge shotgun or .300 mag rifle)".[213]

Campers are often told to wear bright-colored red ribbons and bells and carry whistles to ward off bears. They are told to look for grizzly bear dung in camping areas and be careful to carry the bells and whistles in those areas. Grizzly bear dung is difficult to differentiate from American black bear dung, as diet is in a constant state of flux depending on the availability of seasonal food items. If a bear is killed near camp, the bear's carcass must be adequately disposed of, including entrails and blood, if possible. Failure to move the carcass has often resulted in it attracting other bears and further exacerbating a bad situation. Moving camps immediately is another recommended method.[195]

Culture

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

Brown bears often figure into the literature of Europe and North America, in particular that which is written for children. "The Brown Bear of Norway" is a Scottish fairy tale telling the adventures of a girl who married a prince magically turned into a bear and who managed to get him back into a human form by the force of her love and after many trials and difficulties. With "Goldilocks and the Three Bears", a story from England, the Three Bears are usually depicted as brown bears. In German-speaking countries, children are often told the fairytale of "Snow White and Rose Red"; the handsome prince in this tale has been transfigured into a brown bear. In the United States, parents often read their preschool age children the book Brown Bear, Brown Bear, What Do You See? to teach them their colors and how they are associated with different animals.[214]

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.[215]

The grizzly bear is the state animal of Montana.[216] The California golden bear is the state animal of California.[217] Both animals are subspecies of the brown bear and the species was extirpated from the latter state.

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. The brown bear is depicted on the reverse of the Croatian 5 kuna coin, minted since 1993.

The Bundesliga club Bayern Munich has a brown bear mascot named Berni. The Chicago National Football League (NFL) franchise is named the Bears. In this context, no differentiation between American black and brown bears is needed. The school mascot for Bob Jones University, Brown University, Cornell University, George Fox University, the University of Alberta, the University of California, Berkeley, the University of California, Los Angeles, the University of California, Riverside, and numerous American high schools is the brown bear.

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

See also

References

  1. ^ a b c d e f 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. Retrieved 14 January 2022.
  3. ^ a b c d e f 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
  4. ^ Boddington, Craig (2004). Fair Chase in North America. Illustrations by Carlson, Ken. Boone and Crockett Club. p. 45. ISBN 978-0-940864-47-4.
  5. ^ Martin, A. P.; Palumbi, S. R. (1993). "Body size, metabolic rate, generation time, and the molecular clock". Proceedings of the National Academy of Sciences of the United States of America. 90 (9): 4087–4091. Bibcode:1993PNAS...90.4087M. doi:10.1073/pnas.90.9.4087. PMC 46451. PMID 8483925.
  6. ^ a b Christiansen, P. (1999). "What size were Arctodus simus and Ursus spelaeus (Carnivora: Ursidae)?". Annales Zoologici Fennici. 36 (2): 93–102. JSTOR 23735739.
  7. ^ a b Servheen, C.; Darling, L. M.; Archibald, W. R. (1990), The status and conservation of the bears of the world, International Association for Bear Research and Management
  8. ^ (PDF). American Society of Mammalogists, Smith College. 23 April 1993. Archived from the original (PDF) on 31 March 2017.
  9. ^ White, Paul. . Transylvania Wildlife Project. Archived from the original on 19 May 2013.
  10. ^ a b Zedrosser, A.; Dahle, B.; Swenson, J. E.; Gerstl, N. (2001). "Status and management of the brown bear in Europe" (PDF). Ursus. 12: 9–20. Archived (PDF) from the original on 9 October 2022.
  11. ^ Busch, Robert (2004). The Grizzly Almanac. Globe Pequot Press. pp. 11–14. ISBN 978-1-5922-8320-0. Retrieved 21 October 2014.
  12. ^ Elson, C. S. (1954). "Further Evidence about the Barren-Ground Grizzly Bear in Northeast Labrador and Quebec". Journal of Mammalogy. 35 (3): 345–357. doi:10.2307/1375959. JSTOR 1375959.
  13. ^ a b c d e f g h i j k l m n o p q r s t (PDF). American Society of Mammalogists, Smith College. 23 April 1993. Archived from the original (PDF) on 31 March 2017.
  14. ^ a b Himalayan brown bears now critically endangered 16 October 2018 at the Wayback Machine. Euronews.com. 6 January 2014
  15. ^ Boscagli, G. (1990). "Marsican brown bear population in central Italy—Status report 1985". Aquilo, Series Zoologica. 27: 81–83.
  16. ^ Harper, Douglas. "bruin". Online Etymology Dictionary.
  17. ^ Bruin, Encyclopaedia Britannica
  18. ^ Hunting the Grisly and other Sketches. FullTextArchive.com. Retrieved 15 September 2011.
  19. ^ Liddell, Henry George and Scott, Robert. "Ursus." A Latin Dictionary. Perseus Digital Library.
  20. ^ Liddell, Henry George and Scott, Robert."Arktos." A Greek-English Lexicon. Perseus Digital Library.
  21. ^ 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.
  22. ^ 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.
  23. ^ Kurten, Bjorn (1976). The Cave Bear Story. New York: Columbia University Press.
  24. ^ 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. doi:10.1016/s0960-9822(01)00046-x. PMID 11231157. S2CID 14645603.
  25. ^ 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.
  26. ^ a b Geist, Valerius (1989), (PDF), in Clutton-Brock, Juliet (ed.), The Walking larder: patterns of domestication, pastoralism, and predation, Unwin Hyman, pp. 282–294, ISBN 0-0444-5013-3, archived from the original (PDF) on 16 December 2008
  27. ^ Kurten, B.; Anderson, E. (1974). "Association of Ursus arctos and Arctodus simus (Mammalia: Ursidae) in the late Pleistocene of Wyoming". Breviora. 426: 1–6.
  28. ^ Wilson, D. E.; Ruff, S. (1999), The Smithsonian Book of North American Mammals, Washington, D.C.: Smithsonian Institution Press
  29. ^ Harris, Arthur H. (2013). "Pleistocene Vertebrates of Arizona, New Mexico, and Trans-Pecos Texas". UTEP Biodiversity Collections, University of Texas at El Paso.
  30. ^ 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. Alt URL
  31. ^ 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.
  32. ^ 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. doi:10.1111/j.1365-294x.2008.03631.x. PMID 18363668. S2CID 23361337. Archived (PDF) from the original on 9 October 2022.
  33. ^ 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.
  34. ^ 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.
  35. ^ Ursus arctos, ITIS
  36. ^ 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.
  37. ^ 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.
  38. ^ Shields, G. F.; Kocher, T. D. (1991). "Phylogenetic relationships of North American ursids based on analysis of mitochondrial DNA". Evolution. 45 (1): 218–221. doi:10.2307/2409495. JSTOR 2409495. PMID 28564083.
  39. ^ Kurtén, B. (1964). "The evolution of the polar bear, Ursus maritimus (Phipps)". Acta Zoologica Fennica. 108: 1–26.
  40. ^ Ingólfsson, Ólafur; Wiig, Øystein (2009). "Late Pleistocene fossil find in Svalbard: the oldest remains of a polar bear (Ursus maritimus Phipps, 1744) ever discovered". Polar Research. 28 (3): 455. Bibcode:2009PolRe..28..455I. doi:10.1111/j.1751-8369.2008.00087.x. S2CID 56292690.
  41. ^ 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. doi:10.1002/wmon.6. ISSN 1938-5455.
  42. ^ 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.
  43. ^ Waits, L. P.; et al. (1998). "Mitochondrial DNA Phylogeography of the North American Brown Bear and Implications for Conservation". Conservation. 12 (2): 408–417. doi:10.1111/j.1523-1739.1998.96351.x. S2CID 86172292.
  44. ^ 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. doi:10.1046/j.1365-294x.1998.00440.x. ISSN 0962-1083. PMID 9787441. S2CID 21848010.
  45. ^ 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.
  46. ^ 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.
  47. ^ 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.
  48. ^ 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.
  49. ^ 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.
  50. ^ Smith, Richard P. (2007). "Hybrid Black Bear". Black Bear Hunting. Stackpole Books. ISBN 978-0-8117-0269-0.
  51. ^ 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.
  52. ^ 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.
  53. ^ Parker, S. P. (1990). Grzimek's encyclopedia of mammals. McGraw-Hill, New York. ISBN 0-07-909508-9 (set).
  54. ^ a b c Nowak, R. M. (1999). Walker's mammals of the world (Vol. 1). JHU Press.
  55. ^ a b c d e f g h Feldhamer, G. A.; Thompson, B. C.; Chapman, J. A. (2003). Wild mammals of North America: biology, management, and conservation. JHU Press. ISBN 9780801874161.
  56. ^ 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.
  57. ^ Steyaert, Sam M. J. G.; Endrestøl, Anders; Hackländer, Klaus; Swenson, Jon E; Zedrosser, Andreas (2012). "The mating system of the brown bear Ursus arctos". Mammal Review. 42: 12. doi:10.1111/j.1365-2907.2011.00184.x.
  58. ^ a b Jones, S. V. (1923). "Color variations in wild animals". Journal of Mammalogy. 4 (#3): 172–177. doi:10.2307/1373567. JSTOR 1373567.
  59. ^ Swenson, J. E. (2000). Action plan for the conservation of the brown bear in Europe (Ursus arctos) (No. 18-114). Council of Europe.
  60. ^ 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.
  61. ^ a b c d e f g h i j k l m n o p q Bear Anatomy and Physiology from Gary Brown's The Great Bear Almanac, Lyons & Burford, Publishers, 1993
  62. ^ a b c d e f 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., USA: Science Publishers, Inc. ISBN 1-886106-81-9.
  63. ^ a b The living animals of the world; a popular natural history with one thousand illustrations Volume 1: Mammals, by Cornish, C. J., 1858–1906; Selous, Frederick Courteney, 1851–1917; Johnston, Harry Hamilton, Sir, 1858–1927; Maxwell, Herbert, Sir, published by New York, Dodd, Mead and Company. Archive.org. Retrieved 15 September 2011.
  64. ^ 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.
  65. ^ Kurtén, B. (1966). "Pleistocene bears of North America: Genus Tremarctos, spectacled bears". Acta Zoologica Fennica. 115: 1–96.
  66. ^ Slaughter, B. H.; Pine, R. H.; Pine, N. E. (1974). "Eruption of cheek teeth in insectivora and carnivora". Journal of Mammalogy. 55 (#1): 115–25. doi:10.2307/1379261. JSTOR 1379261. PMID 4819587.
  67. ^ 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.
  68. ^ a b Christiansen, P. (2007). "Evolutionary implications of bite mechanics and feeding ecology in bears". Journal of Zoology. 272 (#4): 423. doi:10.1111/j.1469-7998.2006.00286.x.
  69. ^ Heptner, "Sirenia and carnivora (Sea cows; Wolves and Bears)"
  70. ^ Seryodkin, I. V. (2016). Behavior of Brown Bears During Feeding in the Sikhote-Alin. Achievements in the Life Sciences.
  71. ^ Stirling, I., & Guravich, D. (1998). Polar bears. University of Michigan Press.
  72. ^ a b c d e f g h i j k l m n o Carnivores of the World by Dr. Luke Hunter. Princeton University Press (2011), ISBN 9780691152288
  73. ^ Wood, Gerald (1983). The Guinness Book of Animal Facts and Feats. ISBN 978-0-85112-235-9.
  74. ^ Whitaker, J. O., & Elman, R. (1996). National Audubon Society Field Guide to North American Mammals (p. 992). New York: Knopf.
  75. ^ a b c d e f g Novak, M., Baker, J.A., Obbard, M.E. & Malloch, B. (1987). Wild furbearer management and conservation in North America. Ontario Ministry of Natural Resources.
  76. ^ 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.
  77. ^ "Ancient bear made early migration". BBC News. 12 November 2004. Retrieved 2 January 2010.
  78. ^ "Brown Bear | Species | WWF". WWF.
  79. ^ . Rolling Hills Wildlife Adventure. Archived from the original on 10 November 2016. Retrieved 2 December 2013.
  80. ^ "Brown Bear Research in Alaska". Alaska Department of Fish and Game. Retrieved 4 April 2012.
  81. ^ "Neixen quatre cries d'ós bru als Pirineus". elperiodico.cat (in Catalan and Spanish). 2 August 2010. Retrieved 19 November 2010.
  82. ^ "Situació preocupant de l'ós bru als Pirineus" (in Catalan). ecologistasenaccion.org. May 2010. Retrieved 19 November 2010.
  83. ^ "Osos, el desafío de una población creciente". ABC (in Spanish). Spain. 28 September 2013.
  84. ^ Martínez‐Abraín, A.; Jiménez, J.; Oro, D. (2018). "Pax Romana: 'refuge abandonment' and spread of fearless behavior in a reconciling world". Animal Conservation. John Wiley & Sons, Inc (Zoological Society of London (ZSL)). 22 (1): 3–13. doi:10.1111/acv.12429. ISSN 1367-9430. S2CID 89938098.
  85. ^ "Bear". ΑΡΚΤΟΥΡΟΣ. Retrieved 13 October 2022.
  86. ^ Miller, CR; Waits, L.P. (2006). "Phylogeography and mitochondrial diversity of extirpated brown bear (Ursus arctos) populations in the contiguous United States and Mexico". Mol. Ecol. 15 (#14): 4477–85. doi:10.1111/j.1365-294x.2006.03097.x. PMID 17107477. S2CID 7336900.
  87. ^ Calvignac, Sebastien; Hughes, Sandrine; Hanni, Catherine (2009). "Genetic diversity of endangered brown bear (Ursus arctos) populations at the crossroads of Europe, Asia and Africa". Diversity and Distributions. 15 (#5): 742–750. doi:10.1111/j.1472-4642.2009.00586.x. S2CID 21666120.
  88. ^ . North American Bear Center. Archived from the original on 24 May 2013. Retrieved 16 August 2013.
  89. ^ 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.
  90. ^ 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.
  91. ^ 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.
  92. ^ 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.
  93. ^ 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.
  94. ^ 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.
  95. ^ 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.
  96. ^ 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.
  97. ^ Egbert, Allan L.; Stokes, Allen W.; Egbert, A. L. (1974). "The social behaviour of brown bears on an Alaskan salmon stream" (PDF). International Conference Bear Res. And Manage. 3: 41–56. doi:10.2307/3872753. JSTOR 3872753.
  98. ^ 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.
  99. ^ 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.
  100. ^ 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. doi:10.1046/j.1365-2656.2003.00737.x. JSTOR 3505643. PMID 30893970. S2CID 67818528.
  101. ^ Stonorov, Derek and Stokes, Allen W. (1972) "Social Behavior of the Alaska Brown Bear" Panel 4: Bear Behaviour
  102. ^ a b c Egbert, A. L. (1978). The Social Behavior of Brown Bears at McNeil River, Alaska (Ph.D. thesis). Utah State University.
  103. ^ a b Pearson, A. M. (1975). The northern interior grizzly bear Ursus arctos L. Information Canada.
  104. ^ a b c d e f g 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. doi:10.1111/j.1439-0310.2006.01152.x. Archived (PDF) from the original on 9 October 2022.
  105. ^ 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.
  106. ^ Bidon, T.; Janke, A.; Fain, S. R.; Eiken, H. G.; Hagen, S. B.; Saarma, U.; Hallstrom, B. M.; Lecomte, N.; Hailer, F. (2014). "Brown and Polar Bear Y Chromosomes Reveal Extensive Male-Biased Gene Flow within Brother Lineages". Molecular Biology and Evolution. 31 (#6): 1353–1363. doi:10.1093/molbev/msu109. PMID 24667925.
  107. ^ 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.
  108. ^ Types of Bears – Information on Specific Bear Species (2009)
  109. ^ a b c 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.
  110. ^ 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.
  111. ^ Nawaz, Muhammad Ali; Swenson, Jon E.; Zakaria, Vaqar (2008). "Pragmatic management increases a flagship species, the Himalayan brown bears, in Pakistan's Deosai National Park". Biological Conservation. 141 (#9): 2230. doi:10.1016/j.biocon.2008.06.012.
  112. ^ Paralikidis, N. P.; Papageorgiou, N. K.; Kontsiotis, V. J.; Tsiompanoudis, A. C. (2010). "The dietary habits of the brown bear (Ursus arctos) in western Greece". Mammalian Biology – Zeitschrift für Säugetierkunde. 75: 29. doi:10.1016/j.mambio.2009.03.010.
  113. ^ 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.
  114. ^ Naves, J.; Fernández-Gil, A.; Rodríguez, C.; Delibes, M. (2006). "Brown Bear Food Habits at the Border of Its Range: A Long-Term Study". Journal of Mammalogy. 87 (#5): 899. doi:10.1644/05-MAMM-A-318R2.1. hdl:10261/50290. S2CID 85351746.
  115. ^ 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.
  116. ^ 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.
  117. ^ Corbet, G. B. (1966). Terrestrial mammals of Western Europe. G.T. Foulic & Co. Ltd., London, UK.
  118. ^ 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.
  119. ^ 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.
  120. ^ Frković, A.; Huber, D. & Kusak, J. (2001). "Brown bear litter sizes in Croatia". Ursus. 12: 103–105. JSTOR 3873235.
  121. ^ 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.
  122. ^ 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.
  123. ^ Barnes Jr, V.; Smith, R. (1993). "Cub adoption by brown bears (Ursus arctos middendorffi) on Kodiak Island, Alaska". Canadian Field-Naturalist. 107: 365–367.
  124. ^ 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.
  125. ^ 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.
  126. ^ Couturier MA (1954). L'ours Brun, Ursus arctos. L. Couturier, Grenoble, France.
  127. ^ a b Mallinson, J. (1978). The Shadow of Extinction: Europe's Threatened Mammals. Macmillan, London.
  128. ^ 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.
  129. ^ 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.
  130. ^ . Shadowofthebear.com. Archived from the original on 1 May 2008.
  131. ^ 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.
  132. ^ 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.
  133. ^ 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.
  134. ^ a b 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.
  135. ^ . Alaska Office of Economic Development. Dced.state.ak.us. Archived from the original on 15 April 2009. Retrieved 10 October 2009.
  136. ^ a b "Ursus arctos (Brown bear)".
  137. ^ . Yellowstonepark.com. 21 June 2011. Archived from the original on 15 July 2010.
  138. ^ Brown, Susan, A (17 January 2010). "Inherited behavior traits of the domesticated ferret". weaselwords.com. Retrieved 29 January 2010.
  139. ^ 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.
  140. ^ a b Macdonald, D.W.; Barrett, P. (1993). Mammals of Europe. New Jersey: Princeton University Press. p. 107. ISBN 978-0-691-09160-0.
  141. ^ David Prynn (2004). Amur tiger. Russian Nature Press. p. 115.
  142. ^ Frasef, A. (2012). Feline Behaviour and Welfare. CABI. pp. 72–77. ISBN 978-1-84593-926-7.
  143. ^ Seryodkin; et al. (2003). "Denning ecology of brown bears and Asiatic black bears in the Russian Far East". Ursus. 14 (2): 159.
  144. ^ 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. Archived (PDF) from the original on 9 October 2022.
  145. ^ Matthiessen, Peter; Hornocker, Maurice (2001). Tigers In The Snow. North Point Press. ISBN 0-86547-596-2.
  146. ^ Miquelle, D.G.; Smirnov, E.N.; Goodrich, J.M. (2005). "1". Tigers of Sikhote-Alin Zapovednik: ecology and conservation. Vladivostok, Russia: PSP.
  147. ^ Geptner, V. G., Sludskii, A. A. (1972). Mlekopitaiuščie Sovetskogo Soiuza. Vysšaia Škola, Moskva. (In Russian; English translation: Heptner, V. G.; Sludskii, A. A.; Bannikov, A. G.; (1992). Mammals of the Soviet Union. Volume II, Part 2: Carnivora (Hyaenas and Cats). Smithsonian Institution and the National Science Foundation, Washington DC). pp. 95–202.
  148. ^ 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 (english translation ed.). 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.
  149. ^ 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.
  150. ^ 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.
  151. ^ Downey, Betsy. (PDF). International Wolf Center. Archived from the original (PDF) on 27 February 2008.
  152. ^ 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.
  153. ^ ADW: Ursus arctos: Information 12 May 2013 at the Wayback Machine. Arlis.org. Retrieved 9 August 2012.
  154. ^ Hornocker, M., and S. Negri (eds.) (2009). Cougar: ecology and conservation. University of Chicago Press. Chicago, IL, ISBN 0226353443.
  155. ^ 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.
  156. ^ 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.
  157. ^ 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.
  158. ^ The Intellectual observer: review of natural history, microscopic research, and recreative science, Groombridge, 1865
  159. ^ Adams, Andrew Leith (1867) Wanderings of a naturalist in India: the western Himalayas, and Cashmere, Edmonston and Douglas
  160. ^ Dough O'Hara . Anchorage Daily News. 24 April 2005
  161. ^ "ABC News: Grizzlies Encroaching on Polar Bear Country". ABC News. Retrieved 10 October 2009.
  162. ^ 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.
  163. ^ a b c d McDonald, D., & Norris, S. (2001). The new encyclopedia of mammals. Oxford University Press.
  164. ^ 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.
  165. ^ a b Servheen, C. (1987). Grizzly bear compendium. Interagency Grizzly Bear Committee.
  166. ^ Palomares, F.; Caro, T. M. (1999). "Interspecific Killing among Mammalian Carnivores" (PDF). The American Naturalist. 153 (#5): 492–508. doi:10.1086/303189. hdl:10261/51387. PMID 29578790. S2CID 4343007. Archived (PDF) from the original on 9 October 2022.
  167. ^ "Brown/Grizzly Bear Hunting in Alaska". Alaska Department of Fish and Game.
  168. ^ Huber, D.; Kusak, J.; Frkovic, A. (1998). "Traffic kills of brown bears in Gorski kotar, Croatia". Ursus. 10: 167–171. JSTOR 3873124.
  169. ^ Servheen, C., Waller, J., & Kasworm, W. (1998). Fragmentation effects of high-speed highways on grizzly bear populations shared between the United States and Canada. International Conference on Wildlife Ecology and Transportation (ICOWET 1998).
  170. ^ Kaczensky, P.; Blazic, M.; Gossow, H. (2004). "Public attitudes towards brown bears (Ursus arctos) in Slovenia". Biological Conservation. 118 (5): 661. doi:10.1016/j.biocon.2003.10.015.
  171. ^ Woodroffe, R. (2000). "Predators and people: Using human densities to interpret declines of large carnivores" (PDF). Animal Conservation. 3 (2): 165. doi:10.1111/j.1469-1795.2000.tb00241.x. S2CID 84430290. Archived (PDF) from the original on 9 October 2022.
  172. ^ Martin, J.; Basille, M.; Van Moorter, B.; Kindberg, J.; Allaine, D.; Swenson, J. E. (2010). "Coping with human disturbance: spatial and temporal tactics of the brown bear (Ursus arctos)". Canadian Journal of Zoology. 88 (9): 875. doi:10.1139/Z10-053. S2CID 44121835.
  173. ^ Larivière, S. (2001). (PDF). Mammalian Species (647): 1–11. doi:10.1644/1545-1410(2001)647<0001:ua>2.0.co;2. S2CID 198968922. Archived from the original (PDF) on 16 March 2016. Retrieved 14 December 2016.
  174. ^ Mattson, D. J. (1990). "Human impacts on bear habitat use" (PDF). Bears: Their Biology and Management. 8: 33–56. doi:10.2307/3872901. JSTOR 3872901. Archived (PDF) from the original on 9 October 2022.
  175. ^ Elgmork, K. (1978). "Human impact on a brown bear population (Ursus arctos L.)". Biological Conservation. 13 (2): 81. doi:10.1016/0006-3207(78)90063-0.
  176. ^ Lamb, C.T.; Mowat, G.; Reid, A.; Smit, L.; Proctor, M.; McLellan, B.N.; Nielsen, S.E.; Boutin, S. (2018). "Effects of habitat quality and access management on the density of a recovering grizzly bear population". Journal of Applied Ecology. 55 (3): 1406–1417. doi:10.1111/1365-2664.13056.
  177. ^ Stringham, S. F. (1986). "Effects of climate, dump closure, and other factors on Yellowstone grizzly bear litter size". Bears: Their Biology and Management. 6: 33–39. doi:10.2307/3872803. JSTOR 3872803.
  178. ^ Peirce, K. N.; Van Daele, L. J. (2006). "Use of a garbage dump by brown bears in Dillingham, Alaska". Ursus. 17 (2): 165–177. doi:10.2192/1537-6176(2006)17[165:uoagdb]2.0.co;2. JSTOR 3873094. S2CID 54802857.
  179. ^ Robbins, C. T.; Schwartz, C. C.; Felicetti, L. A. (2004). "Nutritional ecology of ursids: A review of newer methods and management implications" (PDF). Ursus. 15 (2): 161–171. doi:10.2192/1537-6176(2004)015<0161:NEOUAR>2.0.CO;2. S2CID 54680694. Archived (PDF) from the original on 9 October 2022.
  180. ^ Clevenger, A. P.; Purroy, F. J.; Pelton, M. R. (1992). "Food habits of brown bears (Ursus arctos) in the Cantabrian Mountains, Spain". Journal of Mammalogy. 73 (2): 415–421. doi:10.2307/1382077. JSTOR 1382077.
  181. ^ Chauhan, N. P. S. (2003). "Human casualties and livestock depredation by black and brown bears in the Indian Himalaya, 1989–98" (PDF). Ursus: 84–87. Archived (PDF) from the original on 9 October 2022.
  182. ^ Sagør, J. T.; Swenson, J. E.; Røskaft, E. (1997). "Compatibility of brown bear Ursus arctos and free-ranging sheep in Norway". Biological Conservation. 81 (1–2): 91. doi:10.1016/S0006-3207(96)00165-6.
  183. ^ Herrero, S. (2002). Bear attacks: their causes and avoidance. Globe Pequot. ISBN 978-1585745579.
  184. ^ Lamb, C.T..; Mowat, G.; McLellan, B.N.; Nielsen, S.E.; Boutin, S. (2016). "Forbidden fruit: human settlement and abundant fruit create an ecological trap for an apex omnivore". Journal of Animal Ecology. 86 (1): 55–65. doi:10.1111/1365-2656.12589. PMID 27677529. S2CID 3347450.
  185. ^ "Relocation". Get Bear Smart Society. Archived from the original on 13 May 2006.
  186. ^ Gunther, K. A.; Haroldson, M. A.; Frey, K.; Cain, S. L.; Copeland, J.; Schwartz, C. C. (2004). "Grizzly bear-human conflicts in the Greater Yellowstone ecosystem, 1992–2000" (PDF). Ursus. 15 (#1): 10–22. doi:10.2192/1537-6176(2004)015<0010:gbcitg>2.0.co;2. S2CID 86821625. Archived (PDF) from the original on 9 October 2022.
  187. ^ Blanchard, B. M.; Knight, R. R. (1995). "Biological consequences of relocating grizzly bears in the Yellowstone ecosystem". The Journal of Wildlife Management. 59 (#3): 560–565. doi:10.2307/3802463. JSTOR 3802463.
  188. ^ Brannon, R. D. (1987). . Canadian Field-Naturalist. 101 (#4): 569–575. Archived from the original on 19 December 2019. Retrieved 12 December 2016.
  189. ^ Mustoni, A.; Carlini, E.; Chiarenzi, B.; et al. (2003). "Planning the Brown Bear Ursus arctos reintroduction in the Adamello Brenta Natural Park". Hystrix: The Italian Journal of Mammalogy. 14 (1–2). doi:10.4404/hystrix-14.1-2-4313.
  190. ^ Dahle, B.; Sørensen, O. J.; Wedul, E. H.; Swenson, J. E.; Sandegren, F. (1998). "The diet of brown bears Ursus arctos in central Scandinavia: effect of access to free-ranging domestic sheep Ovis aries". Wildlife Biology. 4 (3): 147–158. doi:10.2981/wlb.1998.017. S2CID 90983953.
  191. ^ Mertens, A.; Promberger, C. (2001). "Economic aspects of large carnivore-livestock conflicts in Romania". Ursus. 12: 173–180. JSTOR 3873246.
  192. ^ Kavčič, I.; Adamič, M.; Kaczensky, P.; Krofel, M.; Kobal, M.; Jerina, K. (2015). "Fast food bears: brown bear diet in a human-dominated landscape with intensive supplemental feeding". Wildlife Biology. 21: 1. doi:10.2981/wlb.00013. S2CID 83906946.
  193. ^ Kavčič, I.; Adamič, M.; Kaczensky, P.; Krofel, M.; Jerina, K. (2013). "Supplemental feeding with carrion is not reducing brown bear depredations on sheep in Slovenia". Ursus. 24 (#2): 111. doi:10.2192/URSUS-D-12-00031R1.1. S2CID 85579033.
  194. ^ 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.
  195. ^ a b c d e Herrero, Stepehen (1985). Bear Attacks: Their Causes and Avoidance. Edmonton: Hurtig Publishers Ltd.
  196. ^ a b Herrero, Stephen (2002), Bear Attacks: Their Causes and Avoidance (revised ed.), Lyons Press, ISBN 1-5857-4557-X
  197. ^ Smith, Tom S.; Herrero, Steven. . Alaska Science Center – Biological Science Office. Archived from the original on 14 August 2009.
  198. ^ , Digital Collegian, 4 April 2004, archived from the original on 9 March 2005
  199. ^ Rogers, Lynn L., , Bear.org, archived from the original on 16 October 2002
  200. ^ Medred, Craig (26 July 2011) Alaska bear attack: NOLS kids did a 'phenomenal job', Alaska Dispatch
  201. ^ 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.
  202. ^ Herrero, Stephen; Fleck, Susan (1990). "Injury to People Inflicted by Black, Grizzly or Polar Bears: Recent Trends and New Insights" (PDF). Bears: Their Biology and Management. 8: 25–32. doi:10.2307/3872900. JSTOR 3872900. Archived (PDF) from the original on 9 October 2022.
  203. ^ Ustinov, S. K. (1972). Cannibalism and attacks on humans by brown bears in Eastern Siberia. In Proc. Conf. Bear Ecology, Morphology, Protection, and Use. Soviet Union Acad. Sci., Moscow, USSR (pp. 85–87).
  204. ^ Bears besiege Russian mine after killing guards. The Times. 24 July 2008
  205. ^ Dinets, Vladimir. . Archived from the original on 15 December 2012.
  206. ^ . Tooth & Claw. Archived from the original on 16 May 2008.
  207. ^ 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.
  208. ^ Bear Caused Human Injuries and Deaths In Yellowstone National Park. Yellowstone-bearman.com (1 January 2000). Retrieved 15 September 2011.
  209. ^ Herrero, S.; Higgins, A. (2003). "Human injuries inflicted by bears in Alberta: 1960–98". Ursus. 14 (1): 44–54. JSTOR 3872956.
  210. ^ Smith, Herrero; DeBruyn, Wilde (2008). . North American Bear Center. Archived from the original on 1 January 2011.
  211. ^ Smith, Tom S. . Alaska Science Center – Biological Science Office. Archived from the original on 14 August 2009.
  212. ^ . Alaska Department of Public Safety. 19 November 2007. Archived from the original (Case Number: 07-96958) on 17 December 2007.
  213. ^ "Safety: bears and you". State of Alaska Department of Natural Resources.
  214. ^ 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.
  215. ^ "Karhu on Suomen kansalliseläin". yle.fi.
  216. ^ . Montana Historical Society. Archived from the original on 18 December 2009. Retrieved 18 August 2016.
  217. ^ . California State Library. Archived from the original on 5 January 2019. Retrieved 24 September 2011.

Notes

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

Bibliography

  • Heptner V.G.; Sludskii, A.A. (1992). Mammals of the Soviet Union, Volume II, Part 2. Leiden u.a.: Brill. ISBN 978-90-04-08876-4.
  • 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

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

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, north, america, populations, brown, bears, call. 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 1 3 In North America the populations of brown bears are called grizzly bears while the subspecies that inhabits the Kodiak Islands of Alaska is known as the Kodiak bear It is one of the largest living terrestrial members of the order Carnivora rivaled in size only by its closest relative the polar bear Ursus maritimus which is much less variable in size and slightly bigger on average 4 5 6 7 8 The brown bear s range includes parts of Russia Central Asia the Himalayas China Canada the United States Hokkaido Scandinavia Finland the Balkans the Picos de Europa and the Carpathian region especially Romania Iran Anatolia and the Caucasus 1 9 The brown bear is recognized as a national and state animal in several European countries 10 Brown bearTemporal range 0 5 0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Middle Pleistocene HoloceneKodiak bear on Kodiak IslandConservation statusLeast Concern IUCN 3 1 1 CITES Appendix II CITES 2 note 1 Scientific classificationKingdom AnimaliaPhylum ChordataClass MammaliaOrder CarnivoraFamily UrsidaeGenus UrsusSpecies U arctosBinomial nameUrsus arctosLinnaeus 1758Subspecies15 see text and articleBrown bear range mapWhile 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 As of 2012 update this and the American black bear are the only bear species not classified as threatened by the IUCN though the large sizes of both bears may be a disadvantage due to increased competition with humans 1 3 7 Populations that were hunted to extinction in the 19th and 20th centuries are the Atlas bear of North Africa and the Californian Ungavan 11 12 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 1 13 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 14 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 50 to 60 bears 10 15 Contents 1 Evolution and taxonomy 1 1 Generalized names and evolution 1 2 Scientific taxonomy 1 3 Hybrids 2 Description 2 1 Color 2 2 Cranial morphology and size 2 3 Claws and feet 3 Distribution and habitat 3 1 Conservation status 4 Behavior and life history 4 1 Communication 4 2 Home ranges 4 3 Reproduction 4 4 Dietary habits 4 5 Interspecific predatory relationships 4 6 Longevity and mortality 5 Relationship with humans 5 1 Conflicts between bears and humans 5 2 Bear encounters and attacks 5 3 History of defense from bears 5 4 Culture 6 See also 7 References 8 Notes 9 Bibliography 10 External linksEvolution and taxonomyThe 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 16 17 In the mid 19th century United States the brown bear was termed Old Ephraim and sometimes as Moccasin Joe 18 The scientific name of the brown bear Ursus arctos comes from the Latin ursus meaning bear 19 and from ἄrktos arktos the Greek word for bear 20 Generalized names and evolution Brown bears are thought to have evolved from Ursus etruscus in Asia 21 22 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 23 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 24 The oldest fossils positively identified as from this species occur in China from about 0 5 million years ago Brown bears entered Europe about 250 000 years ago and North Africa shortly after 21 25 Brown bear remains from the Pleistocene period are common in the British Isles where it is thought they might have outcompeted cave bears Ursus spelaeus The species entered Alaska 100 000 years ago though they did not move south until 13 000 years ago 21 It is speculated that brown bears were unable to migrate south until the extinction of the much larger giant short faced bear Arctodus simus 26 27 Several paleontologists suggest the possibility of two separate brown bear migrations inland brown bears also known as grizzlies are thought to stem from narrow skulled bears which migrated from northern Siberia to central Alaska and the rest of the continent while Kodiak bears descend from broad skulled bears from Kamchatka which colonized the Alaskan peninsula Brown bear fossils discovered in Ontario Ohio Kentucky and Labrador show that the species occurred farther east than indicated in historic records 21 In North America two types of the subspecies Ursus arctos horribilis are generally recognized the coastal brown bear and the inland grizzly bear these two types broadly define the range of sizes of all brown bear subspecies 13 Scientific taxonomy Main article Subspecies of brown bear Adult female Eurasian brown bear the nominate subspecies There are many methods used by scientists to define bear species and subspecies as no one method is always effective Brown bear taxonomy and subspecies classification has been described as formidable and confusing with few authorities listing the same specific set of subspecies 28 Genetic testing is now perhaps the most important way to scientifically define brown bear relationships and names Generally genetic testing uses the word clade rather than species because a genetic test alone cannot define a biological species Most genetic studies report on how closely related the bears are or their genetic distance There are hundreds of obsolete brown bear subspecies each with its own name and this can become confusing Hall 1981 lists 86 different types and even as many as 90 have been proposed 29 30 However recent DNA analysis has identified as few as five main clades which contain all extant brown bears 31 32 while a 2017 phylogenetic study revealed nine clades including one representing polar bears 33 As of 2005 update 15 extant or recently extinct subspecies were recognized by the general scientific community 34 35 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 36 Under some definitions the brown bear can be construed as the paraspecies for the polar bear 37 38 39 40 DNA analysis shows that apart from recent human caused population fragmentation 41 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 42 43 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 44 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 45 Brown bears are apparently divided into five different clades some of which coexist or co occur in different regions 3 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 known either as a pizzly bear or a grolar bear 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 46 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 Analyses of the genomes of bears have shown that introgression between species was widespread during the evolution of the genus Ursus 47 48 49 including the introgression of polar bear DNA introduced to brown bears during the Pleistocene A bear shot in autumn 1986 in Michigan US was thought by some to be a grizzly American black bear hybrid due to its unusually large size and its proportionately larger braincase and skull DNA testing was unable to determine whether it was a large American black bear or a grizzly bear 50 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 also 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 51 52 Skeleton 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 53 54 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 6 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 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 13 55 56 57 Color in Slovenia Brown bears are often not fully brown 58 They have long thick fur with a moderately long mane at the back of the neck which varies somewhat across the types 59 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 58 60 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 61 The winter fur is very thick and long especially in northern subspecies and can reach 11 to 12 centimetres 4 to 5 in at the withers The winter hairs are thin yet rough to the touch The summer fur is much shorter and sparser and its length and density varies geographically 62 Cranial morphology and size Skull Adults have massive heavily built concave skulls which are large in proportion to the body The forehead is high and rises steeply 61 The projections of the skull are well developed when compared to those of Asian black bears Ursus thibetanus the latter have sagittal crests not exceeding more than 19 20 of the total length of the skull while the former have sagittal crests comprising up to 40 41 of the skull s length Skull projections are more weakly developed in females than in males The braincase is relatively small and elongated There is a great deal of geographical variation in the skull and presents itself chiefly in dimensions 62 Grizzlies for example tend to have flatter profiles than European and coastal American brown bears 63 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 62 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 62 The teeth of brown bears reflect their dietary plasticity and are broadly similar to other bears excluding the two most herbivorous living bears the giant panda Ailuropoda melanoleuca and the spectacled bear Tremarctos ornatus which have blunt small premolars ideal for grinding down fibrous plants compared to the jagged premolars of ursid bears that at least seasonally often rely on flesh as a food source 64 65 The teeth are reliably larger than American black bears but average smaller in molar length than polar bears 66 67 Brown bears have the broadest skull of any extant ursine bear only the aforementioned most herbivorous living bears exceed them in relative breadth of the skull 13 55 68 Another extant ursine bear the sloth bear Melursus ursinus has a proportionately longer skull than the brown bear and can match the skull length of even large brown bear subtypes presumably as an aid for foraging heavily on insect colonies for which a long muzzle is helpful as an evolved feature in several unrelated mammalian groups 61 68 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 69 They are generally dark with a light tip with some forms having completely light claws 62 Brown bear claws are longer and straighter than those of American black bears Ursus americanus 61 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 both species of black bear although in rare cases adult female brown bears have been seen in trees 70 The claws of a polar bear are also quite different being notably shorter but broader with a strong curve and sharper point presumably both as an aid to traveling over ice sometimes nearly vertically and procuring active prey 25 71 The paws of the brown bear are quite large The rear feet of adult bears have been found to typically measure 21 to 36 cm 8 3 to 14 2 in long while the forefeet tend to measure about 40 less in length All four feet in average sized brown bears tend to be about 17 5 to 20 cm 6 9 to 7 9 in in width In large coastal or Kodiak bear males the hindfoot may measure up to 40 cm 16 in in length 28 5 cm 11 2 in in width while outsized Kodiak bears having had confirmed measurements of up to 46 cm 18 in along their rear foot 72 73 74 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 most bears of the species and also used heavily in den construction prior to hibernation 61 75 The brown bear s strength has been roughly estimated as 2 5 to 5 times that of a human 76 Distribution and habitatMain 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 77 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 78 The largest populations are in Russia with 120 000 79 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 32 000 individuals 80 In the lower 48 states they are repopulating slowly but steadily along the Rockies and the western Great Plains 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 81 82 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 83 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 84 85 in the south 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 They can also 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 72 This species inhabits the broadest range of habitats of any living bear species 72 They seem to have no altitudinal preferences and have been recorded from sea level to an elevation of 5 000 m 16 000 ft the latter in the Himalayas 72 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 72 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 update this and the American black bear are the only bear species not classified as threatened by the IUCN 1 3 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 86 The Syrian brown bear Ursus arctos syriacus is very rare and it has been extirpated from more than half of its historic range 87 One of the smallest bodied subspecies the Himalayan brown bear Ursus arctos isabellinus is critically endangered occupying only 2 of its former range and threatened by uncontrolled poaching for its body parts 14 The Marsican brown bear in central Italy is believed to have a population of just 50 60 bears 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 88 Studies have shown that activity throughout the range 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 89 90 91 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 92 93 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 94 95 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 e g using a barnacle covered rock to scratch its neck which requires advanced cognitive abilities 96 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 97 98 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 99 100 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 101 During combat bears use their paws to strike their opponents in the chest or shoulders and bite the head or neck 102 Communication 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 makes 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 102 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 102 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 13 100 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 103 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 13 75 55 104 In areas where food is abundant and concentrated such as coastal Alaska home ranges for females are up to 24 km2 9 3 sq mi and for males are up to 89 km2 34 sq mi Similarly in British Columbia bears of the two sexes travel relatively compact home ranges of 115 km2 44 sq mi and 318 km2 123 sq mi In Yellowstone National Park home ranges for females are up to 281 km2 108 sq mi and up to 874 km2 337 sq mi for males In Romania the largest home range was recorded for adult males 3 143 km2 1214 sq mi 105 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 13 72 75 100 A study of male inherited Y chromosome DNA sequence found that brown bears over the past few 10 000 years have shown strong male biased dispersal 106 That study found surprisingly similar Y chromosomes in brown bear populations as far apart as Norway and coastal Alaska indicating extensive gene flow across Eurasia and North America Notably this contrasts with genetic signals from female inherited mitochondrial DNA mtDNA where brown bears of different geographic regions typically show strong differences in their mtDNA a result of female philopatry Reproduction 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 72 55 107 Being serially monogamous brown bears remain with the same mate from a couple of days to a couple of weeks 13 108 Outside of this narrow time frame adult male and female brown bears show no sexual interest in each other 13 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 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 109 103 110 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 54 110 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 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 75 55 111 112 113 114 104 115 Paternity DNA tests have shown that littermates do not share the same father in up to 29 of litters 104 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 55 104 Copulation is vigorous and prolonged and can last up to an hour although the mean time is about 23 24 minutes 55 116 Grizzly bear cubs often imitate their mothers closely Males take no part in raising their cubs parenting is left entirely to the females 100 117 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 109 118 119 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 109 120 121 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 72 122 123 Older and larger females within a population tend to give birth to larger litters 124 The size of a litter also depends on factors such as geographic location and food supply 125 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 126 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 127 128 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 75 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 129 The cubs learn by following and imitating their mother s actions during the period they are with her 130 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 55 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 131 132 Brown bears practice infanticide as an adult male bear may kill the cubs of another 104 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 104 Cubs flee up a tree if available when they see a strange male bear and the mother often successfully defends them even though the male may be twice as heavy as she although females have been known to die in these confrontations 104 133 134 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 72 Throughout life this species is regularly curious about the potential of eating virtually any organism or object that they encounter Food that is both abundant and easily accessed or caught is preferred Their jaw structure has evolved to fit their dietary habits Their diet varies enormously throughout their differing areas based on opportunity Despite their reputation most brown bears are not highly carnivorous as they derive up to 90 of their dietary food energy from vegetable matter 135 They often feed on a variety of plant life including berries grasses flowers acorns and pine cones as well as fungi such as mushrooms 13 Among all bears brown bears are uniquely equipped to dig for tough foods such as roots bulbs 136 and shoots They use their long strong claws to dig out earth to reach the roots and their powerful jaws to bite through them 13 In spring winter provided carrion grasses shoots sedges moss 136 and forbs are the dietary mainstays for brown bears internationally 72 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 72 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 137 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 61 This species may eat birds and their eggs including almost entirely ground or rock nesting species 61 The diet may be supplemented by rodents or similar smallish mammals including marmots ground squirrels mice rats lemmings and voles 13 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 138 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 61 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 61 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 61 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 61 Large mammals preyed on can include various ungulate species such as elk moose caribou muskoxen and wild boar 13 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 139 It will also bite or swipe some prey to stun it enough to knock it over for consumption 140 To pick out young or infirm individuals bears will charge at herds so the slower moving and more vulnerable individuals will be made apparent Brown bears may also ambush young animals by finding them via scent 13 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 140 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 13 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 61 They also 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 13 Cannibalism is not unheard of though predation is not normally believed to be the primary motivation when brown bears attack each other 61 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 13 Plants and fruit farmed by humans are readily consumed as well including corn wheat sorghum melons and any form of berries 61 They will also feed at domestic bee yards readily consuming both honey and the brood grubs and pupae of the honey bee colony 61 Human foods and trash or refuse is eaten when possible When an open garbage dump was kept in Yellowstone brown bears were one of the most voracious and regular scavengers The dump was closed after both brown and American black bears came to associate humans with food and lost their natural fear of them 61 Interspecific predatory relationships Brown bear being followed by a wolf Adult bears are generally immune to predatory attacks except from tigers and other bears Siberian tigers prefer preying on young Ussuri brown bears but smaller adult female brown bears outside their dens may also be taken generally when lethargic from hibernation 141 142 143 Of 44 recorded encounters between the two predators 20 resulted in confrontations in 50 of these the bears were killed in 27 the tigers were killed and 23 of the cases ended with both animals surviving and parting ways 144 Some bears emerging from hibernation seek out tigers in order to steal their kills 145 Some large brown bears may actually benefit from the tiger s presence by appropriating tiger kills that the bears may not be able to successfully hunt themselves and follow tiger tracks 146 Geptner et al 1972 stated bears are generally afraid of tigers and change their path after coming across tiger trails 147 In the winters of 1970 1973 Yudakov and Nikolaev recorded 1 case of brown bear showing no fear of the tigers and another case of brown bear changing path upon crossing tiger tracks 148 Other researchers have observed bears following tiger tracks for various reasons 149 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 case of fully grown wolves being killed by a grizzly bear 150 Given the opportunity however both species will prey on the other s cubs 151 Conclusively the individual power of the bear against the collective strength of the wolf pack usually results in a long battle for kills or domination In some areas grizzly bears also regularly displace cougars from their kills 152 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 153 154 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 61 There is one record of a golden eagle preying on a brown bear cub 155 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 156 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 157 They will eat the fruit dropped by the Asian black bear from trees as they themselves are too large and cumbersome to climb 158 Improbably in the Himalayas Brown bears are reportedly intimidated by Asian black bears in confrontations 159 There has been a recent 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 160 and dead polar bear cubs have been found in brown bear dens 161 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 72 162 The lifespan of brown bears of both sexes within minimally hunted populations is estimated at an average of 25 years 163 The oldest wild brown bear on record was nearly 37 years old 163 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 13 54 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 164 Annual mortality for bears of any age is estimated at 10 in most protected areas 72 however the average annual mortality rate rises to an estimated 38 in hunted populations 72 Around 13 to 44 of cubs die within their first year even in well protected areas 13 Mortality rates of 75 100 among the cubs of any given year are not uncommon 165 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 13 75 134 166 By the second and third years of their lives the annual mortality rate among cubs in the care of their mothers drops to 10 15 75 Even in populations living in protected areas humans are still the leading cause of mortality for brown bears 3 163 167 The largest amount of legalized brown bear hunting occurs in Canada Finland Russia Slovakia and Alaska 72 Hunting is unregulated in many areas within the range of the brown bear Even where hunting is legally permitted most biologists feel that the numbers hunted are excessive considering the low reproduction rate and sparse distribution of the species 3 127 165 Brown bears are also killed in collisions with automobiles which is a significant cause of mortality in the United States and Europe 168 169 Relationship with humansConflicts between bears and humans Ancient depiction of a brown bear in the arena Papyrus 3053 Brown bears usually avoid areas where extensive development or urbanization has occurred unlike the smaller more inoffensive American black bear which can adapt to peri urban regions 170 171 172 173 Under many circumstances extensive human development may cause brown bears to alter their home ranges 174 175 High road densities both paved and gravel roads are often associated with higher mortality habitat avoidance and lower bear density 176 However brown bears can easily lose their natural cautiousness upon being attracted to human created food sources such as garbage dumps litter bins and dumpsters Brown bears may even venture into human dwellings or barns in search of food as humans encroach on bear habitats 177 In other areas such as Alaska dumps may continue to be an attractant for brown bears 178 179 In different parts of their distribution brown bears sometimes kill and eat domesticated animals 180 181 182 The saying A fed bear is a dead bear has come into use to popularize the idea that allowing a bear to scavenge human garbage such as trash cans and campers backpacks pet food or other food sources that draw the bear into contact with humans can result in the bear s death 183 Results of a 2016 study performed in a southeastern British Columbian valley indicate that areas where attractive bear food and concentrated human settlements overlap human bear conflict can create an ecological trap resulting in a lower apparent survival rate for brown bears as well as attracting additional bears and thereby causing overall population declines 184 When brown bears come to associate human activity with a food reward they are likely to continue to become emboldened the likelihood of human bear encounters increases as they may return to the same location despite relocation Relocation of the bear has been used to separate the bear from the human environment but it does not address the problem of the bear s newly learned association of humans with food or the environmental situations which created the human habituated bear Placing a bear in habitat used by other bears may lead to competition and social conflict and result in the injury or death of the less dominant bear 185 Yellowstone National Park a reserve located in the western United States contains prime habitat for the grizzly bear Ursus arctos horribilis and due to the enormous number of visitors human bear encounters are common The scenic beauty of the area has led to an influx of people moving into the area In addition because there are so many bear relocations to the same remote areas of Yellowstone and because male bears tend to dominate the center of the relocation zone female bears tend to be pushed to the boundaries of the region and beyond As a result a large proportion of repeat offenders bears that are killed for public safety are females This creates a further depressive effect on an already endangered subspecies The grizzly bear is officially described as Threatened in the U S Although the problem is most significant with regard to grizzlies these issues affect the other types of brown bears as well 186 187 188 In Europe part of the problem lies with shepherds over the past two centuries many sheep and goat herders have gradually abandoned the more traditional practice of using dogs to guard flocks which have concurrently grown larger Typically they allow the herds to graze freely over sizeable tracts of land As brown bears reclaim parts of their range they may eat livestock as sheep and goats are relatively easy for a bear to kill In some cases the shepherds shoot the bear thinking their livelihood is under threat Many are now better informed about the ample compensation available and will make a claim when they lose livestock to a bear 189 190 191 Another issue in several parts of their range in Europe is supplemental feeding stations where various kind of animal carrion is offered which are set up mainly in Scandinavia and eastern Europe both to support the locally threatened species and so humans can enjoy watching bears that may otherwise prove evasive Despite that most stations were cautiously set in remote areas far from human habitations some brown bears in such areas have become conditioned to associate humans with food and become excessively bold problem bears Also supplemental feeding appears to cause no decrease in livestock predation 192 193 Bear encounters and attacks A statue of the Ussuri brown bear from Hokkaido which perpetrated the worst brown bear attack in Japanese history killing seven people Brown bears seldom attack humans on sight and usually avoid people In Russia it is estimated that 1 in 1 000 on foot encounters with brown bears results in an attack 194 They are however unpredictable in temperament and may attack if they are surprised or feel threatened 195 There are an average of two fatal attacks by bears per year in North America 196 In Scandinavia there are only four known cases since 1902 of bear encounters which have resulted in death The two most common causes for bear attack are surprise and curiosity 197 Some types of bears such as polar bears are more likely to attack humans when searching for food while American black bears are much less likely to attack Despite their boldness and potential for predation if the bear is hungry polar bears rarely attack humans because they are infrequently encountered in the Arctic sea 196 Aggressive behavior in brown bears is favored by numerous selection variables Increased aggressiveness also assists female brown bears in better ensuring the survival of their young to reproductive age 198 Mothers defending cubs are the most prone to attacking being responsible for 70 of brown bear caused human fatalities in North America 199 Sows with cubs account for many attacks on humans by brown bears in North America Habituated or food conditioned bears can also be dangerous as their long term exposure to humans causes them to lose their natural shyness and in some cases to associate humans with food Small parties of one or two people are more often attacked by brown bears than large groups with only one known case of an attack on a group of six or more In that instance it is thought that due to surprise the grizzly bear may not have recognized the size of the group 200 In the majority of attacks resulting in injury brown bears precede the attack with a growl or huffing sound 195 In contrast to injuries caused by American black bears which are usually minor brown bear attacks more often tend to result in serious injury and in some cases death 195 Brown bears seem to confront humans as they would when fighting other bears they rise up on their hind legs and attempt to disarm their victims by biting and holding on to the lower jaw to avoid being bitten in turn 26 Due to the bears enormous physical strength even a single bite or swipe can be deadly as in tigers with some human victims having had their heads completely crushed by a bear bite 63 201 failed verification Most attacks occur in the months of July August and September the time when the number of outdoor recreationalists such as hikers or hunters is higher People who assert their presence through noises tend to be less vulnerable as they alert bears to their presence In direct confrontations people who run are statistically more likely to be attacked than those who stand their ground Violent encounters with brown bears usually last only a few minutes though they can be prolonged if the victims fight back 195 In Alberta two common behaviors by human hunters imitating the calls of deer to attract them and carrying ungulate carcasses seem to court aggressive behavior and lead to a higher rate of attack from grizzly bears 202 Attacks on humans are considered extremely rare in the former Soviet Union though exceptions exist in districts where they are not as often pursued by hunters 62 East Siberian brown bears for example tend to be much bolder toward humans than their shyer more persecuted European counterparts The delineation in Eurasia between areas where aggressiveness of brown bears tends to increase is the Ural Mountains although the brown bears of eastern Europe are somewhat more aggressive than those of western Europe 163 203 In 2008 a platinum mining compound in the Olyotorsky district of northern Kamchatka was besieged by a group of 30 bears who killed two guards and prevented workers from leaving their homes 204 10 people a year on average are killed by brown bears in Russia more than all the other parts of the brown bear s international range combined although Russia also holds more brown bears than all other parts of the world combined 205 In Scandinavia only three fatal attacks were recorded in the 20th century 206 In Japan a large brown bear nicknamed Kesagake 袈裟懸け kesa style slasher made history for causing 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 with possibly another three previous fatalities to its credit before being gunned down after a large scale beast hunt Today there is still a shrine at Rokusensawa 六線沢 where the event took place in memory of the victims of the incident 207 Within Yellowstone National Park injuries caused by grizzly attacks in developed areas averaged approximately one per year during the 1930s through to the 1950s though it increased to four per year during the 1960s They then decreased to one injury every two years during the 1970s Between 1980 and 2002 there have been only two human injuries caused by grizzly bears in a developed area Although grizzly attacks were rare in the backcountry before 1970 the number of attacks increased to an average of approximately one per year during the 1970s 1980s and 1990s 208 In Alberta from 1960 to 1998 the number of attacks by grizzly bears ending in injury were nearly three times more common than attacks ending in injury by American black bears despite the American black bear being an estimated 38 times more numerous in the province than the grizzly bear 209 History of defense from bears See also Bear danger can of bear spray 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 210 Carrying pepper spray is highly recommended by many authorities when traveling in bear country however carrying two means of deterrent one of which is a large caliber gun is also advised Solid shotgun slugs or three buckshot rounds or a pistol of 44 caliber or more is suggested if a heavy hunting rifle is not available Guns remain a viable last resort option to be used in defense of life from aggressive brown bears 211 Too often people do not carry a proper caliber weapon to neutralize the bear According to the Alaska Science Center a 12 gauge shotgun with slugs has been the most effective weapon There have been fewer injuries as a result of only carrying lethal loads in the shotgun as opposed to deterrent rounds State of Alaska Defense of Life or Property DLP laws require one to report the kill to the authorities and salvage the hide skull and claws 212 A page at the State of Alaska Department of Natural Resources website offers information about how to select a gun that will stop a bear 12 gauge shotgun or 300 mag rifle 213 Campers are often told to wear bright colored red ribbons and bells and carry whistles to ward off bears They are told to look for grizzly bear dung in camping areas and be careful to carry the bells and whistles in those areas Grizzly bear dung is difficult to differentiate from American black bear dung as diet is in a constant state of flux depending on the availability of seasonal food items If a bear is killed near camp the bear s carcass must be adequately disposed of including entrails and blood if possible Failure to move the carcass has often resulted in it attracting other bears and further exacerbating a bad situation Moving camps immediately is another recommended method 195 Culture The Story of the Three Bears illustration from Childhood s Favorites and Fairy Stories Brown bears often figure into the literature of Europe and North America in particular that which is written for children The Brown Bear of Norway is a Scottish fairy tale telling the adventures of a girl who married a prince magically turned into a bear and who managed to get him back into a human form by the force of her love and after many trials and difficulties With Goldilocks and the Three Bears a story from England the Three Bears are usually depicted as brown bears In German speaking countries children are often told the fairytale of Snow White and Rose Red the handsome prince in this tale has been transfigured into a brown bear In the United States parents often read their preschool age children the book Brown Bear Brown Bear What Do You See to teach them their colors and how they are associated with different animals 214 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 215 The grizzly bear is the state animal of Montana 216 The California golden bear is the state animal of California 217 Both animals are subspecies of the brown bear and the species was extirpated from the latter state 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 The brown bear is depicted on the reverse of the Croatian 5 kuna coin minted since 1993 The Bundesliga club Bayern Munich has a brown bear mascot named Berni The Chicago National Football League NFL franchise is named the Bears In this context no differentiation between American black and brown bears is needed The school mascot for Bob Jones University Brown University Cornell University George Fox University the University of Alberta the University of California Berkeley the University of California Los Angeles the University of California Riverside and numerous American high schools is the brown bear In the town of Prats de Mollo in Vallespir French Catalonia a bear festival festa de l os is celebrated annually at the beginning of spring in which the locals dress up as bears cover themselves with soot or coal and oil and attack the onlookers attempting to get everyone dirty The festival ends with the ball de l os bear dance See alsoList of fatal bear attacks in North America Sankebetsu brown bear incidentReferences a b c d e f 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 Retrieved 14 January 2022 a b c d e f 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 Boddington Craig 2004 Fair Chase in North America Illustrations by Carlson Ken Boone and Crockett Club p 45 ISBN 978 0 940864 47 4 Martin A P Palumbi S R 1993 Body size metabolic rate generation time and the molecular clock Proceedings of the National Academy of Sciences of the United States of America 90 9 4087 4091 Bibcode 1993PNAS 90 4087M doi 10 1073 pnas 90 9 4087 PMC 46451 PMID 8483925 a b Christiansen P 1999 What size were Arctodus simus and Ursus spelaeus Carnivora Ursidae Annales Zoologici Fennici 36 2 93 102 JSTOR 23735739 a b Servheen C Darling L M Archibald W R 1990 The status and conservation of the bears of the world International Association for Bear Research and Management Mammalian Species Ursus arctos PDF American Society of Mammalogists Smith College 23 April 1993 Archived from the original PDF on 31 March 2017 White Paul Brown Bear Transylvania Wildlife Project Archived from the original on 19 May 2013 a b Zedrosser A Dahle B Swenson J E Gerstl N 2001 Status and management of the brown bear in Europe PDF Ursus 12 9 20 Archived PDF from the original on 9 October 2022 Busch Robert 2004 The Grizzly Almanac Globe Pequot Press pp 11 14 ISBN 978 1 5922 8320 0 Retrieved 21 October 2014 Elson C S 1954 Further Evidence about the Barren Ground Grizzly Bear in Northeast Labrador and Quebec Journal of Mammalogy 35 3 345 357 doi 10 2307 1375959 JSTOR 1375959 a b c d e f g h i j k l m n o p q r s t Mammalian Species Ursus arctos PDF American Society of Mammalogists Smith College 23 April 1993 Archived from the original PDF on 31 March 2017 a b Himalayan brown bears now critically endangered Archived 16 October 2018 at the Wayback Machine Euronews com 6 January 2014 Boscagli G 1990 Marsican brown bear population in central Italy Status report 1985 Aquilo Series Zoologica 27 81 83 Harper Douglas bruin Online Etymology Dictionary Bruin Encyclopaedia Britannica Hunting the Grisly and other Sketches FullTextArchive com Retrieved 15 September 2011 Liddell Henry George and Scott Robert Ursus A Latin Dictionary Perseus Digital Library Liddell Henry George and Scott Robert Arktos A Greek English Lexicon Perseus Digital Library 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 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 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 a b Geist Valerius 1989 Did Large Predators keep Humans out of North America PDF in Clutton Brock Juliet ed The Walking larder patterns of domestication pastoralism and predation Unwin Hyman pp 282 294 ISBN 0 0444 5013 3 archived from the original PDF on 16 December 2008 Kurten B Anderson E 1974 Association of Ursus arctos and Arctodus simus Mammalia Ursidae in the late Pleistocene of Wyoming Breviora 426 1 6 Wilson D E Ruff S 1999 The Smithsonian Book of North American Mammals Washington D C Smithsonian Institution Press 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 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 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 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 Shields G F Kocher T D 1991 Phylogenetic relationships of North American ursids based on analysis of mitochondrial DNA Evolution 45 1 218 221 doi 10 2307 2409495 JSTOR 2409495 PMID 28564083 Kurten B 1964 The evolution of the polar bear Ursus maritimus Phipps Acta Zoologica Fennica 108 1 26 Ingolfsson olafur Wiig Oystein 2009 Late Pleistocene fossil find in Svalbard the oldest remains of a polar bear Ursus maritimus Phipps 1744 ever discovered Polar Research 28 3 455 Bibcode 2009PolRe 28 455I doi 10 1111 j 1751 8369 2008 00087 x S2CID 56292690 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 doi 10 1002 wmon 6 ISSN 1938 5455 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 Waits L P et al 1998 Mitochondrial DNA Phylogeography of the North American Brown Bear and Implications for Conservation Conservation 12 2 408 417 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 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 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 Smith Richard P 2007 Hybrid Black Bear Black Bear Hunting Stackpole Books ISBN 978 0 8117 0269 0 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 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 Parker S P 1990 Grzimek s encyclopedia of mammals McGraw Hill New York ISBN 0 07 909508 9 set a b c Nowak R M 1999 Walker s mammals of the world Vol 1 JHU Press a b c d e f g h Feldhamer G A Thompson B C Chapman J A 2003 Wild mammals of North America biology management and conservation JHU Press ISBN 9780801874161 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 Steyaert Sam M J G Endrestol Anders Hacklander Klaus Swenson Jon E Zedrosser Andreas 2012 The mating system of the brown bear Ursus arctos Mammal Review 42 12 doi 10 1111 j 1365 2907 2011 00184 x 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 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 q Bear Anatomy and Physiology from Gary Brown s The Great Bear Almanac Lyons amp Burford Publishers 1993 a b c d e f 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 USA Science Publishers Inc ISBN 1 886106 81 9 a b The living animals of the world a popular natural history with one thousand illustrations Volume 1 Mammals by Cornish C J 1858 1906 Selous Frederick Courteney 1851 1917 Johnston Harry Hamilton Sir 1858 1927 Maxwell Herbert Sir published by New York Dodd Mead and Company Archive org Retrieved 15 September 2011 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 Slaughter B H Pine R H Pine N E 1974 Eruption of cheek teeth in insectivora and carnivora Journal of Mammalogy 55 1 115 25 doi 10 2307 1379261 JSTOR 1379261 PMID 4819587 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 a b Christiansen P 2007 Evolutionary implications of bite mechanics and feeding ecology in bears Journal of Zoology 272 4 423 doi 10 1111 j 1469 7998 2006 00286 x Heptner Sirenia and carnivora Sea cows Wolves and Bears Seryodkin I V 2016 Behavior of Brown Bears During Feeding in the Sikhote Alin Achievements in the Life Sciences Stirling I amp Guravich D 1998 Polar bears University of Michigan Press a b c d e f g h i j k l m n o Carnivores of the World by Dr Luke Hunter Princeton University Press 2011 ISBN 9780691152288 Wood Gerald 1983 The Guinness Book of Animal Facts and Feats ISBN 978 0 85112 235 9 Whitaker J O amp Elman R 1996 National Audubon Society Field Guide to North American Mammals p 992 New York Knopf a b c d e f g 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 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 Retrieved 2 January 2010 Brown Bear Species WWF WWF Brown Bear Rolling Hills Wildlife Adventure Archived from the original on 10 November 2016 Retrieved 2 December 2013 Brown Bear Research in Alaska Alaska Department of Fish and Game Retrieved 4 April 2012 Neixen quatre cries d os bru als Pirineus elperiodico cat in Catalan and Spanish 2 August 2010 Retrieved 19 November 2010 Situacio preocupant de l os bru als Pirineus in Catalan ecologistasenaccion org May 2010 Retrieved 19 November 2010 Osos el desafio de una poblacion creciente ABC in Spanish Spain 28 September 2013 Martinez Abrain A Jimenez J Oro D 2018 Pax Romana refuge abandonment and spread of fearless behavior in a reconciling world Animal Conservation John Wiley amp Sons Inc Zoological Society of London ZSL 22 1 3 13 doi 10 1111 acv 12429 ISSN 1367 9430 S2CID 89938098 Bear ARKTOYROS 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 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 doi 10 1111 j 1472 4642 2009 00586 x S2CID 21666120 Brown Grizzly Bear Facts North American Bear Center Archived from the original on 24 May 2013 Retrieved 16 August 2013 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 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 Egbert Allan L Stokes Allen W Egbert A L 1974 The social behaviour of brown bears on an Alaskan salmon stream PDF International Conference Bear Res And Manage 3 41 56 doi 10 2307 3872753 JSTOR 3872753 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 doi 10 1046 j 1365 2656 2003 00737 x JSTOR 3505643 PMID 30893970 S2CID 67818528 Stonorov Derek and Stokes Allen W 1972 Social Behavior of the Alaska Brown Bear Panel 4 Bear Behaviour a b c Egbert A L 1978 The Social Behavior of Brown Bears at McNeil River Alaska Ph D thesis Utah State University a b Pearson A M 1975 The northern interior grizzly bear Ursus arctos L Information Canada a b c d e f g 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 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 Bidon T Janke A Fain S R Eiken H G Hagen S B Saarma U Hallstrom B M Lecomte N Hailer F 2014 Brown and Polar Bear Y Chromosomes Reveal Extensive Male Biased Gene Flow within Brother Lineages Molecular Biology and Evolution 31 6 1353 1363 doi 10 1093 molbev msu109 PMID 24667925 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 Types of Bears Information on Specific Bear Species 2009 a b c 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 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 Nawaz Muhammad Ali Swenson Jon E Zakaria Vaqar 2008 Pragmatic management increases a flagship species the Himalayan brown bears in Pakistan s Deosai National Park Biological Conservation 141 9 2230 doi 10 1016 j biocon 2008 06 012 Paralikidis N P Papageorgiou N K Kontsiotis V J Tsiompanoudis A C 2010 The dietary habits of the brown bear Ursus arctos in western Greece Mammalian Biology Zeitschrift fur Saugetierkunde 75 29 doi 10 1016 j mambio 2009 03 010 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 Naves J Fernandez Gil A Rodriguez C Delibes M 2006 Brown Bear Food Habits at the Border of Its Range A Long Term Study Journal of Mammalogy 87 5 899 doi 10 1644 05 MAMM A 318R2 1 hdl 10261 50290 S2CID 85351746 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 G B 1966 Terrestrial mammals of Western Europe G T Foulic amp Co Ltd London UK 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 Frkovic A Huber D amp Kusak J 2001 Brown bear litter sizes in Croatia Ursus 12 103 105 JSTOR 3873235 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 107 365 367 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 Couturier MA 1954 L ours Brun Ursus arctos L Couturier Grenoble France a b Mallinson J 1978 The Shadow of Extinction Europe s Threatened Mammals Macmillan London 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 Brown Bear Reproduction Shadowofthebear com Archived from the original on 1 May 2008 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 a b 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 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 Retrieved 29 January 2010 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 David Prynn 2004 Amur tiger Russian Nature Press p 115 Frasef A 2012 Feline Behaviour and Welfare CABI pp 72 77 ISBN 978 1 84593 926 7 Seryodkin et al 2003 Denning ecology of brown bears and Asiatic black bears in the Russian Far East Ursus 14 2 159 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 amp Management International Association for Bear Research and Management p 64 Archived PDF from the original on 9 October 2022 Matthiessen Peter Hornocker Maurice 2001 Tigers In The Snow North Point Press ISBN 0 86547 596 2 Miquelle D G Smirnov E N Goodrich J M 2005 1 Tigers of Sikhote Alin Zapovednik ecology and conservation Vladivostok Russia PSP Geptner V G Sludskii A A 1972 Mlekopitaiuscie Sovetskogo Soiuza Vyssaia Skola Moskva In Russian English translation Heptner V G Sludskii A A Bannikov A G 1992 Mammals of the Soviet Union Volume II Part 2 Carnivora Hyaenas and Cats Smithsonian Institution and the National Science Foundation Washington DC pp 95 202 Yudakov A G Nikolaev I G 2004 Hunting Behavior and Success of the Tigers Hunts The Ecology of the Amur Tiger based on Long Term Winter Observations in 1970 1973 in the Western Sector of the Central Sikhote Alin Mountains english translation ed Institute of Biology and Soil Science Far Eastern Scientific Center Academy of Sciences of the USSR Archived from the original on 19 February 2020 Retrieved 17 September 2014 Seryodkin I V Goodrich J M Kostyrya A V Schleyer B O Smirnov E N Kerley L L 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 Goodrich J M eds Tigers of Sikhote Alin Zapovednik Ecology and Conservation in Russian Vladivostok Russia PSP pp 156 163 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 M and S Negri eds 2009 Cougar ecology and conservation University of Chicago Press Chicago IL ISBN 0226353443 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 The Intellectual observer review of natural history microscopic research and recreative science Groombridge 1865 Adams Andrew Leith 1867 Wanderings of a naturalist in India the western Himalayas and Cashmere Edmonston and Douglas 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 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 a b c d McDonald D amp Norris S 2001 The new encyclopedia of mammals Oxford University Press 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 a b Servheen C 1987 Grizzly bear compendium Interagency Grizzly Bear Committee Palomares F Caro T M 1999 Interspecific Killing among Mammalian Carnivores PDF The American Naturalist 153 5 492 508 doi 10 1086 303189 hdl 10261 51387 PMID 29578790 S2CID 4343007 Archived PDF from the original on 9 October 2022 Brown Grizzly Bear Hunting in Alaska Alaska Department of Fish and Game Huber D Kusak J Frkovic A 1998 Traffic kills of brown bears in Gorski kotar Croatia Ursus 10 167 171 JSTOR 3873124 Servheen C Waller J amp Kasworm W 1998 Fragmentation effects of high speed highways on grizzly bear populations shared between the United States and Canada International Conference on Wildlife Ecology and Transportation ICOWET 1998 Kaczensky P Blazic M Gossow H 2004 Public attitudes towards brown bears Ursus arctos in Slovenia Biological Conservation 118 5 661 doi 10 1016 j biocon 2003 10 015 Woodroffe R 2000 Predators and people Using human densities to interpret declines of large carnivores PDF Animal Conservation 3 2 165 doi 10 1111 j 1469 1795 2000 tb00241 x S2CID 84430290 Archived PDF from the original on 9 October 2022 Martin J Basille M Van Moorter B Kindberg J Allaine D Swenson J E 2010 Coping with human disturbance spatial and temporal tactics of the brown bear Ursus arctos Canadian Journal of Zoology 88 9 875 doi 10 1139 Z10 053 S2CID 44121835 Lariviere S 2001 Ursus americanus PDF Mammalian Species 647 1 11 doi 10 1644 1545 1410 2001 647 lt 0001 ua gt 2 0 co 2 S2CID 198968922 Archived from the original PDF on 16 March 2016 Retrieved 14 December 2016 Mattson D J 1990 Human impacts on bear habitat use PDF Bears Their Biology and Management 8 33 56 doi 10 2307 3872901 JSTOR 3872901 Archived PDF from the original on 9 October 2022 Elgmork K 1978 Human impact on a brown bear population Ursus arctos L Biological Conservation 13 2 81 doi 10 1016 0006 3207 78 90063 0 Lamb C T Mowat G Reid A Smit L Proctor M McLellan B N Nielsen S E Boutin S 2018 Effects of habitat quality and access management on the density of a recovering grizzly bear population Journal of Applied Ecology 55 3 1406 1417 doi 10 1111 1365 2664 13056 Stringham S F 1986 Effects of climate dump closure and other factors on Yellowstone grizzly bear litter size Bears Their Biology and Management 6 33 39 doi 10 2307 3872803 JSTOR 3872803 Peirce K N Van Daele L J 2006 Use of a garbage dump by brown bears in Dillingham Alaska Ursus 17 2 165 177 doi 10 2192 1537 6176 2006 17 165 uoagdb 2 0 co 2 JSTOR 3873094 S2CID 54802857 Robbins C T Schwartz C C Felicetti L A 2004 Nutritional ecology of ursids A review of newer methods and management implications PDF Ursus 15 2 161 171 doi 10 2192 1537 6176 2004 015 lt 0161 NEOUAR gt 2 0 CO 2 S2CID 54680694 Archived PDF from the original on 9 October 2022 Clevenger A P Purroy F J Pelton M R 1992 Food habits of brown bears Ursus arctos in the Cantabrian Mountains Spain Journal of Mammalogy 73 2 415 421 doi 10 2307 1382077 JSTOR 1382077 Chauhan N P S 2003 Human casualties and livestock depredation by black and brown bears in the Indian Himalaya 1989 98 PDF Ursus 84 87 Archived PDF from the original on 9 October 2022 Sagor J T Swenson J E Roskaft E 1997 Compatibility of brown bear Ursus arctos and free ranging sheep in Norway Biological Conservation 81 1 2 91 doi 10 1016 S0006 3207 96 00165 6 Herrero S 2002 Bear attacks their causes and avoidance Globe Pequot ISBN 978 1585745579 Lamb C T Mowat G McLellan B N Nielsen S E Boutin S 2016 Forbidden fruit human settlement and abundant fruit create an ecological trap for an apex omnivore Journal of Animal Ecology 86 1 55 65 doi 10 1111 1365 2656 12589 PMID 27677529 S2CID 3347450 Relocation Get Bear Smart Society Archived from the original on 13 May 2006 Gunther K A Haroldson M A Frey K Cain S L Copeland J Schwartz C C 2004 Grizzly bear human conflicts in the Greater Yellowstone ecosystem 1992 2000 PDF Ursus 15 1 10 22 doi 10 2192 1537 6176 2004 015 lt 0010 gbcitg gt 2 0 co 2 S2CID 86821625 Archived PDF from the original on 9 October 2022 Blanchard B M Knight R R 1995 Biological consequences of relocating grizzly bears in the Yellowstone ecosystem The Journal of Wildlife Management 59 3 560 565 doi 10 2307 3802463 JSTOR 3802463 Brannon R D 1987 Nuisance grizzly bear Ursus arctos translocations in the greater Yellowstone area Canadian Field Naturalist 101 4 569 575 Archived from the original on 19 December 2019 Retrieved 12 December 2016 Mustoni A Carlini E Chiarenzi B et al 2003 Planning the Brown Bear Ursus arctos reintroduction in the Adamello Brenta Natural Park Hystrix The Italian Journal of Mammalogy 14 1 2 doi 10 4404 hystrix 14 1 2 4313 Dahle B Sorensen O J Wedul E H Swenson J E Sandegren F 1998 The diet of brown bears Ursus arctos in central Scandinavia effect of access to free ranging domestic sheep Ovis aries Wildlife Biology 4 3 147 158 doi 10 2981 wlb 1998 017 S2CID 90983953 Mertens A Promberger C 2001 Economic aspects of large carnivore livestock conflicts in Romania Ursus 12 173 180 JSTOR 3873246 Kavcic I Adamic M Kaczensky P Krofel M Kobal M Jerina K 2015 Fast food bears brown bear diet in a human dominated landscape with intensive supplemental feeding Wildlife Biology 21 1 doi 10 2981 wlb 00013 S2CID 83906946 Kavcic I Adamic M Kaczensky P Krofel M Jerina K 2013 Supplemental feeding with carrion is not reducing brown bear depredations on sheep in Slovenia Ursus 24 2 111 doi 10 2192 URSUS D 12 00031R1 1 S2CID 85579033 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 d e Herrero Stepehen 1985 Bear Attacks Their Causes and Avoidance Edmonton Hurtig Publishers Ltd a b Herrero Stephen 2002 Bear Attacks Their Causes and Avoidance revised ed Lyons Press ISBN 1 5857 4557 X Smith Tom S Herrero Steven Ursus arctos californicus Alaska Science Center Biological Science Office Archived from the original on 14 August 2009 Why are grizzly bears more aggressive than our black bears Digital Collegian 4 April 2004 archived from the original on 9 March 2005 Rogers Lynn L How Dangerous are Black Bears Bear org archived from the original on 16 October 2002 Medred Craig 26 July 2011 Alaska bear attack NOLS kids did a phenomenal job Alaska Dispatch 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 Herrero Stephen Fleck Susan 1990 Injury to People Inflicted by Black Grizzly or Polar Bears Recent Trends and New Insights PDF Bears Their Biology and Management 8 25 32 doi 10 2307 3872900 JSTOR 3872900 Archived PDF from the original on 9 October 2022 Ustinov S K 1972 Cannibalism and attacks on humans by brown bears in Eastern Siberia In Proc Conf Bear Ecology Morphology Protection and Use Soviet Union Acad Sci Moscow USSR pp 85 87 Bears besiege Russian mine after killing guards The Times 24 July 2008 Dinets Vladimir Brown Bears of Russia Archived from the original on 15 December 2012 Brown Bear Ursus arctos Tooth amp Claw Archived from the original on 16 May 2008 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 Bear Caused Human Injuries and Deaths In Yellowstone National Park Yellowstone bearman com 1 January 2000 Retrieved 15 September 2011 Herrero S Higgins A 2003 Human injuries inflicted by bears in Alberta 1960 98 Ursus 14 1 44 54 JSTOR 3872956 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 Smith Tom S Brown Bear Projects at the Alaska Science Center Alaska Science Center Biological Science Office Archived from the original on 14 August 2009 Alaska State Troopers Press Release of Monday November 19 2007 Alaska Department of Public Safety 19 November 2007 Archived from the original Case Number 07 96958 on 17 December 2007 Safety bears and you State of Alaska Department of Natural Resources 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 Karhu on Suomen kansalliselain yle fi 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 Notes Populations of Bhutan China Mexico and Mongolia are included in Appendix I All other populations are included in Appendix II BibliographyHeptner V G Sludskii A A 1992 Mammals of the Soviet Union Volume II Part 2 Leiden u a Brill ISBN 978 90 04 08876 4 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 Wikimedia Commons has media related to Ursus arctos category 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 1132848624, wikipedia, wiki, book, books, library,

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