Brown bear

The brown bear (Ursus arctos) is a large bear native to Eurasia and North America.
Of the carnivorans, it is rivaled in size only by its closest relative, the polar
bear, which is much less variable in size and slightly bigger on average. The brown
bear is a sexually dimorphic species, as adult males are larger and more compactly
built than females. The fur ranges in color from cream to reddish to dark brown. It
has evolved large hump muscles, unique among bears, and paws up to 21 cm (8.3 in)
wide and 36 cm (14 in) long, to effectively dig through dirt. Its teeth are similar
to those of other bears and reflect its dietary plasticity.

Throughout the brown bear's range, it inhabits mainly forested habitats in

elevations of up to 5,000 m (16,000 ft). It is omnivorous, and consumes a variety
of plant and animal species. Contrary to popular belief, the brown bear derives 90%
of its diet from plants. When hunting, it will target animals as small as rodents,
to those as large as moose or muskoxen. In parts of coastal Alaska, brown bears
predominantly feed on spawning salmon that come near shore to lay their eggs. For
most of the year, it is a usually solitary animal that associates only when mating
or raising cubs. Females give birth to an average of one to three cubs that remain
with their mother for 1.5 to 4.5 years. It is a long-lived animal, with an average
lifespan of 25 years in the wild. Relative to its body size, the brown bear has an
exceptionally large brain. This large brain allows for high cognitive abilities,
such as tool use. Attacks on humans, though widely reported, are generally rare.

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

The brown bear is considered to be one of the most popular of the world's
charismatic megafauna. It has been kept in zoos since ancient times, and has been
tamed and trained to perform in circuses and other acts. For thousands of years,
the brown bear has had a role in human culture, and is often featured in
literature, art, folklore, and mythology.

Etymology
The brown bear is sometimes referred to as the bruin, from Middle English. This
name originated in the fable History of Reynard the Fox, translated by William
Caxton, from the Middle Dutch word bruun or bruyn, meaning "brown".[3][4] In the
mid-19th-century United States, the brown bear was given the nicknames "Old
Ephraim" and "Moccasin Joe".[5]

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

Evolution and taxonomy

Taxonomy and subspecies
Main article: Subspecies of brown bear
A bear in a wooded area
Adult female Eurasian brown bear, the nominate subspecies
Carl Linnaeus scientifically described the species under the name Ursus arctos in
the 1758 edition of Systema Naturae.[8] Brown bear taxonomy and subspecies
classification has been described as "formidable and confusing", with few
authorities listing the same set of subspecies.[9] There are hundreds of obsolete
brown-bear subspecies. As many as 90 subspecies have been proposed.[10][11] A 2008
DNA analysis identified as few as five main clades, which comprise all extant brown
bear species, while a 2017 phylogenetic study revealed nine clades, including one
representing polar bears.[12][13] As of 2005, 15 extant, or recently extinct,
subspecies were recognized by the general scientific community.[14]

DNA analysis shows that, apart from recent, human-caused population fragmentation,
[15] 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.[16][17] 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.[18]

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 the movement of males, to
the point where their nuclear genomes indicate more than 90% brown bear ancestry.
[19] MtDNA analysis revealed that brown bears are apparently divided into five
different clades, some of which coexist or co-occur in different regions.[20]

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

A possible phylogeny based on complete mitochondrial DNA sequences from Yu et al.

(2007).[21]
Ursidae
Giant panda (Ailuropoda melanoleuca)

Spectacled bear (Tremarctos ornatus)

Ursinae
Sloth bear (Melursus ursinus)

Sun bear (Helarctos malayanus)

Asian black bear (Ursus thibetanus)

American black bear (Ursus americanus)

Polar bear (Ursus maritimus)

Brown bear (Ursus arctos)

The polar bear and the brown bear form a close grouping, while the relationships of
the other species are not very well resolved.[22]
A more recent phylogeny based on the genetic study of Kumar et al. (2017).[23]
Ursidae
Giant panda (Ailuropoda melanoleuca)

Spectacled bear (Tremarctos ornatus)

Ursinae
Sloth bear (Melursus ursinus)

Sun bear (Helarctos malayanus)

Asian black bear (Ursus thibetanus)

American black bear (Ursus americanus)

Polar bear (Ursus maritimus)

Brown bear (Ursus arctos)

The study concludes that Ursine bears originated around five million years ago and
show extensive hybridization of species in their lineage.[23]

The brown bear is thought to have evolved from the Etruscan bear (Ursus etruscus)
in Asia during the early Pliocene.[24][25] 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.[26] The oldest brown bear fossils occur in
Asia from about 500,000 to 300,000 years ago.[27][28] They entered Europe 250,000
years ago and North Africa shortly after.[24] Brown bear remains from the
Pleistocene period are common in the British Isles, where, amongst other factors,
they may have contributed to the extinction of cave bears (Ursus spelaeus).[29]

Brown bears first emigrated to North America from Eurasia via Beringia during the
Illinoian Glaciation.[30] Genetic evidence suggests that several brown bear
populations migrated into North America, aligning with the glacial cycles of the
Pleistocene. The founding population of most North American brown bears arrived
first, with the genetic lineage developing around ~177,000 BP. Genetic divergences
suggest that brown bears first migrated south during MIS-5 (~92,000–83,000 BP),
upon the opening of the ice-free corridor,[30][31][32] After a local extinction in
Beringia ~33,000 BP, two new but closely related lineages repopulated Alaska and
northern Canada from Eurasia after the Last Glacial Maximum (>25,000 BP).[30]

Brown-bear fossils discovered in Ontario, Ohio, Kentucky, and Labrador show that
the species occurred farther east than indicated in historic records.[24] In North
America, two types of the subspecies Ursus arctos horribilis are generally
recognized—the coastal brown bear and the inland grizzly bear.[33]

Hybrids
See also: Grizzly–black bear hybrid and Grizzly–polar bear hybrid
A photo of a bear walking in tall grass
Possible grizzly–black bear hybrid in the Yukon Territory
A grizzly–polar bear hybrid is a rare ursid hybrid resulting from a crossbreeding
of a brown bear with a polar bear. It has occurred both in captivity and in the
wild. In 2006, the occurrence of this hybrid 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.[34] 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).[35] Analyses of the genomes of bears have shown
that introgression between species was widespread during the evolution of the genus
Ursus,[36] including the introgression of polar-bear DNA introduced to brown bears
during the Pleistocene.[37]

Description
Size
A photo of a smiling brown bear standing by a lake
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, as most accepted subtypes vary widely in
size. This is in part due to sexual dimorphism, as male brown bears average at
least 30% larger than females in most subtypes. Individual bears vary in size
seasonally, weighing the least in spring due to lack of foraging during
hibernation, and the most in late fall, after a period of hyperphagia to put on
additional weight to prepare for hibernation.[38][39]

Brown bear skeleton

Brown bears generally weigh 80 to 600 kg (180 to 1,320 lb), with males outweighing
females.[40] They have 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 (28 to 60 in). The tail is relatively
short, as in all bears, ranging from 6 to 22 cm (2.4 to 8.7 in) in length.[41][42]
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.[43] Brown bears of the interior are generally smaller,
being around the same weight as an average lion, at an 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.[44] The average weight of adult male bears, from 19
populations, was found to be 217 kg (478 lb) while adult females from 24
populations were found to average 152 kg (335 lb).[33][45]

Coloration

A brown bear at Whipsnade Zoo

Brown bears are often not fully brown.[46] They have long, thick fur, with a
moderately elongated mane at the back of the neck which varies somewhat across bear
types.[47] 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.[46][48] Even within well-defined subspecies,
individuals may show highly variable hues of brown. North American grizzlies can be
from 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.[49] The brown bear's winter fur is very thick and long, especially in
northern subspecies, and can reach 11 to 12 cm (4.3 to 4.7 in) at the withers. The
winter hairs are thin, yet rough to the touch. The summer fur is much shorter and
sparser, with its length and density varying among geographic ranges.[50]

Cranial morphology and size

Skull: side view (top), upper view (left) and lower view (right)
Adults have massive, heavily built, concave skulls, which are large in proportion
to the body.[49] The projections of the skull are well developed.[50] Skull lengths
of Russian brown bears tend to be 31.5 to 45.5 cm (12.4 to 17.9 in) for males, and
27.5 to 39.7 cm (10.8 to 15.6 in) for females. Brown bears have the broadest skull
of any extant ursine bear.[44] The width of the zygomatic arches in males is 17.5
to 27.7 cm (6.9 to 10.9 in), and 14.7 to 24.7 cm (5.8 to 9.7 in) in females.[50]
Brown bears have strong jaws: the incisors and canine teeth are large, with the
lower canines 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 its alveolus in the jaw. The first three molars of the
lower jaw are very weak, and are often lost at an early age.[50] The teeth of brown
bears reflect their dietary plasticity and are broadly similar to those of other
bears.[51][52] They are reliably larger than teeth of American black bears, but
average smaller in molar length than those of polar bears.[53]

Claws and feet

Front paws
Brown bears have large, curved claws, with the front ones being larger than the
back. They may reach 5 to 6 cm (2.0 to 2.4 in) and measure 7 to 10 cm (2.8 to 3.9
in) along the curve.[50] Compared with the American black bear (Ursus americanus),
the brown bear has longer and stronger claws, with a blunt curve.[49] Due to their
claw structure, in addition to their excessive weight, adult brown bears are not
able to climb trees as well as black bears. In rare cases adult female brown bears
have been seen scaling trees.[54] The claws of a polar bear are quite different,
being notably shorter but broader with a strong curve and sharper point.[27][55]
The species has large paws; the rear feet measure 21 to 36 cm (8.3 to 14.2 in)
long, while the forefeet tend to measure 40% less.[56] Brown bears are the only
extant bears with a hump at the top of their shoulder, which is made entirely of
muscle. This feature developed presumably to impart more force in digging, which
helps during foraging and facilitates den construction prior to hibernation.[49]

Distribution and habitat

Main article: Distribution of brown bears
A bear standing in flowing water
Brown bear at Brooks Falls
Brown bears inhabit the broadest range of habitats of any living bear species.[56]
They seem to have no altitudinal preferences and have been recorded from sea level
to an elevation of 5,000 m (16,000 ft) in the Himalayas.[56] In most of their
range, brown bears seem to prefer semi-open 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.[56]

This species was once native to Europe, much of Asia, the Atlas Mountains of
Africa, and North America,[57] 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.[58] The largest populations are in Russia with
130,000,[59] the United States with 32,500, and Canada with around 25,000. Brown
bears live in Alaska, east through the Yukon and Northwest Territories, south
through British Columbia, and through the western half of Alberta. The Alaskan
population is estimated at a healthy 30,000 individuals.[60] In the lower 48
states, they are repopulating slowly, but steadily along the Rockies and the
western Great Plains.[61]

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,[62] 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[63]) in the west, to Russia in the east, and from Sweden and Finland in the
north to Romania (5,000–6,000), Bulgaria (900–1,200), Slovakia (with about 600–800
animals), Slovenia (500–700 animals), and Greece (with Karamanlidis et al. 2015
estimating >450 animals) in the south.[64][65] In Asia, brown bears are found
primarily throughout Russia, thence more spottily southwest to parts of the Middle
East, including the Eastern Black Sea Region, Turkey which has 5,432[66]
individuals of brown bear, to as far south as southwestern Iran, and to the
southeast in Northeast China. Brown bears are also found in Western China,
Kyrgyzstan, North Korea, Pakistan, Afghanistan, and India. A population of brown
bears can be found on the Japanese island of Hokkaidō, which holds the largest
number of non-Russian brown bears in eastern Asia, with about 2,000–3,000 animals.
[56]

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, the brown bear and the American black bear are
the only bear species not classified as threatened by the IUCN.[1][20] 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.[67] The Syrian brown bear (U. a. syriacus) is
very rare and it has been extirpated from more than half of its historic range.[68]
One of the smallest-bodied subspecies, the Himalayan brown bear (U. a.
isabellinus), is critically endangered: it occupies only 2% of its former range and
is threatened by uncontrolled poaching for its body parts.[69] The Marsican brown
bear in central Italy is believed to have a population of just 50 bears.[70]

The smallest populations are most vulnerable to habitat loss and fragmentation,
whereas the largest are primarily threatened by overhunting. The use of land for
agriculture may negatively effect brown bears. Additionally, roads and railway
tracks could pose a serious threat, as oncoming vehicles may collide with crossing
animals.[1] Poaching has been cited as another mortality factor.[71] In one
instance, a 3-year-long survey in the Russian Far East detected the illegal
shipping of brown bear gallbladders to Southeast Asian countries. The purpose and
motive behind the trade is unknown.[72]

An action plan in 2000 aimed to conserve brown bears in Europe by mitigating human–
wildlife conflict, educating farm owners as to sustainable practices, and
preserving and expanding remaining forests.[73] Compensation was given to people
who suffered losses of livestock, food supplies, or shelter.[74] Growing bear
populations were recorded in some countries, such as Sweden, where an increase of
1.5% per annum occurred between the 1940s and 1990s.[75] Brown bears in Central
Asia are primarily threatened by climate change. In response to this,
conservationists plan on building wildlife corridors to promote easy access from
one brown bear population to another.[76] In Himalayan Nepal, farmers may kill
brown bears in revenge for livestock predation.[77]

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
A 2014 study revealed that brown bears peaked in activity around the morning and
early evening hours.[78] Although activity can happen day or night, bears that live
in locations where they are apt to interact with humans are more likely to be fully
nocturnal.[79] In areas with little interaction, many adult bears are primarily
crepuscular, while yearlings and newly independent bears appear to be most active
throughout the day.[80][81] From summer through autumn, a brown bear can double its
weight from what it was in the spring, gaining up to 180 kg (400 lb) of fat, on
which it relies to make it through winter, when it becomes lethargic.[82][83]
Although they are not full hibernators and can be woken easily, both sexes prefer
to den in a protected spot during the winter months.[84] Hibernation dens may be
located at 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.[85]
Brown bears have one of the largest brains of any extant carnivoran relative to
their body size and have been shown to engage in tool use, which requires advanced
cognitive abilities.[86] This species is mostly solitary, although bears may gather
in large numbers at major food sources (e.g., open garbage dumps or rivers
containing spawning salmon) and form social hierarchies based on age and size.[87]
[88] Adult male bears are particularly aggressive and are avoided by adolescent and
subadult males, both at concentrated feeding opportunities and chance encounters.
Females 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.[89][90]
Dominance between bears is asserted by making a frontal orientation, showing off
canine teeth, 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.[87] During combat, bears use their paws to strike their
opponents in the chest or shoulders and bite the head or neck.[91]

Communication

Brown bear sounds

Duration: 1 minute and 32 seconds.1:32
Brown bear sounds including huffing, jaw popping and growling
Brown bear roars
Duration: 55 seconds.0:55
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 during everyday activities, a face where the ears pointed to
the sides and the mouth closed or slackly open. During social play, bears make
"relaxed open-mouth face" in which the mouth is open, with a curled upper lip and
hanging lower lip, and the ears alert and shifting. When looking at another animal
at a distance, the bear makes an "alert face" as the ears are cocked and alert, the
eyes wide open with 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 that 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 when the bear is aggressive and can
quickly switch between them.[91]

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 (approximately twice per
second). Growls and roars are made when aggressive. Growling is "harsh" and
"guttural" and can range from a simple grrr to a rumble. A rumbling growl can
escalate to 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!.[91]

Home ranges
Brown bears usually inhabit vast home ranges; however, they are not highly
territorial. Several adult bears roam freely over the same vicinity without
contention, unless rights to a fertile female or food sources are being contested.
[33][90] Despite their lack of traditional territorial behavior, adult males seem
to have a "personal zone" within which other bears are not tolerated if they are
seen.[92] Males always wander further than females, due to such behavior giving
increasing access to both females and food sources. Females have the advantage of
inhabiting smaller territories, which decreases the likelihood of encounters with
male bears who may endanger their cubs.[33][93]

In areas where food is abundant, such as coastal Alaska, home ranges for females
and males are up to 24 km2 (9.3 sq mi) and 89 km2 (34 sq mi), respectively.
Similarly, in British Columbia, bears of the two sexes travel in relatively compact
home ranges of 115 and 318 km2 (44 and 123 sq mi). In Yellowstone National Park,
home ranges for females are up to 281 km2 (108 sq mi) and up to 874 km2 (337 sq mi)
for males. In Romania, the largest home range was recorded for adult males (3,143
km2; 1,214 sq mi).[94] In the central Arctic of Canada, where food sources are
quite scarce, home ranges range up to 2,434 km2 (940 sq mi) for females and 8,171
km2 (3,155 sq mi) for males.[95][90]

Reproduction

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

The mating season occurs from mid-May to early July, shifting to later in the year
the farther north the bears are found.[96] Being serially monogamous, brown bears
remain with the same mate for a couple of days to a couple of weeks. Outside of
this narrow time frame, adult male and female brown bears show no sexual interest
in each other.[33] Females mature sexually between the ages of four and eight.[97]
Males first mate about a year later, when they are large and strong enough to
compete with other males for mating rights.[98] Males will try to mate with as many
females as they can; usually a successful male will mate with two females in a span
of one to three weeks.[42][98] The adult female brown bear is similarly
promiscuous, mating with up to four, sometimes even eight, males while in heat and
potentially mating with two in a single day.[99] Females come into oestrus every
three to four years, with an outside range of 2.4 to 5.7 years. The urine markings
of a female in oestrus can attract several males via scent.[100] 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.[44][93] Copulation is
prolonged and lasts for over 20 minutes.[44][101]

A mother bear standing next to her cubs

Grizzly bear cubs often imitate their mothers closely.
Males take no part in raising cubs – parenting is left entirely to the females.[90]
[102] 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.[97][103][104] Litters consist of as many as six cubs, though litters of one
to three are more typical.[105] The size of a litter depends on factors such as
geographic location and food supply.[106] At birth, cubs are blind, toothless and
hairless and may weigh 350 to 510 g (0.77 to 1.12 lb). 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 from a smaller one).[56][107][108]
Older and larger females within a population tend to give birth to larger litters.
[109] The cubs feed on their mother's milk until spring or 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 and forage for solid food with their mother over
long distances.[33][110]

Brown bear cub in Finland

The cubs are 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.
[95] Increased brain size in large carnivores has been positively linked to whether
a given species is solitary, as is the brown bear, or raises offspring communally.
Thus, the relatively large, well-developed brain of a female brown bear is
presumably key in teaching behavior.[111] The cubs learn by following and imitating
their mother's actions during the period they are with her.[56] Cubs remain with
their mother for an average of 2.5 years in North America, and gain independence
from as early as 1.5 years of age to as late as 4.5 years.[44] 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 in Hokkaido, and in Sweden most yearlings
were their own.[112][113] Brown bears practice infanticide, as an adult male bear
may kill the cubs of another.[93] 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.[93] Cubs may flee
up a tree when they see a strange male bear approaching. The mother often
successfully defends them, even though the male may be twice as heavy as she.
However, females have been known to die in such confrontations.[93][114][115]

Dietary habits
Main article: Dietary biology of the brown bear

Brown bear hunting salmon

The brown bear is one of the most omnivorous animals and has been recorded as
consuming the greatest variety of foods of any bear.[56] Despite their reputation,
most brown bears are not highly carnivorous, as they derive up to 90% of their
dietary food energy from vegetable matter.[116] They often feed on a variety of
plant life, including berries, grasses, flowers, acorns, and pine cones, as well as
fungi such as mushrooms.[33] Among all bears, brown bears are uniquely equipped to
dig for tough foods such as roots, bulbs,[117] and shoots. They use their long,
strong claws to dig out earth to reach roots and their powerful jaws to bite
through them.[33] In spring, winter-provided carrion, grasses, shoots, sedges,
moss,[117] and forbs are the dietary mainstays for brown bears internationally.[56]
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.[56]

Brown bear digging for squirrels

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.[118] 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.[49] This species may eat birds and their
eggs, including almost entirely ground- or rock-nesting species.[49] The diet may
be supplemented by rodents or similar small mammals, including marmots, ground
squirrels, mice, rats, lemmings, and voles.[33] With particular regularity, bears
in Denali National Park will wait at burrows of Arctic ground squirrels, hoping to
pick off a few of those 1 kg (2.2 lb) rodents.[119]

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 those 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 water, 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 salmon 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 closely in numbers at good spawning sites. The
largest and most powerful males claim the most fruitful fishing spots and will
sometimes fight over the rights to them.[49]

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.[49] While perhaps a majority of bears of the species will charge
at large prey at one point in their lives, many predation attempts start with the
bear clumsily and half-heartedly pursuing the prey and end with the prey escaping
alive.[49] On the other hand, some brown bears are quite self-assured predators who
habitually pursue and catch large prey. Such bears are usually taught how to hunt
by their mothers from an early age.[49] Large mammals preyed on can include various
ungulate species such as elk, moose, caribou, muskoxen, and wild boar.[33] When
brown bears attack these large animals, they usually target young or infirm ones,
which are easier to catch. Typically when hunting (especially young prey), the bear
pins its prey to the ground and then immediately tears at and eats it alive.[120]
It will also bite or swipe some prey to stun it enough to knock it over for
consumption.[121] To pick out young or infirm individuals, bears will charge at
herds so the more vulnerable, and thus slower-moving, individuals will become
apparent. Brown bears may ambush young animals by finding them via scent.[33]

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.[121] 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.[33] 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.[49] They feed on carrion, and use their size to intimidate
other predators – such as wolves, cougars, tigers, and American black bears – away
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.
[33] Cannibalism is not unheard of, though predation is not normally believed to be
the primary motivation when brown bears attack each other.[49]

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.[33] Plants and fruit
farmed by humans are readily consumed as well, including corn, wheat, sorghum,
melons, and any form of berries.[49] They may feed on domestic bee yards, readily
consuming both honey and the brood (grubs and pupae) of the honey bee colony.[49]
Human foods and trash are eaten when possible. When an open garbage dump was kept
in Yellowstone, brown bears were one of the most voracious and regular scavengers.
The dump was closed after both brown and American black bears came to associate
humans with food and lost their natural fear of them.[49]

Relations with other predators

Brown bear being followed by a wolf

Adult bears are generally immune to predatory attacks except from large Siberian
(Amur) tigers and other bears. Following a decrease of ungulate populations from
1944 to 1959, 32 cases of Siberian tigers attacking both Ussuri brown bears (Ursus
arctos lasiotus) and Ussuri black bears (U. thibetanus ussuricus) were recorded in
the Russian Far East, and bear hairs were found in several tiger scat samples.
Tigers attack black bears less often than brown bears, since the brown bears live
in more open habitats and are not able to climb trees. In the same time period,
four cases of brown bears killing female tigers and young cubs were reported, both
in disputes over prey and in self-defense.[50] In rare cases, when Amur tigers prey
on brown bears, they usually target young and sub-adult bears, besides small female
adults taken outside their dens, generally when lethargic from hibernation.[122]
Predation by tigers on denned brown bears was not detected during a study carried
out between 1993 and 2002.[123] Ussuri brown bears, along with the smaller black
bears constitute 2.1% of the Siberian tiger's annual diet, of which 1.4% are brown
bears.[124][125]

Brown bear and wolf pack squabbling over a carcass

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 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 at the same kill. To date, there is a single recorded
case of fully-grown wolves being killed by a grizzly bear.[126] Given the
opportunity, however, both species will prey on the other's cubs.[127] In some
areas, grizzly bears regularly displace cougars from their kills.[128] Cougars kill
small bear cubs on rare occasions, but there was only one report of a bear killing
a cougar, of unknown age and condition, between 1993 and 1996.[129][130]

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 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.[131] 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.[132] As of the 21st century, there has been an increase in
interactions between brown bears and polar bears, theorized to be caused by climate
change. Brown and grizzly bears have been seen moving increasingly northward into
territories formerly claimed by polar bears. They tend to dominate polar bears in
disputes over carcasses,[133] and dead polar bear cubs have been found in brown
bear dens.[134]

Longevity and mortality

Front paw imprint

Rear paw imprint

The brown bear has a naturally long life. Wild females have been observed
reproducing at 28 years, 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.[56][135] The lifespan of both sexes within minimally hunted populations
is estimated at an average of 25 years. The oldest recorded wild individual was
nearly 37 years old.[136] In captivity, the oldest recorded female was around 40
years old, while males have been known to live up to 47 years.[33][42]

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.[137] Annual mortality for bears of any age is
estimated at 10% in most protected areas.[56] Around 13% to 44% of cubs die within
their first year.[33] 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 cause of mortality
for first-year cubs is malnutrition.[33]

Brown bears are susceptible to parasites such as flukes, ticks, tapeworms,

roundworms, and biting lice.[138][139] It is thought that brown bears may catch
canine distemper virus (CDV) from other caniforms such as stray dogs and wolves.
[140] A captive individual allegedly succumbed to Aujeszky's disease.[141]

Hibernation physiology
A study conducted by the Brown Bear Research Project did a proteomic analysis of
the brown bear's blood, organs, and tissues to pinpoint proteins and peptides that
either increased or decreased in expression in the winter and summer months. One
major finding was that the presence of the plasma protein sex hormone-binding
globulin (SHBG) increased by 45 times during the brown bear's hibernation period.
Although scientists do not yet understand the role of SHBG in maintaining the brown
bear's health, some believe these findings could potentially help in understanding
and preventing human diseases that come from a sedentary life style.[142]

Relations with humans

Attacks on humans
Main article: Bear attack
See also: Bear danger
A statue of a roaring bear looking over a fence
A statue of the Ussuri brown bear from Hokkaido, Japan
Brown bears usually avoid areas where extensive development or urbanization has
occurred.[143] They usually avoid people and rarely attack on sight.[144] They are,
however, unpredictable in temperament, and may attack if threatened or surprised.
[145] Mothers defending cubs are the most prone to attacking, being responsible for
70% of brown bear-caused human fatalities in North America.[146] Attacks tend to
result in serious injury and, in some cases, death.[145] Due to the bears' enormous
physical strength, a single bite or swipe can be deadly.[147] Violent encounters
with brown bears usually last a few minutes, though they can be prolonged if the
victims fight back.[145]

A study conducted in 2019 found that 664 bear attacks were reported during a 15-
year period (2000–2015) throughout North America and Eurasia. There were 568
injuries and 95 fatalities.[148] Around 10 people a year are killed by brown bears
in Russia, more than all the other parts of the brown bear's range combined.[149]
In Japan, a large brown bear nicknamed Kesagake ("kesa-style slasher") caused the
worst brown bear attack in Japanese history in Tomamae, Hokkaidō, during numerous
encounters during December 1915. It killed seven people and wounded three others
before being gunned down during a large-scale beast-hunt.[150] 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.[151]

Bear hunting
Main article: Bear hunting

Hunters with a bear killed by a rifle (1904)

Humans have been recorded hunting brown bears for over 10,300–9,300 years.[152] In
Europe, between the 17th and 18th centuries, humans sought to control brown bear
numbers by awarding those who managed to kill one. This bounty scheme pushed the
brown bear population to the brink of extinction before comprehensive protection
was offered in the 1900s. Despite this, a 2018 study found hunting to be one of the
contributing factors to the drop in brown bear numbers in northern Europe.[153]
[148]

The earliest known case of a European killing a grizzly bear dates back to 1691.
Their arrival in western United States led to the extirpation of local brown bear
populations in the 19th and early 20th centuries. During the early years of
European settlement in North America, bears were usually killed with a spear or
lasso rope. The introduction of rifles in the mid-19th century largely facilitated
bear hunts, which allowed for an increasing trend. Bears were also pitted into
fights against male cattle, often ending with either animal grievously injured or
dead. The last two decades of the 19th century saw an increase in bounties.
Conflicts with farmers also contributed to its rapid decline. It wasn't until the
1920s that grizzly bears received some type of protection from the US government.
[154] Today, brown bears are legally hunted in some American states, such as
Alaska. However, a hunting license is required and killing females with cubs will
result in a prison sentence.[155]

Their meat is sometimes consumed and used in recipes such as dumplings, hams and
stews. The Indigenous people of James Bay (Eastern) Cree use their flesh in
traditional dishes. In Asia and Romania, the paws are consumed as exotic
delectables; brown bear paws have been a prevalent component of traditional Chinese
food since 500 BC. The total weight of commercially sold brown bear meat is
estimated at 17 tons annually.[156]

In captivity

Brown bear at Canton Zoo, US

Bears have been recorded in captivity as early as 1,500 BC.[157] As of 2017, there
are more than 700 brown bears in zoos and wildlife parks worldwide. Captive bears
are largely lethargic and spend a considerable amount of time doing nothing. When
active, captive bears may engage in repetitive back and forth motion, known widely
as pacing. This behavior is most prevalent in bears kept in small, cramped cages
often with no natural setting. Pacing is a way of coping with stress that comes
with being trapped in unnaturally small spaces.[158] These stereotyped behaviors
have decreased due to better and larger enclosures being built, and more
sustainable management from zoo staff.[159]

Starting from infancy, brown bears may also be exploited as dancing bears. Cubs,
for example, are positioned on hot metal plates, causing them to "dance" to the
sound of violin music running in the background. The process is repeated, resulting
in bears being trained to "dance" when a violin is played. Similarly, brown bears
are displayed in tiny enclosures near a restaurant, mainly for the purpose of
luring customers. Privately-owned bears are usually placed in insufficient
environments and often suffer from malnutrition and obesity.[160]

Animal trainer with a bear

Brown bears have been popular attractions at circuses and other acts since ancient
times. Due to their large size and imposing demeanor, the Romans used brown bears
in the execution of criminals, and pitted bears in fights with other animals.
Gladiators would fight bears, in what was essentially a fight to the death. Such
events occurred in amphitheaters housing thousands of spectators. In later times,
street performances became popular in Medieval times; acts included "dancing" and
"sleeping on command". These performances became increasingly widespread, and from
the 1700s to 1800s, traveling circuses would perform in the streets of many
European and Asian countries. Such circuses made use of bears that wore special
clothing. They were usually run by musicians, and still continue today. Modern
circuses began utilizing bears around the second half of the 18th century. Brown
bears were said to be the easiest bear species to train due to their intelligence,
unique personalities, and exceptional stability.[157] According to a 2009 analysis,
the brown bear was the second most exploited circus animal after the tiger.[161]

Culture
Further information: Cultural depictions of bears
The image shows three bears standing in a house
"The Story of the Three Bears", illustration from Childhood's Favorites and Fairy
Stories
The image shows a bear behind a house of cards
Russia is often represented by a bear, as in this Puck political cartoon from 1904.
Bears have been popular subjects in art, literature, folklore, and mythology. The
image of the mother bear was prevalent throughout societies in North America and
Eurasia, based on the female's devotion and protection of her cubs.[162] The
earliest cave paintings of bears occurred in the Paleolithic, with over 100
recorded paintings.[163] Brown bears often figure in the literature of Europe and
North America as "cute and cuddly", in particular that which is written for
children. "The Brown Bear of Norway" is a Scottish fairy tale telling of 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 after many
trials and tribulations. 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.[164] Smokey Bear, the famous mascot of U.S. Forest Service, has
since the 1940s been used to educate people on the dangers of human-caused
wildfire.[165]

Brown bears have been extensively featured in the culture of Native Americans, and
are considered sacred. To stop a bear's spirit from escaping after it was killed,
the Denaa people severed all 4 of its paws. They delayed consuming brown-bear
flesh, owing to the belief that the bear's spirit was overwhelming in fresh kills.
Additionally, community members that wore bear claw necklaces were highly
respected, as wearing one was seen as a sign of bravery and honor. The clattering
caused by repeatedly shaking these necklaces were believed to bring have
therapeutic powers. In Haida culture, one legend has it that a marriage between a
woman and a grizzly bear commenced the lineage of Indian people. This is thought to
have allowed the Haida to thrive in bear country.[154]

There is evidence of prehistoric bear worship, though this is disputed by

archaeologists.[166] It is possible that bear worship existed in early Chinese and
Ainu cultures.[167] The Romans built small carved figures of bears that were used
during the burials of infants.[168] In Ancient Greek mythology, bears were
considered similar to humans, mainly due to their ability to stand upright. In many
western stories and older fables the portrayed attributes of bears are
sluggishness, foolishness, and gullibility, which contradicts the actual behavior
of the species. For example, bears have been reported tricking hunters by
backtracking in the snow.[169]

In North America, the brown bear is considered a charismatic megafauna and has long
piqued people's interest. The death of Bear 148 at the hands of a trophy hunter in
2017 sparked media outrage and the continued disapproval of trophy hunting.[170]
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.[171][172] The grizzly bear is
the state animal of Montana.[173] The California golden bear is the state animal of
California, despite being extinct.[174] The coat of arms of Madrid depicts a bear
reaching up into a madroño or strawberry tree (Arbutus unedo) to eat some of its
fruit. The Swiss city of Bern's coat of arms depicts a bear and the city's name is
popularly thought to derive from the German word for bear.[175][176] The brown bear
is depicted on the reverse of the Croatian 5-kuna coin, minted since 1993.[177]