The brown bear (Ursus arctos) is a large bear species found across Eurasia and North America.

[1][4] 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.[5][6][7][8][9] 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][10] The brown bear is recognized as a national and state
animal in several European countries.[11]

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][4][8] Populations that were hunted to
extinction in the 19th and 20th centuries are the Atlas bear of North Africa and the Californian,
Ungavan[12][13] 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][14] 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.[15] 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.

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).[17][better  source  needed] 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. [14]

Scientific taxonomy
Main article: Subspecies of brown bear

Eurasian brown bear (Ursus arctos arctos), 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, 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.[4]

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.[7] 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).[14][55][56][57]

Color

Brown bear on a rock

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. [14][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]