The brown bear (Ursus arctos) is a large bear species found across Eurasia and North America. 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. The brown bear's range includes parts of Russia, Central Asia, China, Canada, the United States, Hokkaido, Scandinavia, the Balkans, the Picos de Europa and the Carpathian region (especially Romania), Iran, Anatolia, and the Caucasus. The brown bear is recognized as a national and state animal in several European countries.
While the brown bear's range has shrunk, and it has faced local extinctions, 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. Populations that were hunted to extinction in the 19th and 20th centuries are the Atlas bear of North Africa and the Californian, Ungavan and Mexican populations of the grizzly bear of North America. Many of the populations in the southern parts of Eurasia are highly endangered as well. One of the smaller-bodied forms, the Himalayan brown bear, is critically endangered, occupying only 2% of its former range and threatened by uncontrolled poaching for its body parts. The Marsican brown bear of central Italy is one of several currently isolated populations of the Eurasian brown bear and is believed to have a population of just 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).[better source needed] In the mid-19th century United States, the brown bear was termed "Old Ephraim" and sometimes as "Moccasin Joe".
Generalized names and evolution
Brown bears are thought to have evolved from Ursus etruscus in Asia. 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." 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. 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. 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. It is speculated that brown bears were unable to migrate south until the extinction of the much larger giant short-faced bear (Arctodus simus).
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. 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.
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. 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. However, recent DNA analysis has identified as few as five main clades which contain all extant brown bears, while a 2017 phylogenetic study revealed nine clades, including one representing polar bears. As of 2005, 15 extant or recently extinct subspecies were recognized by the general scientific community.
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. Under some definitions, the brown bear can be construed as the paraspecies for the polar bear.
DNA analysis shows that, apart from recent human-caused population fragmentation, 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. 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, 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.
Brown bears are apparently divided into five different clades, some of which coexist or co-occur in different regions.
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. 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, 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.
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.
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. 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. 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).
Brown bears are often not fully brown. They have long, thick fur, with a moderately long mane at the back of the neck which varies somewhat across the types. 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. 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. 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.
Cranial morphology and size
Adults have massive, heavily built concave skulls, which are large in proportion to the body. The forehead is high and rises steeply. 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. Grizzlies, for example, tend to have flatter profiles than European and coastal American brown bears. 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. 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. 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. The teeth are reliably larger than American black bears, but average smaller in molar length than polar bears. 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. 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.
Claws and feet
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. They are generally dark with a light tip, with some forms having completely light claws. Brown bear claws are longer and straighter than those of American black bears (Ursus americanus). 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. 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. 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. 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.
Distribution and habitat
Brown bears were once native to Europe, much of Asia, the Atlas Mountains of Africa, and North America, 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. The largest populations are in Russia with 120,000, 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. In the lower 48 states, they are repopulating slowly, but steadily along the Rockies and the western Great Plains.
In Europe, there are 14,000 brown bears in ten fragmented populations, from Spain (estimated at only 20–25 animals in the Pyrenees in 2010, 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) in the west, to Russia in the east, and from Sweden and Finland in the north to Romania (4000–5000), Bulgaria (900–1200), Slovakia (with about 600–800 animals), Slovenia (500–700 animals) and Greece (with about 200 animals) in the south.
In Asia, brown bears are found primarily throughout Russia, thence more spottily southwest to parts of the Middle East, to as far south as southwestern Iran, and to the southeast in a small area of Northeast China, Western China, and parts of North Korea, Pakistan, Afghanistan and India. 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.
This species inhabits the broadest range of habitats of any living bear species. 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). 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.
While the brown bear's range has shrunk and it has faced local extinctions, it remains listed as a Least concern species by the IUCN, with a total population of approximately 200,000. As of 2012, this and the American black bear are the only bear species not classified as threatened by the IUCN. However, the California grizzly 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. The Syrian brown bear (Ursus arctos syriacus) is very rare and it has been extirpated from more than half of its historic range. 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. The Marsican brown bear in central Italy is believed to have a population of just 50–60 bears.
Behavior and life history
The brown bear is often described as nocturnal. However, it frequently seems to peak in activity in the morning and early evening hours. 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. 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. 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.
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. 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. 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. 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. During combat, bears use their paws to strike their opponents in the chest or shoulders and bite the head or neck.
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" expert 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.
Brown bears also produce various vocalizations. Huffing occurs when the animal is tense while woofing is made when it is startled. Both sounds are produce by exhalations though huffing is harsher and is made repeatedly (two per second) while a woof is made only once. Growls and roars are made in aggression. Growling is harsh and gutteral and can range from short and low grrr to a continuous rolling rumble. Roaring is described as "thunderous" and can be heard 2 km (1.2 mi) away. A rumbling growl can escalate into a roar when the bear is charging. Bears will bawl when seeking contact, such as between mothers and cubs or a male to a potential mate. This sound is heard as waugh!, waugh!.
Brown bears usually occur over vast home ranges; however, they are not highly territorial. Several adult bears often roam freely over the same vicinity without issue, unless rights to a fertile female or food sources are being contested. Males always cover more area than females each year. 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. 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. 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). 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.
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. 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.
The mating season is from mid-May to early July, shifting later the further north the bears are found. Being serially monogamous, brown bears remain with the same mate from 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. 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. 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. 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. Paternity DNA tests have shown that littermates do not share the same father in up to 29% of litters.  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. Copulation is vigorous and prolonged and can last up to an hour, although the mean time is about 23–24 minutes.
Males take no part in raising their cubs – parenting is left entirely to the females. Through the process of delayed implantation, a female's fertilized egg divides and floats freely in the uterus for six months. During winter dormancy, the fetus attaches to the uterine wall. The cubs are born eight weeks later while the mother sleeps. If the mother does not gain enough weight to survive through the winter while gestating, the embryo does not implant and is reabsorbed into the body. 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. 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). Older and larger females within a population tend to give birth to larger litters The size of a litter also depends on factors such as geographic location and food supply. At birth, the cubs are blind, toothless and hairless and may weigh from 350 to 510 g (0.77 to 1.12 lb), again reportedly based on the age and condition of the mother. They feed on their mother's milk until spring or even early summer, depending on climate conditions. At this time, the cubs weigh 7 to 9 kg (15 to 20 lb) and have developed enough to follow her over long distances and begin to forage for solid food.
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. 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. The cubs learn by following and imitating their mother's actions during the period they are with her. 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. 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. Brown bears practice infanticide, as an adult male bear may kill the cubs of a female bear. 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. 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 lose their lives in these confrontations.
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. 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. They often feed on a variety of plant life, including berries, grasses, flowers, acorns and pine cones, as well as fungi such as mushrooms. Among all bears, brown bears are uniquely equipped to dig for tough foods such as roots and shoots. They use their long, strong claws to dig out earth to reach the roots and their powerful jaws to bite through them. In spring, winter-provided carrion, grasses, shoots, sedges and forbs are the dietary mainstays for brown bears internationally. 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. 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. 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. This species may eat birds and their eggs, including almost entirely ground- or rock-nesting species. The diet may be supplemented by rodents or similar smallish mammals, including marmots, ground squirrels, mice, rats, lemmings and voles. 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.
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.
Beyond the regular predation of salmon, most brown bears are not particularly active predators. 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. 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. Large mammals preyed on can include various ungulate species such as elk, moose, caribou, muskoxen and wild boar. 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. It will also bite or swipe some prey to stun it enough to knock it over for consumption. 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. 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. 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. 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. 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 comprised by winter-killed big game. Cannibalism is not unheard of, though predation is not normally believed to be the primary motivation when brown bears attack each other.
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. Plants and fruit farmed by humans are readily consumed as well, including corn, wheat, sorghum, melons and any form of berries. They will also feed at domestic bee farms, readily consuming both honey and the contents of the honey bee colony. 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.
Interspecific predatory relationships
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. 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. Some bears emerging from hibernation seek out tigers in order to steal their kills. 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. Geptner et al. (1972) stated bears are generally afraid of tigers and change their path after coming across tiger trails. 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. Other researchers have observed bears following tiger tracks for various reasons.
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. Given the opportunity, however, both species will prey on the other's cubs. 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. 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. 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. There is one record of a golden eagle predating on a brown bear cub.
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. 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. They will eat the fruit dropped by the Asian black bear from trees, as they themselves are too large and cumbersome to climb. Improbably, in the Himalayas Brown bears are reportedly intimidated by Asian black bears in confrontations.
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, and dead polar bear cubs have been found in brown bear dens.
Longevity and mortality
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. The lifespan of brown bears of both sexes within minimally hunted populations is estimated at an average of 25 years. The oldest wild brown bear on record was nearly 37 years old. 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.
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. Annual mortality for bears of any age is estimated at around 10% in most protected areas; however, the average annual mortality rate rises to an estimated 38% in hunted populations. Around 13% to 44% of cubs die within their first year even in well-protected areas. Mortality rates of 75–100% among the cubs of any given year are not uncommon. 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. 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%.
Even in populations living in protected areas, humans are still the leading cause of mortality for brown bears. The largest amount of legalized brown bear hunting occurs in Canada, Finland, Russia, Slovakia and Alaska. 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. Brown bears are also killed in collisions with automobiles, which is a significant cause of mortality in the United States and Europe.
Relationship with humans
Conflicts between bears and humans
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. Under many circumstances, extensive human development may cause brown bears to alter their home ranges. High road densities (both paved and gravel roads) are often associated with higher mortality, habitat avoidance and lower bear density. 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. In other areas, such as Alaska, dumps may continue to be an attractant for brown bears. In different parts of their distribution, brown bears sometimes kill and eat domesticated animals. 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. 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.
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." 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.
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. 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.
Bear encounters and attacks
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. They are, however, unpredictable in temperament, and may attack if they are surprised or feel threatened. There are an average of two fatal attacks by bears per year in North America. 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. 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. 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. Mothers defending cubs are the most prone to attacking, being responsible for 70% of brown bear-caused human fatalities in North America.
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. In the majority of attacks resulting in injury, brown bears precede the attack with a growl or huffing sound. 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. 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. 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. 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. 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.
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. 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. 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. 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. In Scandinavia, only three fatal attacks were recorded in the 20th century.
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.
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. 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.
History of defense from bears
A study by U.S. and Canadian researchers has found pepper spray to be more effective at stopping aggressive bear behavior than guns, working in 92% of studied incidents versus 67% for guns. 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. 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. 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)".
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.
Brown bears often figure into the literature of Europe and North America, in particular that which is written for children. "The Brown Bear of Norway" is a Scottish fairy tale telling the adventures of a girl who married a prince magically turned into a bear and who managed to get him back into a human form by the force of her love and after many trials and difficulties. With "Goldilocks and the Three Bears", a story from England, the Three Bears are usually depicted as brown bears. In German-speaking countries, children are often told the fairytale of "Snow White and Rose Red"; the handsome prince in this tale has been transfigured into a brown bear. In the United States, parents often read their preschool age children the book Brown Bear, Brown Bear, What Do You See? to teach them their colors and how they are associated with different animals.
The Russian bear is a common national personification for Russia (as well as the former Soviet Union), despite the country having no appointed national animal. The brown bear is Finland's national animal.
The grizzly bear is the state animal of Montana. The California golden bear is the state animal of California. 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).
- McLellan, B.N.; Proctor, M.F.; Huber, D. & Michel, S. (2017). "Ursus arctos". The IUCN Red List of Threatened Species. IUCN. 2017: e.T41688A121229971. doi:10.2305/IUCN.UK.2017-3.RLTS.T41688A121229971.en.
- Servheen, C.; Herrero, S.; Peyton, B.; Pelletier, K.; Moll, K.; Moll, J., eds. (1999), Bears: status survey and conservation action plan (PDF), 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.
- Christiansen, P. (1999). "What size were Arctodus simus and Ursus spelaeus (Carnivora: Ursidae)?". Annales Zoologici Fennici. 36 (2): 93–102. JSTOR 23735739.
- 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.
- Zedrosser, A.; Dahle, B.; Swenson, J. E.; Gerstl, N. (2001). "Status and management of the brown bear in Europe" (PDF). Ursus. 12: 9–20.
- 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.
- "Mammalian Species- Ursus arctos" (PDF). American Society of Mammalogists, Smith College. 23 April 1993. Archived from the original (PDF) on 31 March 2017.
- Himalayan brown bears now critically endangered. 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.
- 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.
- 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.
- 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.
- Kurten, Bjorn (1976). The Cave Bear Story. New York: Columbia University Press.
- 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.
- 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.
- 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". Molecular Ecology. 17 (8): 1962–1970. doi:10.1111/j.1365-294x.2008.03631.x. PMID 18363668. S2CID 23361337.
- 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.; 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.
- 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.
- Kurtén, B. (1964). "The evolution of the polar bear, Ursus maritimus (Phipps)". Acta Zoologica Fennica. 108: 1–26.
- 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.
- 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.
- 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).
- Nowak, R. M. (1999). Walker's mammals of the world (Vol. 1). JHU Press.
- Fitzgerald, Christopher S.; Krausman, Paul R. (2002). "Helarctos malayanus" (PDF). Mammalian Species. 696: 1–5. doi:10.1644/1545-1410(2002)696<0001:HM>2.0.CO;2. Archived from the original (PDF) on 4 March 2016. Retrieved 12 December 2016.
- "Ursus arctos – Brown Bear species". Wildpro. Archived from the original on 17 May 2012. Retrieved 14 June 2012.
- 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.; 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.
- 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.
- 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 & Justice. 48 (#1): 8–15. doi:10.1016/j.scijus.2007.05.001. PMID 18450212.
- Bear Anatomy and Physiology from Gary Brown's The Great Bear Almanac, Lyons & Burford, Publishers, 1993
- 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)]. II, Part 1a. Washington, D.C., USA: Science Publishers, Inc. ISBN 1-886106-81-9.
- ''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.
- Kurtén, 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.
- 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., & Guravich, D. (1998). Polar bears. University of Michigan Press.
- 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., & Elman, R. (1996). National Audubon Society Field Guide to North American Mammals (p. 992). New York: Knopf.
- Novak, M., Baker, J.A., Obbard, M.E. & Malloch, B. (1987). Wild furbearer management and conservation in North America. Ontario Ministry of Natural Resources.
- "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'ós bru als Pirineus". elperiodico.cat (in Catalan and Spanish). 2 August 2010. Retrieved 19 November 2010.
- "Situació preocupant de l'ós bru als Pirineus" (in Catalan). ecologistasenaccion.org. May 2010. Retrieved 19 November 2010.
- "Osos, el desafío de una población creciente". ABC (in Spanish). Spain.
- "Brown Bear – Population & Distribution: A Truly International Species". wwf.panda.org. Retrieved 19 November 2010.
- 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.
- "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.
- 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-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.
- 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.; 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.
- 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.
- 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.
- 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
- Egbert, A. L. (1978). The Social Behavior of Brown Bears at McNeil River, Alaska (Ph.D. thesis). Utah State University.
- Pearson, A. M. (1975). The northern interior grizzly bear Ursus arctos L. Information Canada.
- 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.
- 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)
- 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.
- 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 für Säugetierkunde. 75: 29. doi:10.1016/j.mambio.2009.03.010.
- 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.
- 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 & Co. Ltd., London, UK.
- 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.
- 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.
- Frković, A.; Huber, D. & Kusak, J. (2001). "Brown bear litter sizes in Croatia". Ursus. 12: 103–105. JSTOR 3873235.
- Pazetnov, V.A. & Pazetnov, S.V. (2005). "Female brown bear with six cubs" (PDF). International Bear News. 14 (#2): 17.
- 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.
- 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.
- 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<1026:RCOBBO>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<0536:FBIBBA>2.0.CO;2.
- Bellemain, E.; Zedrosser, A.; Manel, S.; Waits, L. P.; Taberlet, P.; Swenson, J. E. (2006). "The dilemma of female mate selection in the brown bear, a species with sexually selected infanticide". Proceedings of the Royal Society of London B: Biological Sciences. 273 (#1, 584): 283–291. doi:10.1098/rspb.2005.3331. PMC 1560043. PMID 16543170.
- 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.
- 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.
- "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.
- "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. "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.
- 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 & Management. International Association for Bear Research and Management. p. 64.
- 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). 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.
- 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.
- 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.
- 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.
- 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. 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.
- 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.
- 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.
- 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.
- 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.
- McDonald, D., & 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.
- 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.
- "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., & 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". . 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.
- 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.
- Larivière, S. (2001). "Ursus americanus" (PDF). Mammalian Species. 647: 1–11. doi:10.1644/1545-1410(2001)647<0001:ua>2.0.co;2. 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.
- 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.
- 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.
- 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.
- 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.
- 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<0010:gbcitg>2.0.co;2.
- 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.; 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.
- Mertens, A.; Promberger, C. (2001). "Economic aspects of large carnivore-livestock conflicts in Romania". Ursus. 12: 173–180. JSTOR 3873246.
- 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.
- 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.
- 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.
- Herrero, Stepehen (1985). Bear Attacks: Their Causes and Avoidance. Edmonton: Hurtig Publishers Ltd.
- 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.
- 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 & 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 kansalliseläin". yle.fi.
- "Symbols of Montana". Montana Historical Society.
- "History and Culture – State Symbols". California State Library.
- 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.