Ursus arctos (Brown bear)

Brown bears (Ursus arctos) are powerful, adaptable mammals known for their muscular shoulder hump, long front claws, and a coat that ranges from black to blonde. They have the widest distribution of all bear species, and their size and weight vary greatly depending on habitat and food resources.

Table of Contents

Fast Facts

Body length: 100 to 190 cm

Average weight: 92 to 357 kg (203 to 787 lbs), some individuals up to 700 kg (1,543 lb)

Lifespan: Oldest recorded female in the wild is 37 years

Distribution: Northern Eurasia, India, and North America

Status: Least Concern, but many small isolated populations are in danger of local extirpation while other populations are expanding (IUCN). Appendix I (China and Mongolia), Appendix II (all other populations, CITES)

Taxonomy

Kingdom: Animalia

Phylum: Chordata

Class: Mammalia

Order: Carnivora

Family: Ursidae

Subfamily: Ursinae

Genus: Ursus

Species: Ursus arctos Linnaeus, 1758

Common names

Albanian: Ariu i murrmë

Arabic: لدب البني

Armenian: գորշ արջ

Azerbaijan: qonur ayi

Belarusian: буры мядзведзь

Bulgarian: кафява мечка

Catalan: ós cafè

Chinese: 马熊

Croatian: medvjed

Czech: medvěd hnědý

English: Brown bear, Grizzly bear

Estonian: pruunkaru

French: Ours brun

Georgian: მურა დათვი

German: Braunbär

Greek: καφέ αρκούδα

Hindi: भूरे भालू

Italian: orso bruno

Japanese: ヒグマ

Korean: 갈색 곰

Latvian: brūnais lācis

Macedonian: кафеавата мечка

Mongolian: хүрэн баавгай

Nepali: खैरो भालु

Norwegian: brunbjørn

Persian: خرس قهوه ای

Polish: niedźwiedź brunatny

Romanian: ursul brun

Russian: бурый медведь

Serbian: мрки медвед

Slovak: medveď hnedý

Slovenian: rjavi medved

Spanish: Oso Café

Swedish: brunbjörn

Turkish: bozayı

Ukranian: бурий ведмідь

Urdu: بھورا ریچھ

Subspecies

The exact number of brown bear subspecies is under debate. A large number of subspecies were previously identified using morphometrics (differences in physical traits), but more recent investigations using genetic analyses and other techniques such as microsatellite markers conclude that there are fewer subspecies than previously identified. In North America, two current distinct subspecies have been identified (1) the mainland grizzly bear (U. a. horribilis), and (2) the Kodiak Island brown bear or Kodiak bear (U. a. middendorffi), [1]. However, in all of Eurasia, only two distinct groups with different ancestors have been identified. The number and distribution of subspecies is still debated, with all brown bears in northwesten Eurasia being attributed to the (U. a. arctos) subspecies [2]. Although still tentative, there is new evidence that the small, isolated brown bear population dwelling in the Gobi Desert of Mongolia is a separate subspecies.

Description

The brown bear is the second-most numerous of the bear species after the American black bear (Ursus americanus). Following polar bears (U. maritimus), brown bears are the second-largest bear species, but large males along the coast of mainland Alaska and Kodiak Island, Alaska, USA, can sometimes rival the size of polar bears. Male brown bears are generally ≥20% larger than females, but sometimes even twice as large (a trait referred to as sexual dimorphism; [3, 4]). Adult body length can range from 1 m to 3 m but is usually between 1.8 m and 2.2 m [5]. Body size for brown bears varies with latitude, population density, competition for food resources, availability and quality of foods, and environmental conditions that affect food resources [6]. Adult males commonly weigh approximately 200 kg (440 lbs), but the average weight in different populations can be as low as 90 kg (200 lbs; Jasper National Park, Alberta, Canada) or as high as 360 kg (794 lbs; Coastal Alaska, USA). Some coastal males with access to salmon (Oncorhynchus spp.) can exceed 770 kg (1,700 lbs), [4, 5, 7]. Female average weight can be as little as 55 kg (120 lbs; Jasper National Park) but is generally approximately 135 kg (300 lbs), [4, 7]. Brown bears also go through large annual fluctuations in body mass because they store fat during summer and autumn to prepare for winter hibernation [6].

Coat color can range from black to blond but is usually brown. Individuals of the grizzly bear subspecies (U. arctos horribilis) found in North America are often observed with a silver-tipped brown coat that gives them a “grizzled” appearance. Brown bears have a prominent hump of muscle between their shoulders, a disk-shaped face, and long front claws; these traits serve as trademarks in the identification of brown bears. Their front claws can reach up to 8 cm in length [4], and are adapted to digging into hard soil for rodents and roots, but not for climbing trees. Although brown bears are commonly found in forested areas, they evolved in and often prefer open habitats such as alpine areas, northern plains, and meadows, and are more adapted to non-forest environments [8]. Brown bears have powerful jaws and strong teeth with long, curved canines but, unlike wolves and large felids, they are unable to break open large bones (dental formula: I 3/3, C 1/1, P 4/4, M 2/3 = 42 [4, 7]).

Morphology

Total lengtha,bTailFront PadRear padBody mass
Adult ♂164.3a, 186b, 191.57d, 145e17b7.7a18.9a192a, 140-320b, 92c, 191.43d, 201-273e, 357f
Adult ♀151.1a, 160b, 136.25d16b6.7a16.3a135a, 100-200b, 55c, 170.5d, 96-158e, 226f, 98g

Length measurements in cm and mass in kg. a [9] Yellowstone National Park (USA), b [7] Eastern Siberia, c [7] Jasper National Park (Canada), d [10] Turkey, e [11] Sweden, Norway, Slovenia, and Croatia, f [4] Coastal Alaska (USA), g [4] Yukon (Canada).

Biology

Brown bears are generally most active during daylight hours or crepuscular periods (dawn and dusk), but can be mostly diurnal or nocturnal depending on the region, climate, and the level of human activity where they live [12]. Brown bears are mostly solitary except in mother–offspring groups or when mating, but are known to tolerate the presence of several other bears when foraging from highly abundant food sources. Home-range overlap is common for related females, and individuals of both sexes can spend a considerable amount of time interacting with each other [13, 14, 15]. During salmon spawning events in Canada, the United States, and the eastern coast of Russia (Kamchatka), it is possible to observe >30 bears feeding on salmon in close proximity to each other [16]. In Glacier and Yellowstone National Parks, USA, aggregations of bears have also been observed feeding on army cutworm moths (Euxoa auxiliaris) in alpine sites [4, 17, 18]. Brown bears are terrestrial omnivores and have a highly adaptable diet composed mainly of plants, roots, fruits, nuts, insects, salmon where available, and small and large live prey. They can also prey on livestock, which is often a source of conflict with people. Their diet follows the growing season and availability of key foods. The availability and abundance of high-quality foods dictates body size and reproductive rates [6, 8, 19, 20], and individuals in populations with access to high-quality foods such as salmon are larger than interior populations with lower habitat productivity.

Mating occurs in the spring and continues until mid-summer (May to early Jul), and both males and females may breed with multiple partners. The embryo does not implant in the uterus until autumn, a strategy referred to as delayed implantation. Following this strategy, the embryo will not usually implant if the female did not accumulate ≥20% of her body weight as fat reserves for that year [21]. Gestation lasts approximately 6–8 weeks [4, 22]. Cubs are quite vulnerable at birth, they are born in the winter den and weigh only approximately 450 g (1 lb). Litter sizes typically vary between 1 and 3 cubs, with 2 cubs most commonly observed in healthy populations. However, in a few instances, females have been seen with 4 cubs [4]. Females can start reproducing between ages 3 and 12, depending on the region. They most commonly start at 5–7 years old, and rarely reproduce past 28 years of age [7, 20, 23, 24]. Males generally reach sexual maturity at approximately 5 years old [4]. Maternal care usually lasts 1–3 years. Infanticide (large males killing cubs) has been reported in several populations, and is thought to be used as a tactic for males to mate with females sooner because females that lose their cubs can go into estrus sooner than females with cubs [25]. Brown bears have one of the lowest reproductive rates of any mammals [3], and reproductive output varies with differences in latitude, habitat productivity, population health, population density, and management practices and regulations [20, 26]. Adult brown bears do not have any natural predators other than humans, although in countries where tiger (Panthera tigris) range overlaps with brown bear range, there have been reports of tiger-killed adult brown bears [27, 28]. Other than infanticide, the causes of brown bear cub mortalities are largely unknown [4] but wolves (Canis lupus) and golden eagles (Aquila chrysaetos) have been known to kill brown bear cubs opportunistically [29, 30]. The oldest brown bear observed in the wild was 37 years old [31].

When in close proximity, brown bears maintain a hierarchical social-dominance structure and communicate their dominance by using behaviors including body posture and orientation; head, neck, and mouth postures; and vocalization [32]. Although large males are most dominant, females with cubs can rival dominant males aggressively [16, 32]. At a distance, brown bears also communicate with each other by marking trees with their teeth and claws, and by rubbing and leaving their scent on the trunk, especially during the mating season [33].

Although rare, attacks on humans do occur [34]. Brown bears that feel threatened, especially females with cubs, can act aggressively to defend themselves against a perceived threat [34]. A brown bear that is acting defensively may first swat at the ground, pop its jaw, blow and snort, or even bluff-charge (lunge at the perceive threat) [34]. Human–bear interactions that result in physical contact most commonly involve a brown bear being surprised at close range [34]. Brown bears usually avoid people [35, 36], and the most effective way to avoid an encounter is to travel in groups and make noise [34]. Even though firearms are the only lethal deterrent that can stop an aggressive bear, bear spray (red pepper spray) is the most effective deterrent on brown bears during close encounters and firearms generally fail to protect people [36, 37, 38].

Reproduction and Lifespan
Gestation (weeks)Maximum litter sizeJuvenile lifespan (years)Adult lifespan (years)Maximum age (years)
6-8b,c4 (rare)b,d,e,f,g0-36h, 16-25i37j

Lifespan varies widely. b [4] Coastal Alaska (USA), c [31] Alberta (Canada), d [39] Croatia, e [40] France, f [41] Sweden and Norway, g [42] China, h [7] Yellowstone National Park (USA), i [43] Slovenia, j [31] Continental United States.

Breeding interval: Can breed once per year, but the interval between cub production is generally ≥3 years.

Breeding season: Spring to mid-summer (May to early July)

Independence: Most often 3 years (1.5–4.5 yr)

Denning and Hibernation

Hibernation has evolved as a strategy to avoid harsh environmental conditions and a lack of resources usually associated with winter [44]. Hibernation in bears is unique because bears do not undergo deep torpor bouts typical of rodent hibernators such as squirrels (Spermophilus sp.) and marmots (Marmota sp.), and they can easily become active if disturbed while hibernating [45, 46]. During hibernation, brown bears lower their metabolism and heart rate without causing adverse effects on cardiovascular function; this exceptional feat, when better understood, could have important benefits in the treatment of heart disease in humans [46].

Brown bears typically enter hibernation in the autumn (Oct–Nov) and emerge from winter dens in the spring (usually approx. Apr) [47, 48, 49, 50, 51]. In most cases, gestating females enter dens first and hibernate the longest, followed by females with cubs, and adult and subadult males [47, 48, 49, 50, 52]. The availability of food in the autumn likely triggers hibernation, and environmental conditions during winter and spring have been linked to den emergence [49, 50, 53]. On Kodiak Island in the United States, some adult males reduce their activity but do not den throughout the whole winter [54], and there is some evidence of non-denning for whole family groups in the Cantabrian range and Croatia [55, 56]. Brown bears den in natural caves or excavate directly into the ground on dry, steep slopes away from roads and other human disturbances [57, 58, 59, 60, 61]. Lactating females can lose nearly twice as much weight as non-lactating females while denning [62], and because brown bears are easily disturbed while in dens, human disturbances can have negative impacts, especially on cub survival [45, 63].

Population Status

As of 2013, brown bears are 1 of only 2 species of bears that are not considered at least globally vulnerable by the International Union for the Conservation of Nature (IUCN). Globally, the species as a whole is listed as “Least Concern” on the IUCN Red List; however, at the population level, status of brown bears varies widely with geographic location [64]. Northern populations are generally abundant and cover large areas whereas southern populations are highly fragmented and often quite small [64]. Biological populations (i.e., bears in areas with limited opportunities to breed with neighboring bears) often do not correspond to country boundaries. The largest populations are found in Russia (with populations overlapping adjacent countries), Canada, and the northern United States (one population), but brown bears are also fairly common in the Carpathian region (especially in Romania), in the Dinaric Mountains (southern Europe), and in Scandinavia. Populations most likely persist in China and definitely in Japan, although robust population estimates are still unavailable [42].

Population Numbers
Countries and RegionsPopulation Size
Worldwide~200,0001
North America60,078–61,478
Canada~27,578–28,5782
Northern United States, Alaska~31,7003
Southern United States, Lower 48 States1,300–1,6004
Eurasia>62,312–133,121
Europe>17,768–19,970
Carpathian region (total)~8,1005,6
Romania6,6007,8
Slovakia7005
Poland<100a,5,9,10
Ukraine400–9705
SerbiaUnknown6
Alps–Dinaric–Pindos (total)2,100–2,8005,6
Albania, Greece, and Macedonia~250b,5,8
Austria5–8 (introduced)8
Bosnia and Herzegovina12,005
Croatia4005
Czech RepublicUnknown, possibly only vagrants1
Macedonia90–2005,11,†
MontenegroUnknown6
SerbiaUnknown6
Slovenia300–5005,12,*
Scandinavia (total)4,734–5,533a,8
Sweden2,968–3,66713
Norway16614
Finland900–1,000c,14
Estonia7008,15
South Caucasus (Armenia, Azerbaijan, and Georgia)2,000–2,500d,16
Andorra15–21 (recently reoccupied)6,1
Belarus120–2505
Bulgaria5005
France (Pyrenees)~16 (augmented)17
Italy (Alps and Appenines)Min. 43–48,~70† (augmented)18,19,20
Latvia20–405
Spain (Cantabrians)120–1406,8
SwitzerlandUnknown1
TurkeyUnknown21
Asia>44,544–113,151
AfghanistanUnknown22
China1,000–6,300b,23,24,25,†
India500–75025,26,†
IraqUnknown but present27
Islamic Republic of IranPossibly 500–1,00028,†
Japan, Hokkaido1,771–3,6281,24,†
Kazakhstan500–1,000d,22,†
Korea60–210a,24,†
KyrgyzstanUnknown22
Mongolia (Gobi Desert)2329
NepalUnknown, ~40 around the Manasalu Conservation Areac,25
Pakistan150–200, 34 in Deosai National Park25,30
Russia40,000–100,0001,6,14,†
TajikistanUnknown22,†
TurkmenistanUnknown22,†
UzbekistanUnknown22,†

1 [64]; 2 Population estimate for British Columbia in 2004: 16,887 [65], Alberta in 2010: 691 [66], Northern territories: 4,000–5000 [67], and Yukon: 6,000–7,000 [68]; 3 [4]; 4 [69]5 [70]; 6 [71]; 7 [72]; 8 [73]; 9 [74]; 10 [75]; 11 [76]; 12 [77]; 13 [78]; 14 [79]; 15 [2]; 16 [80]; 17 [81]; 18 [82]; 19 [83]; 20 [84]; 21 [85]; 22 [86]; 23 [87]; 24 [41]; 25 [88]; 26 [89]; 27 [90]; 28 [91]; 29 [92]; 30 [93].

a Individuals migrate from Russia to Finland and Norway.

b Population connected with the Alps–Dinara–Pindo region.

c Unknown estimation method.

d Estimates from field transects, interviews, and/or observations (of bear sign).

No robust population estimate.

* Population estimate calculated from a reported average of 13 bears/100 km2 over 95% of their range.

Distribution and Range

Brown bears have the widest distribution of any bear species across the globe, and even though their range was drastically reduced in North America, Europe, and Asia, they are still found in western North America and across Eurasia. Brown bears are currently found in 5,000,000 km2 of North America’s northwestern region, approximately 800,000 km2 of Europe (excluding Russia), and in most of northern Asia [64]. Historically, brown bears were endemic to Europe, Asia, North Africa, and most of North America including Mexico. They were extirpated from Mexico during the 20th Century [64] and from North Africa (Algeria, Morocco, and Tunisia) prior to or during the 19th Century [64, 94]. Brown bears were also extirpated from Belgium, Denmark, Egypt, Germany, Hungary, Ireland, Israel, Jordan, Lebanon, Liechtenstein, Lithuania, Luxembourg, Moldova, Monaco, the Netherlands, Palestine, Portugal, San Marino, Switzerland, the United Kingdom, and probably from Bhutan [64]. Of note, brown bear tracks were observed in Syria in 2004. These tracks are the first evidence of the presence of brown bears in Syria in >40 years [95].

In North America, brown bears were previously found from the Pacific Ocean to the Mississippi River, up into the Arctic and down to central Mexico [3, 96], but they have been extirpated from most of the lower 48 states and the Canadian prairies. They now occupy only 1–2% of their historical range in the lower 48 states [4, 8]; and in Alberta, Canada, they have been designated as threatened since 2010 [66]. In southern Europe, there are a number of very small, isolated populations in the Alps–Dinaric–Pindos region, the Apennine region, the Pyrenees, and the Cantabrian Mountains [64]. In Asia, there are virtually no robust estimates for the many fragmented populations [64], except for bears dwelling in the Gobi Desert of Mongolia [92]. Even though the brown bear range has been reduced on all continents, it is still the most widely distributed bear species on the planet, and can be found at any altitude between sea level and 5,000 m above sea level [64].

Green = Extant, Yellow = Possibly Extant. Map data courtesy of IUCN Redlist.

Habitat

Brown bears are incredibly adaptable. They can be found in a diversity of open and forested habitats from lowlands to mountains, including the arctic tundra, the taiga, boreal forests, temperate rain forests, coastal forests, shrublands and grasslands, and arid regions [1, 7, 64, 97, 98, 99]. Brown bear habitat can be broadly defined by food availability and human avoidance. As a general rule, brown bears tend to persist in areas with low human density, although they are now thriving in Sweden and the Greater Yellowstone Area, USA, because of effective management policies after near-extirpation [100, 101, 102]. Of note, a small population of brown bears persists in the Gobi Desert of Mongolia [92], often using oases complexes where there is little food but also virtually no human activity. Temperatures in the Gobi Desert can fluctuate from −34° to 40°C [103]. Sexual dimorphism and the potential for infanticide also affect habitat selection, with large dominant males utilizing the most productive habitats and females with cubs avoiding sites occupied by large males [104]. Brown bears have a prominent hump of muscles between their shoulders and long claws adapted to digging into hard soil; they therefore prefer open areas but are also commonly found in forested areas, likely because of the cover that forests provide [4, 8]. As a result, brown bears can often be found near natural forest edges or near edges created by roads, forest harvesting, and other human activities [105, 106, 107].

Home Range

Home range sizes vary among populations. Females generally have smaller home ranges than males, and females with cubs-of-the-year have the smallest home ranges [108, 109]. Home range size is also thought to be linked to habitat productivity and food distribution [7]. However, a recent study conducted in Scandinavia linked home range size to population density, but found no relationship with food availability [108]. Overlap among home ranges of related females is common, whereas males tend to have larger home ranges that also overlap those of other males [7, 110, 111].

Home range size (km2)
LocationFemalesMalesSource
Alberta, Canada370-7351824-2152[112]
Jasper National Park, Canada244916[7]
Interior BC, Canada79297[7]
Alaska, USA132710[7]
Kodiak Island, USA1224[7]
Yellowstone NP, USA384828[7]
Mission Mountains, USA2851,402[113]
Scandinavia161–280 (median)833–1,055 (median)[109]
Romania148.6*230.2*[99]
Italy (Alps)178.6* (summer)463.5* (summer)[98]
Slovenia39–63276[114]
Croatia58 (mean)128 (mean)[115]
Japan43NA[55]

Minimum Convex Polygon (MCP) estimating method.

* 95% kernel estimating method.

Estimate based on 3 females.

Diet

Brown bears are opportunistic omnivores and eat an extremely wide variety of food items. Brown bear diet is highly adaptable and follows seasonal fluxes in availability and abundance of foods [88]. Brown bears have a diet similar to American black bears, but compared with other bear species, brown bears are more adapted to digest fibrous foods such as roots [8]. Brown bear diet includes a variety of roots, forbs and grasses, berries (soft mast), nuts and acorns (hard mast), fruit, and insects, as well as small and large live prey including salmon [4, 7]. In the North America interior, fleshy fruits and succulent forbs are the primary food source, but where meat sources are abundant, brown bears are also more carnivorous [64, 116]. Brown bears that feed primarily on fruits and do not have access to salmon are 50% smaller than salmon-feeding bears [117]. Brown bears prefer lipid-rich foods and meat [3]; however, but when given options, they regulate protein and carbohydrate intake to maximize their energy and mass gain [117]. Availability of high-quality foods is especially important in late summer and autumn to prepare for winter dormancy, and because females need to accumulate ≥20% of their body weight as fat to enable the implantation of embryos for reproduction [21, 28]. Brown bears are also known to scavenge food killed by other predators such as tigers, lynx (Lynx sp.), and snow leopards (Uncia uncia) [118]. Brown bears are opportunistic and food generalists; therefore, they are easily attracted to human-related food sources found in remote dwellings, garbage dumps, campgrounds, orchards, and even small to medium-sized settlements. Human-related attractants have been, and remain, a major source of conflict between brown bears and humans.

Natural foods of brown bears include, but are not restricted to the following:

Food typeScientific name
RootsAllium sp., Hedysarum sp., Erythronium sp., Lomatium sp., Oxytropis sp., Perideridia sp.
Forbs, grasses, and sedgesAngelica sp., Arctagrostis sp., Boykinia sp., Brassicaceae sp., Carex sp., Crataegus sp., Eriophorum sp., Equisetum sp., Graminoids, Heracleum sp., Lathyrus sp., Lomatium sp., Malva sp., Medicago sp., Petasites sp., Oxyria sp., Trifolium sp., Taraxacum sp.
Berries–soft mastAmelanchier sp., Arctostaphylos sp., Crataegus sp., Empetrum sp., Helianthus sp., Lonicera sp., Malus sp., Oplopanax sp., Prunus sp., Pyrus sp., Rhamnus sp., Ribes sp., Rosa sp., Rubus sp. Shepherdia canadensis, Sorbus sp., Streptopus sp., Vaccinium sp., Viburnum sp.
Hard mastCorylus sp., Fagus sp., Juglans sp., Pinus sp., Quercus sp.
InvertebratesBombus sp., Camponotus sp., Castanea sp., Chorizagrotis sp., Euxoa sp., Formica sp., Hippodamia sp., Hymenoptera sp., Vespidae sp.
VertebratesAlces alces, Antilocapra sp., Bison bison, Bos grunniens, Castor sp., Cervus sp., Dicrostonyx sp., Equus kiang, Lemmus sp., Lepus sp., Marmota sp., Microtus sp., Myodes sp., Ochotona sp., Odocoileus sp., Oncorhynchus sp., Oreamnos sp., Ovibos sp., Ovis sp., Rangifer sp., Pantholops sp., Phenacomys sp., Peromyscus sp., Pseudois sp., Spermophilus sp., Sus scrofa sp., Synaptomys sp., Tetraoninae sp., Thomomys sp.
FungiRhizopogon sp.

References: [4, 6, 7, 8, 17, 113, 119, 120, 121, 122, 123, 124, 125, 126, 127, 128, 129, 130]

Conservation Status

The brown bear is classified as a species of “Least Concern” on the IUCN Red List. However, all trade in brown bear parts is restricted by the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). Many small isolated populations are at risk of extirpation, but several populations under protection and with good management are expanding, and brown bears are still globally stable. Regional conservation status varies widely among countries and regions. In large populations, brown bears are considered as game and are legally hunted, whereas most of the smaller populations are protected by national and international regulations [64]. Reintroductions and population augmentations have been implemented in several small populations in the United States and Europe [64, 131]. The brown bear populations of Sweden and of the Greater Yellowstone Area, USA, are great examples of populations that have recovered from near extirpation [101, 102].

Threats

Human–bear interactions are the number one cause of brown bear mortalities [132–135]. Major causes of brown bear decline include habitat loss, habitat degradation and fragmentation, industrial and agricultural expansion (overgrazing), inadvertent killings (vehicle or train collisions), sport hunting, illegal hunting, self-defense, removal of “problem bears” (i.e., bears responsible for livestock depredation and human-habituated bears), and illegal trade in bear parts [64, 93, 136–138]. Areas of high human use have the potential to act as population sinks [64, 139], and in some populations, even low numbers of bear deaths can have significant negative impacts on population numbers [130]. Patterns of human settlement are often associated with excessive human-caused mortality and can lead to the fragmentation of populations into small isolated units that face much higher conservation risks [69]. Climate change is not expected to have a direct impact on brown bears, but the indirect impact of climate change on human use of the landscape is expected to increase the potential for human–bear conflicts [49, 140–141].

Brown Bears and Humans

Brown bears are a symbol of power and strength, and often inspire awe as well as instigate fear in humans. Our willingness to coexist with brown bears and our perception of this species will ultimately dictate the extent and success of brown bear distribution and conservation around the globe. Increasing human populations usually result in an increase in human–bear interactions. These interactions are the primary threat to brown bears, often resulting in bear mortality or displacement [118, 135, 139]. Because of this, public awareness and outreach programs are increasingly needed to improve conservation initiatives [142]. As is evident from the successful recovery of brown bear populations in Sweden and the Greater Yellowstone Area, USA, respectful coexistence with brown bears can be achieved and is a worthwhile endeavor. We can learn to successfully coexist with brown bears if we are willing to change our attitudes and behaviors.

Contributor(s)

Karine Pigeon, Ph.D.
Université Laval, Quebec, Canada (affiliation at time of wrtiting).
Current affiliation: http://www.karinepigeon.ca/ Research Gate profile: https://www.researchgate.net/profile/Karine-Pigeon LinkedIn: linkedin.com/in/karine-pigeon-7b773426

References

  1. Paetkau, D., G.F. Shields, and C. Strobeck. 1998. Gene flow between insular, coastal and interior populations of brown bears in Alaska. Molecular Ecology 7:1283–1292. doi: 10.1046/j.1365-294x.1998.00440.x
  2. Tammeleht, E., J. Remm, M. Korsten, J. Davison, I. Tumanov, A. Saveljev, P. Maennil, I. Kojola, and U. Saarma. 2010. Genetic structure in large, continuous mammal populations: The example of brown bears in northwestern Eurasia. Molecular Ecology 19:5359–5370. doi: 10.1111/j.1365-294X.2010.04885.x
  3. Hilderbrand, G.V., C.C. Schwartz, C.T. Robbins, M.E. Jacoby, T.A. Hanley, S.M. Arthur, and C. Servheen. 1999. The importance of meat, particularly salmon, to body size, population productivity, and conservation of North American brown bears. Canadian Journal of Zoology – Revue Canadienne De Zoologie 77:132–138. doi: 10.1139/z98-195
  4. Schwartz, C.C., S.D. Miller, and M.A. Haroldson. 2003. Grizzly bear. Pages 556–586 in G.A. Feldhamer, B.C. Thompson, and J.A. Chapman, editors. Wild mammals of North America: Biology, management, and conservation. Second edition. Johns Hopkins University Press, Baltimore. Maryland, USA.
  5. Whitaker, J.O. 1996. National Audubon Society field guide to North American mammals. National Audubon Society and Alfred A. Knopf, New York, New York, USA.
  6. Zedrosser, A., B. Dahle, and J.E. Swenson. 2006. Population density and food conditions determine adult female body size in brown bears. Journal of Mammalogy 87:510–518.
  7. Pasitschniak-Arts, M. 1993. Ursus arctos. Mammalian Species 439:1–10. doi: 10.1644/05-MAMM-A-218R1.1
  8. Mattson, D.J, S. Herrero, and T. Merrill. 2005. Are black bears a factor in the restoration of North American grizzly bear populations? Ursus 16:11–30. doi: 10.2192/1537-6176(2005)016[0011:ABBAFI]2.0.CO;2
  9. Blanchard, B.M. 1987. Size and growth patterns of the Yellowstone grizzly bear. Ursus 7:99–107. doi: 10.2307/3872615
  10. Ambarli, H. 2006. Analyses of human–bear conflict in Yusufeli, Artvin, Turkey. Thesis, Middle East Technical University, Ankara, Turkey.
  11. Swenson, J.E., M. Adamic, D. Huber, and S. Stokke. 2007. Brown bear body mass and growth in northern and southern Europe. Oecologia 153:37–47. doi: 10.1007/s00442-007-0715-1
  12. Seryodkin, I.V., A.V. Kostyria, J.M. Goodrich, and D.G. Miquelle. 2013. Daily activity patterns of brown bear (Ursus arctos) of the Sikhote–Alin mountain range (Primorskiy Krai, Russia). Russian Journal of Ecology 44:50–55. doi: 10.1134/S1067413613010104
  13. Stenhouse, G.B., J. Boulanger, J. Lee, K. Graham, J. Duval, and J. Cranston. 2005. Grizzly bear associations along the eastern slopes of Alberta. Ursus 16:31–40. doi: 10.2192/1537-6176(2005)016[0031:GBAATE]2.0.CO;2
  14. Støen O.-G., E. Bellemain, S. Saebø, and J.E. Swenson. 2005. Kin-related spatial structure in brown bears Ursus arctos. Behavioral Ecology and Sociobiology 59:191–197. doi: 10.1007/s00265-005-0024-9
  15. Zedrosser, A., O.-G. Støen, S. Saebø, and J.E. Swenson. 2007. Should I stay or should I go? Natal dispersal in the brown bear. Animal Behaviour 74:369–376. doi: 10.1016/j.anbehav.2006.09.015
  16. Egbert, A.L., A.W. Stokes, and A.L. Egbert. 1976. The social behaviour of brown bears on an Alaskan salmon stream. Ursus 3:41–56.
  17. White, D., K.C. Kendall, and H.D. Picton. 1998. Grizzly bear feeding activity at alpine army cutworm moth aggregation sites in northwest Montana. Canadian Journal of Zoology-Revue Canadienne De Zoologie 76:221–227. doi: 10.1139/z97-185
  18. Mattson, D.J., B.M. Blanchard, and R.R. Knight. 1991. Food habits of Yellowstone grizzly bears, 1977–1987. Canadian Journal of Zoology 69:1649–1629. doi: 10.1139/z91-226
  19. Herrero, S. 1978. Comparison of some features of evolution, ecology and behavior of black and grizzly–brown bears. Carnivore 1:7–17.
  20. Garshelis, D.L., M.L. Gibeau, and S. Herrero. 2005. Grizzly bear demographics in and around Banff National Park and Kananaskis Country, Alberta. Journal of Wildlife Management 69:277–297. doi: 10.2193/0022-541X(2005)069<0277:GBDIAA>2.0.CO;2
  21. Robbins, C.T., M. Ben-David, J.K. Fortin, and O.L. Nelson. 2012. Maternal condition determines birth date and growth of newborn bear cubs. Journal of Mammalogy 93:540–546. doi: 10.1644/11-MAMM-A-155.1
  22. Hensel, R.J., W.A. Troyer, and A.W. Erickson. 1969. Reproduction in female brown bear. Journal of Wildlife Management 33:357–365.
  23. Schwartz, C.C., K.A. Keating, H.V. Reynolds, V.G. Barnes, Jr., R.A. Sellers, J.E. Swenson, S.D. Miller, B.N. McLellan, J. Keay, R. McCann, M. Gibeau, W.F. Wakkinen, R.D. Mace, W. Kasworm, R. Smith, and S. Herrero. 2003. Reproductive maturation and senescence in the female brown bear. Ursus 14:109–119.
  24. Wielgus, R.B., and F.L. Bunnell. 2000. Possible negative effects of adult male mortality on female grizzly bear reproduction. Biological Conservation 93:145–154. doi: 10.1016/S0006-3207(99)00152-4
  25. Swenson, J.E., B. Dahle, and F. Sandegren. 2001. Intraspecific predation in Scandinavian brown bears older than cubs-of-the-year. Ursus 12:81–92. Available at: https://www.bearbiology.com/publications/ursus-archive/intraspecific-predation-in-scandinavian-brown-bears-older-than-cubs-of-the-year/
  26. Swenson, J.E., F. Sandegren, S. Brunberg, and P. Segerstrom. 2001. Factors associated with loss of brown bear cubs in Sweden. Ursus 12:69–80. Available at: https://www.bearbiology.com/wp-content/uploads/2017/10/Swenson_Sandegren_Vol_12.pdf
  27. Miquelle, D.G., E.N. Smirnov, H.G. Quigley, I.G. Nikolaev, and E.N. Matyushkin. 1996. Food habits of amur tigers in Sikhote–Alin Zapovednik and the Russian Far East, and implications for conservation. Journal of Wildlife Research 1:138–147.
  28. Van Daele, L. November 2011. International bear news. Quarterly Newsletter of the International Association for Bear Research and Management (IBA) and the IUCN/SSC Bear Specialist Group 20(4):16.
  29. Mech, L.D., and L. Boitani, editors. 2003. Wolves: Behavior, ecology, and conservation. University of Chicago Press, Chicago, Illinois, USA.
  30. Sørenson, J.O., M. Totsas, T. Solståd, and R. Rigg. 2008. Predation by a golden eagle on a brown bear cub. Ursus 19:190–193. doi: 10.2192/08SC008.1
  31. Kasworm, W.F., T.G. Radandt, J.E. Teisberg, M. Proctor, and C. Servheen. 2013. Cabinet–Yaak grizzly bear recovery area 2012 research and monitoring progress report. U.S. Fish & Widlife Service, Missoula, Montana, USA.
  32. Stonorov, D., and A.W. Stokes. 1972. Social behavior of the Alaskan brown bear. Ursus 2:232–242. doi: 10.2307/3872753
  33. Seryodkin, I. 2015. Marking activity of the Kamchatka brown bear. Achievements in the Life Sciences 8:153–161. doi: 10.1016/j.als.2015.04.006
  34. Herrerro, S. 2003. Bear attacks: Their causes and consequences. Second edition. McClelland and Stewart Ltd., Toronto, Ontario, Canada.
  35. Ordiz, A., O.-G. Støen, S. Saebø, V. Sahlen, B.E. Pedersen, J. Kindberg, and J.E. Swenson. 2013. Lasting behavioural responses of brown bears to experimental encounters with humans. Journal of Applied Ecology 50:306–314. doi: 10.1111/1365-2664.12047
  36. Coleman, T.H., C.C. Schwartz, K.A. Gunther, and S. Creel. 2013. Grizzly bear and human interaction in Yellowstone National Park: An evaluation of bear management areas. Journal of Wildlife Management 77:1311–1320. doi: 10.1002/jwmg.602
  37. Smith, T.S., S. Herrero, T.D. Debruyn, and J.M. Wilder. 2008. Efficacy of bear deterrent spray in Alaska. Journal of Wildlife Management 72:640–645. doi: 10.2193/2006-452
  38. Smith, T.S., S. Herrero, C.S. Layton, R.T. Larsen, and K.R. Johnson. 2012. Efficacy of firearms for bear deterrence in Alaska. Journal of Wildlife Management 76:1021–1027. doi: 10.1002/jwmg.342
  39. Frkovic A., D. Huber, and J. Kusak. 2001. Brown bear litter sizes in Croatia. Ursus 12:103–105.
  40. Chapron, G., P.Y. Quenette, S. Legendre, and J. Clobert. 2003. Which future for the French Pyrenean brown bear (Ursus arctos) population? An approach using stage-structured deterministic and stochastic models. Comptes Rendus Biologies 326:174–182. doi: 10.1016/S1631-0691(03)00055-6
  41. Bellemain, E., J.E. Swenson, and P. Taberlet. 2006. Mating strategies in relation to sexually selected infanticide in a non-social carnivore: The brown bear. Ethology 112:238–246. doi: 10.1111/j.1439-0310.2006.01152.x
  42. Japan Bear Network. 2006. Understanding Asian bears to secure their future. Japan Bear Network, Ibaraki, Japan.
  43. Jerina, K., M. Debeljak, S. Dzeroski, A. Kobler, and M. Adamic. 2003. Modeling the brown bear population in Slovenia—A tool in the conservation management of a threatened species. Ecological Modelling 170:453–469. Available at: http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.580.9316&rep=rep1&type=pdf
  44. Watts, P.D., N.A. Øritsland, C. Jonkel, and K. Ronald. 1981. Mammalian hibernation and the oxygen consumption of a denning black bear (Ursus americanus). Comparative Biochemistry and Physiology 69:121–123. doi: 10.1016/0300-9629(81)90645-9
  45. Swenson, J.E., F. Sandegren, S. Brunberg, and P. Wabakken. 1997. Winter den abandonment by brown bears Ursus arctos: Causes and consequences. Wildlife Biology 3:35–38.
  46. Nelson, L.O., and C.T. Robbins. 2015. Cardiovascular function in large to small hibernators: Bears to ground squirrels. Journal of Comparative Physiology B:185:265–279. doi: 10.1007/s00360-014-0881-5
  47. Friebe, A., J.E. Swenson, and F. Sandegren. 2001. Denning chronology of female brown bears in central Sweden. Ursus 12:37–46. doi: 10:2307/3873227
  48. Haroldson, M.A., M.A. Ternent, K.A. Gunther, and C.C. Schwartz. 2002. Grizzly bear denning chronology and movements in the Greater Yellowstone Ecosystem. Ursus 13:29–37. Available at: https://www.bearbiology.com/wp-content/uploads/2017/10/Haroldson_Ternent_gunther_Schwartz_13.pdf
  49. Pigeon, K. 2015. Plasticité comportementale de l’ours grizzli (Ursus arctos horribilis) dans un contexte de changements climatiques. Dissertation, Université Laval, Québec, Canada.
  50. Pigeon, K.E., G.B. Stenhouse, and S.D. Côté. 2016. Drivers of hibernation: Linking food and weather to denning behaviour of grizzly bears. Behavioral Ecology and Sociobiology. 70:1745–1754. doi: 10.1007/s00265-016-2180-5
  51. Nawaz, M.A, J.E. Swenson, and V. Zakaria. 2008. Pragmatic management increases a flagship species, the Himalayan brown bears, in Pakistan’s Deosai National Park. Biological Conservation 141:2230–2241. doi: 10.1016/j.biocon.2008.06.012
  52. Manchi, S., and J.E. Swenson. 2005. Denning behaviour of Scandinavian brown bears Ursus arctos. Wildlife Biology 11:123–132. doi: 10.2981/0909-6396(2005)11[123:DBOSBB]2.0.CO;2
  53. Friebe, A., A.L. Evans, J.M. Arnemo, S. Blanc, S. Brunberg, J.E. Swenson, and A. Zedrosser. 2014. Factors affecting date of implantation, parturition, and den entry estimated from activity and body temperature in free-ranging brown bears. Plos One 9:e101410. doi: 10.1371/journal.pone.0101410
  54. Van Daele, L.J., V.G. Barnes, and R.B. Smith. 1990. Denning characteristics of brown bears on Kodiak Island, Alaska. Ursus 8:257–267. doi: 10.2307/3872927
  55. Sato, Y., Y. Kobayashi, T. Urata, and S. Takatsuki. 2008. Home range and habitat use of female brown bear (Ursus arctos) in Urahoro, eastern Hokkaido, Japan. Mammal Study 33:99–109. doi: 10.3106/1348-6160(2008)33[99:HRAHUO]2.0.CO;2
  56. Nores, C., F. Ballesteros, J.C. Blanco, A. Garcia-Serrano, J. Herrero, and G. Palomero. 2010. Evidence of non-hibernation in Cantabrian brown bears. Acta Theriologica 55:203–209. doi: 10.4098/j.at.0001-7051.085.2008
  57. Petram, W., F. Knauer, and P. Kaczensky. 2004. Human influence on the choice of winter dens by European brown bears in Slovenia. Biological Conservation 119:129–136. doi: 10.1016/j.biocon.2003.07.021
  58. Ciarniello, L.M., M.S. Boyce, D.C. Heard, and D.R. Seip. 2005. Denning behavior and den site selection of grizzly bears along the Parsnip River, British Columbia, Canada. Ursus 16:47–58. doi: 10.2192/1537-6176(2005)016[0047:DBADSS]2.0.CO;2
  59. Elfstrom, M., J.E. Swenson, and J.P. Ball. 2008. Selection of denning habitats by Scandinavian brown bears Ursus arctos. Wildlife Biology 14:176–187. doi: 10.2981/0909-6396(2008)14[176:SODHBS]2.0.CO;2
  60. Goldstein, M.I., A.J. Poe, L.H. Suring, R.M. Nielson, and T.L. McDonald. 2010. Brown bear den habitat and winter Recreation in South-Central Alaska. Journal of Wildlife Management 74:35–42. doi: 10.2193/2008-490
  61. Pigeon, K.E., S.E. Nielsen, G.B. Stenhouse, and S.D. Côté. 2014. Den selection of grizzly bears on a managed landscape. Journal of Mammalogy 9:559–571. doi: 10.1139/z95-262
  62. Farley, S.D., and C.T. Robbins. 1995. Lactation, hibernation, and mass dynamics of American black bears and grizzly bears. Canadian Journal of Zoology-Revue Canadienne De Zoologie 73:2216–2222. doi: 10.1139/z95-262
  63. Evans, A.L., V. Sahlen, O.G. Støen, A. Fahlman, S. Brunberg, K. Madslien, O. Frobert, J. E. Swenson, and J.M. Arnemo. 2012. Capture, anesthesia, and disturbance of free-ranging brown bears (Ursus arctos) during hibernation. Plos One 7:e40520. doi: 10.1371/journal.pone.0040520
  64. McLellan, B.N., C. Servheen, and D. Huber [IUCN SSC Bear Specialist Group]. 2008. Ursus arctos. In IUCN 2013. International Union for the Conservation of Nature Red List of Threatened Species. Version 2013.1. International Union for the Conservation of Nature, Gland, Switzerland; and Cambridge, England, UK. http://www.iucnredlist.org/details/41688/0. Accessed 20 Jun 2013.
  65. Hamilton, A.N., D.C. Heard, and M.A. Austin, 2004. British Columbia grizzly bear (Ursus arctos) population estimate. British Columbia Ministry of Water, Land and Air Protection, Victoria, British Columbia, Canada.
  66. Alberta Sustainable Resource Development and Alberta Conservation Association. 2010. Status of the grizzly bear (Ursus arctos) in Alberta. Wildlife Status Report No. 37: Update 2010. Alberta Sustainable Resource Development, Edmonton, Alberta, Canada.
  67. Northwest Territories Government. 2013. NWT species at risk. Northwest Territories Government, Yellowknife, Northwest Territories, Canada. http://nwtspeciesatrisk.ca/tiki/tiki-index.php?page=GrizzlyBear. Accessed 20 Jun 2013.
  68. Government of Yukon. 2013. Environment, mammals, bear grizzly. May 28, 2013. http://www.env.gov.yk.ca/animals-habitat/mammals/grizzly.php. Accessed 20 Jun 2013.
  69. Proctor, M.F., D. Paetkau, B.N. Mclellan, G.B. Stenhouse, K.C. Kendall, R.D. 4Mace, W.F. Kasworm, C. Servheen, C.L. Lausen, M.L. Gibeau, W.L. Wakkinen, M.A. Haroldson, G. Mowat, C.D. Apps, L.M. Ciarniello, R.M.R. Barclay, M.S. Boyce, C.C. Schwartz, and C. Strobeck. 2012. Population fragmentation and inter-ecosystem movements of grizzly bears in western Canada and the northern United States. Wildlife Monographs 180. doi: 10.1002/wmon.6
  70. Zedrosser, A., B. Dahle, J.E. Swenson, and N. Gerstl. 2001. Status and management of the brown bear in Europe. Ursus 12:9–20. Available at: http://www2.nina.no/lcie_new/pdf/634991427086642093_Zedroser%20URSUS%20Europe%20bear%20status.pdf
  71. Linnell, J., V. Salvatori, and L. Boitani. 2008. Guidelines for population level management plans for large carnivores in Europe. Contract 070501/2005/424162/MAR/B2. Report prepared for the European Commission by A Large Carnivore Initiative for Europe, Rome, Italy. Available at: http://ec.europa.eu/environment/nature/conservation/species/carnivores/pdf/guidelines_for_population_level_management.pdf. Accessed 25 Jun 2013.
  72. Karamanlidis, A.A., M. Straka, E. Drosopoulou, M.D. Hernando, I. Kocijan, L. Paule, and Z. Scouras. 2012. Genetic diversity, structure, and size of an endangered brown bear population threatened by highway construction in the Pindos Mountains, Greece. European Journal of Wildlife Research 58:511–522. doi: 10.1007/s10344-011-0598-7
  73. Swenson, J.E., P. Taberlet, and E. Bellemain. 2011. Genetics and conservation of European brown bears Ursus arctos. Mammal Review 41:87–98. doi: 10.1111/j.1365-2907.2010.00179.x
  74. Gula, R., W. Frackowiak, and K. Perzanowski. 1998. Current status and conservation needs of brown bears in the Polish Carpathians. Ursus 10:81–86. Available at: https://www.bearbiology.com/publications/ursus-archive/current-status-and-conservation-needs-of-brown-bears-in-the-polish-carpathians/
  75. Sergiel, A. May 2011. International bear news. Quarterly Newsletter of the International Association for Bear Research and Management (IBA) and the IUCN/SSC Bear Specialist Group 20 (2):15–18.
  76. Karamanlidis, A.A., A. Stojanov, M. De Gabriel Hernando, G. Ivanov, I. Kocijan, D. Melovski, T. Skrbinsek, and A. Zedrosser. 2014. Distribution and genetic status of brown bear in FYR Macedonia: Implications for conservation. Acta Theriologica 59:119–128. doi: 10.1007/s13364-013-0147-8
  77. Jerina, K., M. Jonozovic, M. Krofel, and T. Skrbinsek. 2013. Range and local population densities of brown bear Ursus arctos in Slovenia. European Journal of Wildlife Research 59:459–467. soi: 10.1007/s10344-013-0690-2
  78. Kindberg, J., J.E. Swenson, G. Ericsson, E. Bellemain, C. Miquel, and P. Taberlet. 2011. Estimating population size and trends of the Swedish brown bear Ursus arctos population. Wildlife Biology 17:114–123. doi: 10.2981/10-100
  79. Kopatz, A., H.G. Eiken, S.B. Hagen, M. Ruokonen, R. Esparza-Salas, J. Schregel, I. Kojola, M.E. Smith, I. Wartiainen, P.E. Aspholm, S. Wikan, A.M. Rykov, O. Makarova, N. Polikarpova, K.F. Tirronen, P.I. Danilov, and J. Aspi. 2012b. Connectivity and population subdivision at the fringe of a large brown bear (Ursus arctos) population in North Western Europe. Conservation Genetics 13:681–692. doi: 10.1007/s10592-012-0317-2
  80. Lortkipanidze, B. 2010. Brown bear distribution and status in the South Caucasus. Ursus 21:97–103. doi: 10.2192/09GR017.1
  81. Camarra, J.J., F. Decaluwe, and P.Y. Quenette. November 2011. International bear news. Quarterly Newsletter of the International Association for Bear Research and Management (IBA) and the IUCN/SSC Bear Specialist Group 20 (4):34–35.
  82. De Barba, M., L.P. Waits, E.O. Garton, P. Genovesi, E. Randi, A. Mustoni, and C. Groff. 2010. The power of genetic monitoring for studying demography, ecology and genetics of a reintroduced brown bear population. Molecular Ecology 19:3938–3951. doi: 10.1111/j.1365-294X.2010.04791.x
  83. Ciucci, P., and L. Boitani. 2008. The Apennine brown bear: A critical review of its status and conservation problems. Ursus 19:130–145. doi: 10.2192/07PER012.1
  84. Groff, C. Summer 2013. International bear news. Quarterly Newsletter of the International Association for Bear Research and Management (IBA) and the IUCN/SSC Bear Specialist Group 22 (2):19–21.
  85. Can, Ö.E., and I. Togan. 2004. Status and management of brown bears in Turkey. Ursus 15:48–53. doi: 10.2192/1537-6176(2004)015<0048:SAMOBB>2.0.CO;2
  86. Moheb, Z., D. Lawson, and S.N. Mostafawi. 2012. Brown bear status and threats in Darwaz, Northern Badakhshan, Afghanistan. Ursus 23:237–240. doi: 10.2307/41933231
  87. Jien, G., and R.B. Harris. 2006. The status of bears in China: Country report. Available at: http://people.montana.com/~rharris/China%20final.pdf. Accessed 25 Jun 2013.
  88. Aryal, A., S. Sathyakumar, and C.C. Schwartz. 2010. Current status of brown bears in the Manasalu Conservation Area, Nepal. Ursus 21:109–114. doi: 10.2192/09GR029.1
  89. Sathyakumar, S. 2006. Status and distribution of Himalayan brown bear (Ursus arctos isabellinus) in India: An assessment of changes over ten years. Indian Forester 132:89–96.
  90. Garshelis, D. February 2011. International Bear News. Quarterly Newsletter of the International Association for Bear Research and Management (IBA) and the IUCN/SSC Bear Specialist Group 20 (1):5–7.
  91. Gutleb, B., and H. Ziaie. 1999. On the distribution and status of the brown bear, Ursus arctos, and the Asiatic black bear, U. thibetanus, in Iran. Zoology in the Middle East 18:5–8. doi: 10.1080/09397140.1999.10637777
  92. Tumendemberel, O., M. Proctor, H. Reynolds, L. Amgalan, T. Tserenbataa, M. Batmunkh, D. Craighead, N. Yanjin, and D. Paetkau. 2012. Gobi bear population survey 2009. Gobi Bear Project, Fairbanks, Alaska, USA. Available at: http://static.squarespace.com/static/528f911de4b01f2a31514e96/t/52d820c5e4b0796add7cb26d/1389895877671/Gobi_Bear_2009_Population_Survey_Report_FINAL_for_website.pdf. Accessed 25 Jun 2013.
  93. Nawaz, M.A. 2007. Status of the brown bear in Pakistan. Ursus 18:89–100. doi: 10.2192/1537-6176(2007)18[89:SOTBBI]2.0.CO;2
  94. Calvignac, S., S. Hughes, C. Tougard, J. Michaux, M. Thevenot, M. Philippe, W. Hamdine, and C. Hanni. 2008. Ancient DNA evidence for the loss of a highly divergent brown bear clade during historical times. Molecular Ecology 17:1962–1970. doi: 10.1111/j.1365-294X.2008.03631.x
  95. Hajjar, I. February 2011. International bear news. Quarterly Newsletter of the International Association for Bear Research and Management (IBA) and the IUCN/SSC Bear Specialist Group 20 (1):7–11.
  96. Servheen, C. 1984. The status of the grizzly bear and the interagency grizzly bear recovery effort. Proceedings of the Annual Conference of the Western Association of Fish and Wildlife Agencies 64:227–234. Available at: https://www.bearbiology.com/fileadmin/tpl/Downloads/URSUS/Vol_10/ServheenB_Vol_10.pdf
  97. Stelmock, J.T., and F.C. Dean. 1986. Brown bear activity and habitat use, Denali National Park: 1980. Paper presented at 6th International Conference on Bear Research and Management, February 1983, Grand Canyon, Arizona, USA.
  98. Preatoni, D., A. Mustoni, A. Martinoli, E. Carlini, B. Chiarenzi, S. Chiozzini, S. Van Dongen, L.A. Wauters, and G. Tosi. 2005. Conservation of brown bear in the Alps: Space use and settlement behavior of reintroduced bears. Acta Oecologica-International Journal of Ecology 28:189–197. doi: 10.1016/j.actao.2005.04.002
  99. Pop, I.M., A. Sallay, L. Bereczky, and S. Chiriac. 2012. Land use and behavioral patterns of brown bears in the South-Eastern Romanian Carpathian Mountains: A case study of relocated and rehabilitated individuals. Procedia Environmental Sciences 14:111–122. doi: 10.1016/j.proenv.2012.03.011
  100. Swenson, J.E., P. Wabakken, F. Sandegren, A. Bjärvall, R. Franzén, and A. Söderberg. 1995. The near extinction and recovery of brown bears in Scandinavia in relation to the bear management policies of Norway and Sweden. Wildlife Biology 1:11–25.
  101. Interagency Conservation Strategy Team. 2007. Final conservation strategy for the grizzly bear in the Greater Yellowstone Area. U.S. Fish & Wildlife Service. http://www.fws.gov/mountain-prairie/species/mammals/grizzly/yellowstoneindex.html. Accessed 20 Jun 2013.
  102. Swenson, J.E., A. Ordiz, J. Martin, S. Steyaert, V. Sahlen, O.-G. Stoen, and J. Kindberg. 2012. How do 3,300 brown bears cope with 9.4 million Swedes? Oral presentation. 21st International Conference on Bear Research and Management, 26–30 Nov 2012, New Delhi, India.
  103. Dugarjav, C., and B. Tsetseg. 2003. Conservation and sustainable use of biodiversity in the Trans–Altai Gobi Desert of Mongolia. Pages 137–156 in J. Lemons, R. Victor, and D. Schaffer, editors. Conserving biodiversity in arid regions. Kluwer Academic Publishers, Boston, Massachusetts, USA.
  104. Rode, K.D., S.D. Farley, and C.T. Robbins. 2006. Sexual dimorphism, reproductive strategy, and human activities determine resource use by brown bears. Ecology 87:2636–2646. doi: 10.1890/0012-9658(2006)87[2636:SDRSAH]2.0.CO;2
  105. Nielsen, S.E., G.B. Stenhouse, and M.S. Boyce. 2006. A habitat-based framework for grizzly bear conservation in Alberta. Biological Conservation 130:217–229. doi: 10.1016/j.biocon.2005.12.016
  106. Stewart, B.P., T.A. Nelson, K. Laberee, S.E. Nielsen, M.A. Wulder, and G. Stenhouse. 2013. Quantifying grizzly bear selection of natural and anthropogenic edges. Journal of Wildlife Management 77:957–964. doi: 10.1002/jwmg.535
  107. Roever, C.L., M.S. Boyce, and G.B. Stenhouse. 2008. Grizzly bears and forestry—II: Grizzly bear habitat selection and conflicts with road placement. Forest Ecology and Management 256:1262–1269. doi: 10.1016/j.foreco.2008.06.006
  108. Dahle, B., O.G. Støen, and J.E. Swenson. 2006. Factors influencing home-range size in subadult brown bears. Journal of Mammalogy 87:859–865.
    doi: 10.1644/05-MAMM-A-352R1.1
  109. Dahle, B., and J.E. Swenson. 2003. Seasonal range size in relation to reproductive strategies in brown bears Ursus arctos. Journal of Animal Ecology 72:660–667. doi: 10.1046/j.1365-2656.2003.00737.x
  110. Støen, O.-G., A. Zedrosser, S. Saebø, and J.E. Swenson. 2006. Inversely density-dependent natal dispersal in brown bears Ursus arctos. Oecologia 148:356–364.
  111. McLellan, B.N., and F.W. Hovey. 2001. Natal dispersal of grizzly bears. Canadian Journal of Zoology 79:838–844. doi: 10.1139/z01-051
  112. Graham, K., and G.B. Stenhouse. 2014. Has advances in radio telemetry increased our knowledge of grizzly bear ecology and benefited grizzly bear conservation in Alberta? The Canadian Field Naturalist 128:223–233.
  113. Servheen, C. 1983. Grizzly bear food habits, movements, and habitat selection in the Mission Mountains, Montana. Journal of Wildlife Management 47:1026–1035. doi: 10.2307/3808161
  114. Kaczensky, P., F. Knauer, B. Krze, M. Jonozovic, M. Adamic, and H. Gossow. 2003. The impact of high speed, high volume traffic axes on brown bears in Slovenia. Biological Conservation 111:191–204. doi: 10.1016/S0006-3207(02)00273-2
  115. Huber, D., I. Kulier, A. Poljak, and B. Devcic-Kuhar. 1993. Food-intake and mass gain of hand-reared brown bear cubs. Zoo Biology 12:525–533. doi: 10.1002/zoo.1430120604
  116. Mattson, D.J., and T. Merrill. 2002. Extirpations of grizzly bears in the contiguous United States, 1850–2000. Conservation Biology 16:1123–1136. doi: 10.1046/j.1523-1739.2002.00414.x
  117. Robbins, C.T., J.K. Fortin, K.D. Rode, S.D. Farley, L.A. Shipley, and L.A. Felicetti. 2007. Optimizing protein intake as a foraging strategy to maximize mass gain in an omnivore. Oikos 116:1675–1682. doi: 10.1111/j.0030-1299.2007.16140.x
  118. Krofel, M., I. Kos, and K. Jerina. 2012. The noble cats and the big bad scavengers: Effects of dominant scavengers on solitary predators. Behavioral Ecology and Sociobiology 66:1297–1304. doi: 10.1007/s00265-012-1384-6
  119. Munro, R.H.M., S.E. Nielsen, M.H. Price, G.B. Stenhouse, and M.S. Boyce. 2006. Seasonal and diel patterns of grizzly bear diet and activity in west-central Alberta. Journal of Mammalogy 87:1112–1121. doi: 10.1644/05-MAMM-A-410R3.1
  120. Martin, J. 2009. Habitat selection and movement by brown bears in multiple-use landscapes. Dissertation, Université Claude-Bernard – Lyon 1, Villeurbanne, France.
  121. McLellan, B.N., and F.W. Hovey. 1995. The diet of grizzly bears in the Flathead River drainage of southeastern British Columbia. Canadian Journal of Zoology-Revue Canadienne De Zoologie 73:704–712. doi: 10.1139/z95-082
  122. Fortin, J.K., C.C. Schwartz, K.A. Gunther, J.E. Teisberg, M.A. Haroldson, M.A. Evans, and C.T. Robbins. 2013. Dietary adjustability of grizzly bears and American black bears in Yellowstone National Park. Journal of Wildlife Management 77:270–281. doi: 10.1002/jwmg.483
  123. Nielsen, S.E., G. McDermid, G.B. Stenhouse, and M.S. Boyce. 2010. Dynamic wildlife habitat models: Seasonal foods and mortality risk predict occupancy-abundance and habitat selection in grizzly bears. Biological Conservation 143:1623–1634. doi: 10.1016/j.biocon.2010.04.007
  124. Hamer, D., and S. Herrero. 1987. Grizzly bear food and habitat in the front ranges of Banff National Park, Alberta. Ursus 7:199–213. doi: 10.2193/0022-541X(2005)069<0277:GBDIAA>2.0.CO;2
  125. MacHutchon, A.G., and D.W. Wellwood. 2003. Grizzly bear food habits in the northern Yukon, Canada. Ursus 14:225–235. Available at: https://www.bearbiology.com/publications/ursus-archive/grizzly-bear-food-habits-in-the-northern-yukon-canada/
  126. Xu, A., Z. Jiang, C. Li, J. Guo, G. Wu, and P. Cai. 2006. Summer food habits of brown bears in Kekexili Nature Reserve, Qinghai-Tibetan Plateau, China. Ursus 17:132–137. doi: 10.2192/1537-6176(2006)17[132:SFHOBB]2.0.CO;2
  127. Bojarska, K., and N. Selva. 2013. Correction factors for important brown bear foods in Europe. Ursus 24:13–15. doi: 10.2192/URSUS-D-12-00017.1
  128. Rodriguez, C., J. Naves, A. Fernandez-Gil, J.R. Obeso, and M. Delibes. 2007. Long-term trends in food habits of a relict brown bear population in northern Spain: The influence of climate and local factors. Environmental Conservation 34:36–44. doi: 10.1017/S0376892906003535
  129. Aryal, A., D. Raubenheimer, S. Sathyakumar, B.S. Poudel, W. Ji, K.J. Kunwar, J. Kok, S. Kohshima, and D. Brunton. 2012. Conservation strategy for brown bear and its habitat in Nepal. Diversity 4:301–317 (open access). doi: 10.3390/d4030301
  130. Wu, L., J. Li, and Z. Lu. Fall 2013. International bear news. Quarterly Newsletter of the International Association for Bear Research and Management (IBA) and the IUCN/SSC Bear Specialist Group 22 (3):20–21.
  131. Clark, J.D., D. Huber, and C. Servheen. 2002. Bear reintroductions: Lessons and challenges. Ursus 13:335–345. Available at: http://www2.nina.no/lcie_new/pdf/634991395729973090_Clark%20URSUS%20Bear%20reintroductions.pdf
  132. McLellan, B.N., F.W. Hovey, R.D. Mace, J.G. Woods, D.W. Carney, M.L. Gibeau, W.L. Wakkinen, and W.F. Kasworm. 1999. Rates and causes of grizzly bear mortality in the interior mountains of British Columbia, Alberta, Montana, Washington, and Idaho. Journal of Wildlife Management 63:911–920. doi: 10.2307/3802805
  133. Benn, B., and S. Herrero. 2002. Grizzly bear mortality and human access in Banff and Yoho National Parks. Ursus 13:213–221. Available at: https://www.bearbiology.com/publications/ursus-archive/grizzly-bear-mortality-and-human-access-in-banff-and-yoho-national-parks-1971-98/
  134. Wakkinen, W.L., and W.F. Kasworm. 2004. Demographics and population trends of grizzly bears in the Cabinet–Yaak and Selkirk ecosystems of British Columbia, Idaho, Montana, and Washington. Ursus 15:65–75. Available at: https://www.bearbiology.com/publications/ursus-archive/demographics-and-population-trends-of-grizzly-bears-in-the-cabinet-yaak-and-selkirk-ecosystems-of-british-columbia-idaho-montana-and-washington/
  135. Can, Ö.E., N. D’Cruze, D.L. Garshelis, J. Beecham, and D.W. Macdonald. 2014. Resolving human–bear conflict: A global survey of countries, experts, and key factors. Conservation Letters 7:501–513. doi: 10.1111/conl.12117
  136. Proctor, M.F., B.N. McLellan, C. Strobeck, and R.M.R. Barclay. 2005. Genetic analysis reveals demographic fragmentation of grizzly bears yielding vulnerably small populations. Proceedings of the Royal Society B-Biological Sciences 272:2409–2416. doi: 10.1098/rspb.2005.3246
  137. Waller, J.S., and C. Servheen. 2005. Effects of transportation infrastructure on grizzly bears in northwestern Montana. Journal of Wildlife Management 69:985–1000. doi: 10.2193/0022-541X(2005)069[0985:EOTIOG]2.0.CO;2
  138. Bischof, R., J.E. Swenson, N.G. Yoccoz, A. Mysterud, and O. Gimenez. 2009. The magnitude and selectivity of natural and multiple anthropogenic mortality causes in hunted brown bears. Journal of Animal Ecology 78:656–665. doi: 10.1111/j.1365-2656.2009.01524.x
  139. Nielsen, S.E., S. Herrero, M.S. Boyce, R.D. Mace, B. Benn, M.L. Gibeau, and S. Jevons. 2004. Modelling the spatial distribution of human-caused grizzly bear mortalities in the Central Rockies ecosystem of Canada. Biological Conservation 120:101–113.
    Available at: http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.533.2686&rep=rep1&type=pdf
  140. Servheen, C. November 2011. International bear news. Quarterly Newsletter of the International Association for Bear Research and Management (IBA) and the IUCN/SSC Bear Specialist Group 20 (4):30–33.
  141. Pigeon, K.E., E. Cardinal, G.B. Stenhouse, and S.D. Côté. 2016. Staying cool in a changing landscape: The influence of maximum daily ambient temperature on grizzly bear habitat selection. Oecologia. 181:1101–1116. doi: 10.1007/s00442-016-3630-5
  142. Sakurai, R., S.K. Jacobson, and G. Ueda. 2013. Public perceptions of risk and government performance regarding bear management in Japan. Ursus 24:70–82. Available at: https://www.bearbiology.com/publications/ursus-archive/public-perceptions-of-risk-and-government-performance-regarding-bear-management-in-japan/
  143.  

Links