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1.
Stereotypical behaviors in captive polar bears (Ursus maritimus) can be detrimental to their welfare. These behaviors can be reduced through enrichment programs but are often not completely eliminated, so identifying potential triggers is important. The present study investigated the influences of seasonal changes, visitor density, and concurrent bear activity on stereotypical behaviors exhibited by 3 captive polar bears at the Toronto Zoo. All bears exhibited these behaviors; however, individual differences were found in duration and form. The male exhibited less stereotypical behavior during spring, and the females exhibited less stereotypical behavior during winter. An increase in visitor density was associated with more stereotypical behavior in 1 female but less stereotypical behavior in the other 2 bears. All bears engaged in more stereotypical behaviors when the other bears were inactive, and 1 female engaged in more stereotypical behaviors when the other bears were out of sight. Further, when conspecifics were active, all bears engaged in less stereotypical behaviors. Given the variability among individual bears, future enrichment programs must be tailored to the needs of individuals to maximize efficacy.  相似文献   

2.
ABSTRACT The distribution of grizzly (Ursus arctos) and American black bears (U. americanus) overlaps in western North America. Few studies have detailed activity patterns where the species are sympatric and no studies contrasted patterns where populations are both sympatric and allopatric. We contrasted activity patterns for sympatric black and grizzly bears and for black bears allopatric to grizzly bears, how human influences altered patterns, and rates of grizzly-black bear predation. Activity patterns differed between black bear populations, with those sympatric to grizzly bears more day-active. Activity patterns of black bears allopatric with grizzly bears were similar to those of female grizzly bears; both were crepuscular and day-active. Male grizzly bears were crepuscular and night-active. Both species were more night-active and less day-active when ≤1 km from roads or developments. In our sympatric study area, 2 of 4 black bear mortalities were due to grizzly bear predation. Our results suggested patterns of activity that allowed for intra- and inter-species avoidance. National park management often results in convergence of locally high human densities in quality bear habitat. Our data provide additional understanding into how bears alter their activity patterns in response to other bears and humans and should help park managers minimize undesirable bear-human encounters when considering needs for temporal and spatial management of humans and human developments in bear habitats.  相似文献   

3.
Previous studies on the mating system of the Asian black bear (Ursus thibetanus) have been limited to observations of captive populations and estimations of multiple paternities. Hence, the mating system of wild bears remains poorly understood. Animal‐borne camera systems (i.e., cameras mounted on animals) provide novel tools to study the behavior of elusive animals. Here, we used an animal‐borne video system to record the activities of wild bears during the mating season. Video camera collars were attached to four adult Asian black bears (male “A” and “B,” and female “A” and “B”) captured in Tokyo, central Japan, in May and June 2018. The collars were retrieved in July 2018, after which the video data were downloaded and analyzed in terms of bear activity and mating behavior. All the bears were found to interact with other uniquely identifiable bears for some of the time (range 9–22 days) during the deployment period (range 36–45 days), and multiple mating in males was documented. Both males and females exhibited different behaviors on social days (i.e., days when the bear interacted with conspecifics) compared with solitary days (i.e., days with no observed interactions with conspecifics). Compared with solitary days, the bears spent a lower proportion of time on foraging activities and higher proportion of time on resting activities on social days. Our results suggest that Asian black bears have a polygamous mating system, as both sexes consort and potentially mate with multiple partners during a given mating season. Furthermore, bears appeared to reduce their foraging activities on social days and engaged more in social interactions.  相似文献   

4.
At least 63 people were killed in 59 incidents by non-captive black bear (Ursus americanus) during 1900–2009. Fatal black bear attacks occurred in Canada and Alaska (n = 49) and in the lower 48 states (n = 14). There were 3.5 times as many fatal attacks in Canada and Alaska but only 1.75 times as many black bears, and much less human contact for black bears in Canada and Alaska. There was a weak positive correlation (rs = 0.56, P ≤ 0.000) between the estimated size of a bear population within a given jurisdiction and the number of fatal black bear attacks. Some jurisdictions had no fatal black bear attacks but had large estimated black bear populations. Of fatal attacks, 86% (54 of 63, 1.08/yr) occurred between 1960 and 2009. There was positive linear relationship between the number of fatal black bear attacks per decade and human population size in the United States and Canada per decade (r2 = 0.92, β = 0.000, P ≤ 0.001). Of fatal attacks, 91% (49 of 54) occurred on parties of 1 or 2 persons. In 38% (15 of 40) of incidents, peoples' food or garbage probably influenced the bear being in the attack location. We judged that the bear involved acted as a predator in 88% (49 of 56) of fatal incidents. Adult (n = 23) or subadult (n = 10) male bears were involved in 92% (33 of 36) of fatal predatory incidents, reflecting biological and behavioral differences between male and female bears. That most fatal black bear attacks were predatory and were carried out by 1 bear shows that females with young are not the most dangerous black bears. As a result of our research agencies managing black bear can more accurately understand the risk of being killed by a black bear, and can communicate this to the public. With training, people can learn to recognize the behaviors of a bear considering them as prey and can act to deter predation. © 2011 The Wildlife Society.  相似文献   

5.
Bears in captivity often show abnormal behaviors such as stereotypies and periods of excessive inactivity. Most of the research aimed at reducing abnormal behaviors in bears has focused on feeding enrichment. However, the effects of feeding enrichment only seem to last as long as it takes a bear to consume the food. This study investigated the effects of inedible, manipulable objects on bears. Two polar bears (Ursus maritimus) received plastic floats. A sloth bear (Melursus ursinus) and a spectacled bear (Treniarctos ornatus) received plastic balls. During a 7-week period, polar bears nearly doubled their activity in the presence of toy floats, and the spectacled bear halved its pacing in the presence of toys. The sloth bear did not respond to the objects. These findings indicate that access to manipulable nonfood objects such as plastic toys can result in prolonged engagement in appetitive behaviors by polar bears.  相似文献   

6.
We combine a recently developed framework for describing dietary generalism with compositional data analysis to examine patterns of omnivory in a large widely distributed mammal. Using the brown bear (Ursus arctos) as a model species, we collected and analyzed data from the literature to estimate the proportions of macronutrients (protein, carbohydrate, and lipid) in the diets of bear populations. Across their range, bears consumed a diversity of foods that resulted in annual population diets that varied in macronutrient proportions, suggesting a wide fundamental macronutrient niche. The variance matrix of pairwise macronutrient log‐ratios indicated that the most variable macronutrient among diets was carbohydrate, while protein and lipid were more proportional or codependent (i.e., relatively more constant log‐ratios). Populations that consumed anthropogenic foods, such agricultural crops and supplementary feed (e.g., corn), had a higher geometric mean proportion of carbohydrate, and lower proportion of protein, in annual diets. Seasonally, mean diets were lower in protein and higher in carbohydrate, during autumn compared to spring. Populations with anthropogenic subsidies, however, had higher mean proportions of carbohydrate and lower protein, across seasons compared to populations with natural diets. Proportions of macronutrients similar to those selected in experiments by captive brown bears, and which optimized primarily fat mass gain, were observed among hyperphagic prehibernation autumn diets. However, the majority of these were from populations consuming anthropogenic foods, while diets of natural populations were more variable and typically higher in protein. Some anthropogenic diets were close to the proportions selected by captive bears during summer. Our results suggest that omnivory in brown bears is a functional adaptation enabling them to occupy a diverse range of habitats and tolerate variation in the nutritional composition and availability of food resources. Furthermore, we show that populations consuming human‐sourced foods have different dietary macronutrient proportions relative to populations with natural diets.  相似文献   

7.
Aim  Middle East brown bears ( Ursus arctos syriacus Hemprich and Ehrenberg, 1828) are presently on the edge of extinction. However, little is known of their genetic diversity. This study investigates that question as well as that of Middle East brown bear relationships to surrounding populations of the species.
Location  Middle East region of south-western Asia.
Methods  We performed DNA analyses on 27 brown bear individuals. Twenty ancient bone samples (Late Pleistocene to 20th century) from natural populations and seven present-day samples obtained from captive individuals were analysed.
Results  Phylogenetic analyses of the mitochondrial sequences obtained from seven ancient specimens identify three distinct maternal clades, all unrelated to one recently described from North Africa. Brown bears from Iran exhibit striking diversity (three individuals, three haplotypes) and form a unique clade that cannot be linked to any extant one. Individuals from Syria belong to the Holarctic clade now observed in Eastern Europe, Turkey, Japan and North America. Specimens from Lebanon surprisingly appear as tightly linked to the clade of brown bears now in Western Europe. Moreover, we show that U. a. syriacus in captivity still harbour haplotypes closely linked to those found in ancient individuals.
Main conclusion  This study brings important new information on the genetic diversity of brown bear populations at the crossroads of Europe, Asia and Africa. It reveals a high level of diversity in Middle East brown bears and extends the historical distribution of the Western European clade to the East. Our analyses also suggest the value of a specific breeding programme for captive populations.  相似文献   

8.
Debra Kutska 《Zoo biology》2009,28(4):292-306
A debate exists among modern zoo staff as to whether or not the addition of un-naturalistic enrichment takes away from, or even defeats, the educational messages designers are trying to incorporate in naturalistic exhibits. A visitor study was conducted at the Central Park Zoo's polar bear exhibit in order to determine whether or not the type of enrichment in an enclosure actually alters guest perceptions. Visitors were exposed to one of two enrichment treatments in the bear enclosure: Naturalistic or Un-naturalistic. The results of this study suggest that enrichment type did not alter the perceptions of visitors. However, it did identify some of the different ways adults and youths perceive animals and zoos. Additionally, the study highlighted the varying perceptions individuals have of the concept of polar bears vs. their perceptions of the captive individuals at the Central Park Zoo. Implications for enrichment usage and exhibit design are discussed. Zoo Biol 28:292–306, 2009. © 2008 Wiley-Liss, Inc.  相似文献   

9.
Grizzly bears (Ursus arctos) and American black bears (U. americanus) are sympatric in much of Yellowstone National Park. Three primary bear foods, cutthroat trout (Oncorhynchus clarki), whitebark pine (Pinus albicaulis) nuts, and elk (Cervus elaphus), have declined in recent years. Because park managers and the public are concerned about the impact created by reductions in these foods, we quantified bear diets to determine how bears living near Yellowstone Lake are adjusting. We estimated diets using: 1) stable isotope and mercury analyses of hair samples collected from captured bears and from hair collection sites established along cutthroat trout spawning streams and 2) visits to recent locations occupied by bears wearing Global Positioning System collars to identify signs of feeding behavior and to collect scats for macroscopic identification of residues. Approximately 45 ± 22% ( ± SD) of the assimilated nitrogen consumed by male grizzly bears, 38 ± 20% by female grizzly bears, and 23 ± 7% by male and female black bears came from animal matter. These assimilated dietary proportions for female grizzly bears were the same as 10 years earlier in the Lake area and 30 years earlier in the Greater Yellowstone Ecosystem. However, the proportion of meat in the assimilated diet of male grizzly bears decreased over both time frames. The estimated biomass of cutthroat trout consumed by grizzly bears and black bears declined 70% and 95%, respectively, in the decade between 1997–2000 and 2007–2009. Grizzly bears killed an elk calf every 4.3 ± 2.7 days and black bears every 8.0 ± 4.0 days during June. Elk accounted for 84% of all ungulates consumed by both bear species. Whitebark pine nuts continue to be a primary food source for both grizzly bears and black bears when abundant, but are replaced by false-truffles (Rhizopogon spp.) in the diets of female grizzly bears and black bears when nut crops are minimal. Thus, both grizzly bears and black bears continue to adjust to changing resources, with larger grizzly bears continuing to occupy a more carnivorous niche than the smaller, more herbivorous black bear. © 2012 The Wildlife Society.  相似文献   

10.
Sympatric black bears (Ursus americanus) and brown bears (Ursus arctos) are common in many boreal systems; however, few predator assemblages are known to coexist on a single seasonally abundant large prey item. In lowland southwestern interior Alaska, black bears and brown bears are considered the primary cause of moose (Alces alces) calf mortality during the first 6 weeks of life. The objective of this study was to document habitat use of global-positioning system (GPS)-collared black bears during peak and non-peak seasons of black bear-induced and brown bear-induced moose calf mortality within southwestern interior Alaska, in spring 2002. We compared habitats of GPS-collared black bears to those of presumably uncollared black bears and brown bears at their moose calf mortality sites. Results from this study suggest that GPS-collared black bears use similar habitat as conspecifics more than expected during the peak period of black bear predation on moose calves, whereas they use habitat in proportion to home range availability during the peak in brown bear predation on moose calves. Sex-specific Ivlev's electivity indices describe greater than expected use of mixed-deciduous forest and needleleaf forest by male GPS-collared black bears during the peak of moose calf predation, whereas females have a tendency to use these habitats less than expected. Juvenile GPS-collared black bears largely use the same habitat as other sympatric predators during the peak of moose calf predation, whereas during the non-peak period juveniles use opposite habitats as adult GPS-collared black bears. The outcome of this study offers possible explanations (e.g., sex, age) for spatial overlap or segregation in one member of a complex predator guild in relation to a seasonal pulse of preferred prey.  相似文献   

11.
In most of Europe, true wilderness areas do not exist and brown bears Ursus arctos generally have to cope with human disturbance and infrastructure. The few studies in Europe that have investigated brown bear activity have demonstrated a predominantly nocturnal and 'shy' behaviour in bears. There is still quite a debate on whether the shy, nocturnal bears of Europe are the result of centuries of persecution by men (genetically fixed trait) or whether hunting and the high disturbance potential in the multi-use landscapes are the driving force (individually learnt trait). We analysed the activity pattern of 16 individual bears monitored for 3372 h between May and October 1982–1998 in the Dinaric Mountains of Slovenia and Croatia. The data were collected via time sampling and basically analysed using two approaches: a general linear model with seasonal component to delineate the most important variables influencing the activity pattern and level and cluster analysis to group bears according to their 24-h activity pattern. Time of day and age were the most important variables predicting activity. Although individual variation in the activity pattern was high among individual bears, in general, yearlings were more diurnal and had a less distinct difference between day- and night-time activity levels than adult bears. Subadults were somewhat intermediate to adults and yearlings. We believe that nocturnal behaviour is most likely driven through negative experiences with humans, giving space for much individual variation. More research is needed to prove the causal relationship of nocturnal behaviour and the degree of disturbance that an individual bear is exposed to.  相似文献   

12.
ABSTRACT

This study investigated the effect animal activity had on information visitors attended to at three bear exhibits. Attention was used as an indirect measure of what information visitors are consciously processing, or learning. It was hypothesized that visitors would be more likely to attend to specific information about animal behavior when bears were active rather than when inactive or out of sight. Visitor conversations were recorded during observations of polar, sloth and spectacled bears. Conversation was coded according to its content (animal-directed, human-focused, behavior, other) and to the corresponding behavior of the bear (active, inactive, pacing, not visible). The energy level of the activity was also considered. The hypothesis was supported for the polar bears, but less so for the other two bears. Behavior conversation was highest and human-directed conversation was lowest in the presence of highly animated polar bears. Behavior content was limited in the presence of the less animated sloth and spectacled bears. However, it was still significantly less frequent, and human content more frequent, when the sloth and spectacled bears were pacing and/or not visible. Therefore, what visitors attended to was influenced by what the bears were doing. The findings suggest that animated activity (rather than the more broadly defined ‘activity’) elicits the most visitor attention to behavior, and this in turn potentially facilitates visitor learning.  相似文献   

13.
Since 1995, at least three polar bears (Ursus maritimus) have died in the area of Churchill (Manitoba, Canada) as a direct result of being suspended in a net during helicopter-assisted translocations. To assess and improve methods of suspending anesthetized polar bears, we conducted a study during November 1997 to determine the cardiopulmonary responses of eight captive polar bears to suspension by net and by sling. Each bear was anesthetized on two occasions in which the sequence of activities followed and the type of data collected was identical, with only the method of suspension differing. Control data obtained from 11 captive polar bears during 1995-96 was included in the statistical analyses of cardiopulmonary data to help clearly differentiate the cardiopulmonary effects of suspension from those of drug metabolism. Suspending polar bears above the ground by net caused acute hypertension (e.g., 17 to 49% increase in mean arterial pressure), possibly as a result of increased venous return due to body compression. Increased arousal (e.g., head, tongue, and limb movement) also occurred consistently during net-suspension and suggested a stress response. Surprisingly, most suspended bears showed little change in blood gas values, but at least one bear became hypoxemic (i.e., PaO2 < 60 mm Hg) with each method of suspension. Because of the potential health risks of hypertension and hypoxemia, we recommend modifying the method by which polar bears are suspended with the goal of reducing body compression.  相似文献   

14.
We compiled, summarized, and reviewed 269 incidents of bear–human conflict involving firearms that occurred in Alaska during 1883–2009. Encounters involving brown bears (Ursus arctos; 218 incidents, 81%), black bears (Ursus americanus; 30 incidents, 11%), polar bears (Ursus maritimus; 6 incidents, 2%), and 15 (6%) unidentified species provided insight into firearms success and failure. A total of 444 people and at least 367 bears were involved in these incidents. We found no significant difference in success rates (i.e., success being when the bear was stopped in its aggressive behavior) associated with long guns (76%) and handguns (84%). Moreover, firearm bearers suffered the same injury rates in close encounters with bears whether they used their firearms or not. Bears were killed in 61% (n = 162) of bear–firearms incidents. Additionally, we identified multiple reasons for firearms failing to stop an aggressive bear. Using logistic regression, the best model for predicting a successful outcome for firearm users included species and cohort of bear, human activity at time of encounter, whether or not the bear charged, and if fish or game meat was present. Firearm variables (e.g., type of gun, number of shots) were not useful in predicting outcomes in bear–firearms incidents. Although firearms have failed to protect some users, they are the only deterrent that can lethally stop an aggressive bear. Where firearms have failed to protect people, we identified contributing causes. Our findings suggest that only those proficient in firearms use should rely on them for protection in bear country. © 2012 The Wildlife Society.  相似文献   

15.
Abstract: We present a comprehensive look at a sample of bear spray incidents that occurred in Alaska, USA, from 1985 to 2006. We analyzed 83 bear spray incidents involving brown bears (Ursus arctos; 61 cases, 74%), black bears (Ursus americanus; 20 cases, 24%), and polar bears (Ursus maritimus; 2 cases, 2%). Of the 72 cases where persons sprayed bears to defend themselves, 50 (69%) involved brown bears, 20 (28%) black bears, and 2 (3%) polar bears. Red pepper spray stopped bears' undesirable behavior 92% of the time when used on brown bears, 90% for black bears, and 100% for polar bears. Of all persons carrying sprays, 98% were uninjured by bears in close-range encounters. All bear—inflicted injuries (n = 3) associated with defensive spraying involved brown bears and were relatively minor (i.e., no hospitalization required). In 7% (5 of 71) of bear spray incidents, wind was reported to have interfered with spray accuracy, although it reached the bear in all cases. In 14% (10 of 71) of bear spray incidents, users reported the spray having had negative side effects upon themselves, ranging from minor irritation (11%, 8 of 71) to near incapacitation (3%, 2 of 71). Bear spray represents an effective alternative to lethal force and should be considered as an option for personal safety for those recreating and working in bear country. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):640–645; 2008)  相似文献   

16.
Most animals concentrate their movement into certain hours of the day depending on drivers such as photoperiod, ambient temperature, inter‐ or intraspecific competition, and predation risk. The main activity periods of many mammal species, especially in human‐dominated landscapes, are commonly set at dusk, dawn, and during nighttime hours. Large carnivores, such as brown bears, often display great flexibility in diel movement patterns throughout their range, and even within populations, striking between individual differences in movement have been demonstrated. Here, we evaluated how seasonality and reproductive class affected diel movement patterns of brown bears of the Dinaric‐Pindos and Carpathian bear populations in Serbia. We analyzed the movement distances and general probability of movement of 13 brown bears (8 males and 5 females) equipped with GPS collars and monitored over 1–3 years. Our analyses revealed that movement distances and probability of bear movement differed between seasons (mating versus hyperphagia) and reproductive classes. Adult males, solitary females, and subadult males showed a crepuscular movement pattern. Compared with other reproductive classes, females with offspring were moving significantly less during crepuscular hours and during the night, particularly during the mating season, suggesting temporal niche partitioning among different reproductive classes. Adult males, solitary females, and in particular subadult males traveled greater hourly distances during the mating season in May‐June than the hyperphagia in July–October. Subadult males significantly decreased their movement from the mating season to hyperphagia, whereas females with offspring exhibited an opposite pattern with almost doubling their movement from the mating to hyperphagia season. Our results provide insights into how seasonality and reproductive class drive intrapopulation differences in movement distances and probability of movement in a recovering, to date little studied, brown bear population in southeastern Europe.  相似文献   

17.
After being extirpated from their historical distribution, black bears (Ursus americanus) have recolonized eastern Oklahoma, USA. As bears continue to establish populations in the region, understanding the cognitive factors that influence human acceptance of black bears will play an important role in facilitating coexistence. We hypothesized that a variety of variables drawing from the cognitive hierarchy and risk perception frameworks would affect human acceptance of black bears in eastern Oklahoma. We collected data using a mail-back questionnaire administered to residents of eastern Oklahoma between August and November 2018. Perceptions of the current black bear population size, attitudes toward bears, valuing Oklahoma wildlife, behavioral intentions, and level of formal education all positively influenced acceptance of bears. Risk-averse respondents and women were less accepting. Outreach to women and encouraging residents to reduce conflict by adopting bear-safe behaviors may help raise acceptance capacity for black bears. © 2021 The Wildlife Society.  相似文献   

18.
Conflicts between humans and wildlife have become increasingly important challenges for resource managers along the urban-wildland interface. Food conditioning (i.e., reliance by an animal on anthropogenic foods) of American black bears (Ursus americanus) is related to conflict behavior (i.e., being bold or aggressive toward humans, consuming human food or garbage, causing property damage) and often occurs in communities adjacent to Great Smoky Mountains National Park (GRSM or Park), USA. The goal of our study was to evaluate black bear space use in GRSM and in exurban areas on surrounding private lands and to identify factors associated with food conditioning and conflict behavior. We radio-collared 53 bears (29 males, 24 females) from 2015 to 2017 to compare space use characteristics and used carbon isotopic signatures (δ13C) from bear hair to assess food conditioning. We then performed an integrated step selection function (iSSF) analysis to characterize and compare movement and resource use as related to food conditioning. Based on the stable isotope analyses, 24 bears were classified as food conditioned (FC; 16 males and 8 females) and 37 were not food conditioned (NFC; 14 males and 23 females). Annual 95% kernel density estimate (KDE) home ranges and 50% KDE core area estimates of female and male bears did not differ by level of food conditioning (i.e., mean δ13C), but 95% and 50% home ranges of FC females were smaller than NFC females when data from 2015, a year of food scarcity and abnormally large home ranges, were excluded. The mean proportion of exurban development (e.g., roads, buildings, openings) within 95% KDE and 50% KDE home ranges of females increased with mean δ13C (i.e., greater food conditioning). The iSSF models indicated that FC bears were more likely to use forest openings associated with higher levels of development than NFC bears. We used those models to demonstrate how landscape modifications can reduce bear use of exurban areas, particularly for NFC bears. Our stable isotope, movement, and resource use data indicate that conflict behaviors displayed by many bears within GRSM were learned in areas outside Park boundaries. © 2020 The Wildlife Society.  相似文献   

19.
The welfare of captive bears became a big issue of concern in Poland when a case of a bear being ill-treated became a high-profile case in the media. This case created a challenge to verify, study, and understand the main problems associated with bear keeping so that zoos could significantly improve the conditions in which they keep bears or ensure they keep bears at the minimum required standards. The results presented here are from 1 of the few countrywide studies of captive bear conditions conducted in all the captive institutions in Poland that keep bears. Thirteen institutions kept bears at the time of the study (2007–2009), including 54 individuals of 5 species. Major welfare problems were identified, and the results have been used to challenge zoos to address the changes required and focus the government's attention on areas that require legislative improvement.  相似文献   

20.
Polar bears are an arctic, marine adapted species that is closely related to brown bears. Genome analyses have shown that polar bears are distinct and genetically homogeneous in comparison to brown bears. However, these analyses have also revealed a remarkable episode of polar bear gene flow into the population of brown bears that colonized the Admiralty, Baranof and Chichagof islands (ABC islands) of Alaska. Here, we present an analysis of data from a large panel of polar bear and brown bear genomes that includes brown bears from the ABC islands, the Alaskan mainland and Europe. Our results provide clear evidence that gene flow between the two species had a geographically wide impact, with polar bear DNA found within the genomes of brown bears living both on the ABC islands and in the Alaskan mainland. Intriguingly, while brown bear genomes contain up to 8.8% polar bear ancestry, polar bear genomes appear to be devoid of brown bear ancestry, suggesting the presence of a barrier to gene flow in that direction.  相似文献   

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