Genomic Evidence for Island Population Conversion Resolves Conflicting Theories of Polar Bear Evolution

Despite extensive genetic analysis, the evolutionary relationship between polar bears (Ursus maritimus) and brown bears (U. arctos) remains unclear. The two most recent comprehensive reports indicate a recent divergence with little subsequent admixture or a much more ancient divergence followed by extensive admixture. At the center of this controversy are the Alaskan ABC Islands brown bears that show evidence of shared ancestry with polar bears. We present an analysis of genome-wide sequence data for seven polar bears, one ABC Islands brown bear, one mainland Alaskan brown bear, and a black bear (U. americanus), plus recently published datasets from other bears. Surprisingly, we find clear evidence for gene flow from polar bears into ABC Islands brown bears but no evidence of gene flow from brown bears into polar bears. Importantly, while polar bears contributed <1% of the autosomal genome of the ABC Islands brown bear, they contributed 6.5% of the X chromosome. The magnitude of sex-biased polar bear ancestry and the clear direction of gene flow suggest a model wherein the enigmatic ABC Island brown bears are the descendants of a polar bear population that was gradually converted into brown bears via male-dominated brown bear admixture. We present a model that reconciles heretofore conflicting genetic observations. We posit that the enigmatic ABC Islands brown bears derive from a population of polar bears likely stranded by the receding ice at the end of the last glacial period. Since then, male brown bear migration onto the island has gradually converted these bears into an admixed population whose phenotype and genotype are principally brown bear, except at mtDNA and X-linked loci. This process of genome erosion and conversion may be a common outcome when climate change or other forces cause a population to become isolated and then overrun by species with which it can hybridize.     

Introgressive hybridization: brown bears as vectors for polar bear alleles

The dynamics and consequences of introgression can inform about numerous evolutionary processes. Biologists have therefore long been interested in hybridization. One challenge, however, lies in the identification of nonadmixed genotypes that can serve as a baseline for accurate quantification of admixture. In this issue of Molecular Ecology, Cahill et al. (2015) analyse a genomic data set of 28 polar bears, eight brown bears and one American black bear. Polar bear alleles are found to be introgressed into brown bears not only near a previously identified admixture zone on the Alaskan Admiralty, Baranof and Chichagof (ABC) Islands, but also far into the North American mainland. Elegantly contrasting admixture levels at autosomal and X chromosomal markers, Cahill and colleagues infer that male‐biased dispersal has spread these introgressed alleles away from the Late Pleistocene contact zone. Compared to a previous study on the ABC Island population in which an Alaskan brown bear served as a putatively admixture‐free reference, Cahill et al. (2015) utilize a newly sequenced Swedish brown bear as admixture baseline. This approach reveals that brown bears have been impacted by introgression from polar bears to a larger extent (up to 8.8% of their genome), than previously known, including the bear that had previously served as admixture baseline. No evidence for introgression of brown bear into polar bear is found, which the authors argue could be a consequence of selection. Besides adding new exciting pieces to the puzzle of polar/brown bear evolutionary history, the study by Cahill and colleagues highlights that wildlife genomics is moving from analysing single genomes towards a landscape genomics approach.     

Phylogeography of mitochondrial DNA variation in brown bears and polar bears

We analyzed 286 nucleotides of the middle portion of the mitochondrial cytochrome b gene of 61 brown bears from three locations in Alaska and 55 polar bears from Arctic Canada and Arctic Siberia to test our earlier observations of paraphyly between polar bears and brown bears as well as to test the extreme uniqueness of mitochondrial DNA types of brown bears on Admiralty, Baranof, and Chichagof (ABC) islands of southeastern Alaska. We also investigated the phylogeography of brown bears of Alaska's Kenai Peninsula in relation to other Alaskan brown bears because the former are being threatened by increased human development. We predicted that: (1) mtDNA paraphyly between brown bears and polar bears would be upheld, (2) the mtDNA uniqueness of brown bears of the ABC islands would be upheld, and (3) brown bears of the Kenai Peninsula would belong to either clade II or clade III of brown bears of our earlier studies of mtDNA. All of our predictions were upheld through the analysis of these additional samples.     

Genetics and conservation of European brown bears Ursus arctos

• 1 We review the genetics research that has been conducted on the European brown bear Ursus arctos, one of the genetically best‐studied mammalian species.
• 2 The first genetics studies on European brown bears were on phylogeography, as a basis for proposed population augmentations. Two major mitochondrial DNA lineages, western and eastern, and two clades within the western lineage were found. This led to a hypothesis that brown bears had contracted to southern refugia during the last glacial maximum. More recent results suggest that gene flow among brown bears blurred this structure and they survived north of these putative refugia. Thus, today's structure might be a result of population fragmentation caused by humans.
• 3 The nuclear diversity of European brown bears is similar in range to that in North American bears: low levels occur in the small populations and high levels in the large populations.
• 4 Many non‐invasive genetic methods, developed during research on brown bears, have been used for individual identification, censusing populations, monitoring migration and gene flow, and testing methods that are easier to use in endangered populations and over large areas.
• 5 Genetics has been used to study many behavioural and population ecological questions that have relevance for the conservation and management of brown bears.
• 6 The European brown bear has served, and will continue to serve, as a model for the development of methods, analyses and hypotheses in conservation genetics.

     

Gene flow between insular, coastal and interior populations of brown bears in Alaska

The brown bears of coastal Alaska have been recently regarded as comprising from one to three distinct genetic groups. We sampled brown bears from each of the regions for which hypotheses of genetic uniqueness have been made, including the bears of the Kodiak Archipelago and the bears of Admiralty, Baranof and Chichagof (ABC) Islands in southeast Alaska. These samples were analysed with a suite of nuclear microsatellite markers. The 'big brown bears' of coastal Alaska were found to be part of the continuous continental distribution of brown bears, and not genetically isolated from the physically smaller 'grizzly bears' of the interior. By contrast, Kodiak brown bears appear to have experienced little or no genetic exchange with continental populations in recent generations. The bears of the ABC Islands, which have previously been shown to undergo little or no female-mediated gene flow with mainland populations, were found not to be genetically isolated from mainland bears. The data from the four insular populations indicate that female and male dispersal can be reduced or eliminated by water barriers of 2–4 km and 7km in width, respectively.     

Bear-hybrids: behaviour and phenotype

The behaviour of a female and a male bear hybrid (brown bear Ursus arctos x polar bear Ursus maritimus) was studied. Behavioural and morphological comparisons between the hybrids and the two parent-species were made.The observation period covered July to November 2007. Different objects were offered to the bears to evoke new patterns of behaviour. While manipulating the offered objects, both bears showed elements of behaviour also found in polar bears, though the male performed those to a larger extend. Not only in the way of manipulating objects but also in the male's type of stereotype parallels to polar bears can be drawn. Phenotypically both bears possess features of both species, brown bear and polar bear. Concerning it lighter colour of fur and the structure of hair the female hybrid resembles more a polar bear than the male. Summarising, the female appears phenotypically more like a polar bear, the male with respect to the behaviour.     

Behavioral response of polar bears to aircraft activity on the northern coast of Alaska

The rapid loss of arctic sea ice is forcing a larger proportion of the Southern Beaufort Sea polar bear (Ursus maritimus) population to spend more time on land, increasing chances of negative interactions between people and bears. In the United States, the Marine Mammal Protection Act (MMPA) protects polar bears from incidental disturbance from human activities. For the remote and roadless areas of northern Alaska, USA, effective management of small aircraft activity is necessary to limit disturbance, but effects of overflights on polar bear behavior are largely unknown. During 2021 and 2022, we intentionally exposed polar bears (n = 115) to systematic aircraft activity (helicopter, fixed-wing) until we observed a disruption of behavior that qualified as a level B take response (e.g., abrupt change in activity or movement) under the MMPA. We used a Bayesian logistic regression to determine what factors influence and can be used to predict when a polar bear will exhibit a level B take response and estimate the probability of an aircraft eliciting a level B take response at different altitudes above the polar bear. Aircraft type, flight altitude, landscape (barrier islands vs. mainland), and bear behavior (active vs. inactive) upon initial aircraft encounter were all important predictors of take. Probability of take rapidly increased with a decrease in flight altitude starting at 450 m for helicopter and 300 m for fixed-wing aircraft. Active (e.g., standing, walking) polar bears on barrier-island landscapes were more likely to experience take than inactive (e.g., bedded) bears on mainland landscapes. Our findings can help with assessments and management plans by quantifying disturbance to polar bears from current and future human activity that involves aircraft use.     

Population substructure and space use of Foxe Basin polar bears

Climate change has been identified as a major driver of habitat change, particularly for sea ice-dependent species such as the polar bear (Ursus maritimus). Population structure and space use of polar bears have been challenging to quantify because of their circumpolar distribution and tendency to range over large areas. Knowledge of movement patterns, home range, and habitat is needed for conservation and management. This is the first study to examine the spatial ecology of polar bears in the Foxe Basin management unit of Nunavut, Canada. Foxe Basin is in the mid-Arctic, part of the seasonal sea ice ecoregion and it is being negatively affected by climate change. Our objectives were to examine intrapopulation spatial structure, to determine movement patterns, and to consider how polar bear movements may respond to changing sea ice habitat conditions. Hierarchical and fuzzy cluster analyses were used to assess intrapopulation spatial structure of geographic position system satellite-collared female polar bears. Seasonal and annual movement metrics (home range, movement rates, time on ice) and home-range fidelity (static and dynamic overlap) were compared to examine the influence of regional sea ice on movements. The polar bears were distributed in three spatial clusters, and there were differences in the movement metrics between clusters that may reflect sea ice habitat conditions. Within the clusters, bears moved independently of each other. Annual and seasonal home-range fidelity was observed, and the bears used two movement patterns: on-ice range residency and annual migration. We predict that home-range fidelity may decline as the spatial and temporal predictability of sea ice changes. These new findings also provide baseline information for managing and monitoring this polar bear population.     

Seasonal and diel movement patterns of brown bears in a population in southeastern Europe

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.     

Efficacy of Bear Deterrent Spray in Alaska

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)     

Phylogeny of the bears (Ursidae) based on nuclear and mitochondrial genes

The taxomic classification and phylogenetic relationships within the bear family remain argumentative subjects in recent years. Prior investigation has been concentrated on the application of different mitochondrial (mt) sequence data, herein we employ two nuclear single-copy gene segments, the partial exon 1 from gene encoding interphotoreceptor retinoid binding protein (IRBP) and the complete intron 1 from transthyretin (TTR) gene, in conjunction with previously published mt data, to clarify these enigmatic problems. The combined analyses of nuclear IRBP and TTR datasets not only corroborated prior hypotheses, positioning the spectacled bear most basally and grouping the brown and polar bear together but also provided new insights into the bear phylogeny, suggesting the sister-taxa association of sloth bear and sun bear with strong support. Analyses based on combination of nuclear and mt genes differed from nuclear analysis in recognizing the sloth bears as the earliest diverging species among the subfamily ursine representatives while the exact placement of the sun bear did not resolved. Asiatic and American black bears clustered as sister group in all analyses with moderate levels of bootstrap support and high posterior probabilities. Comparisons between the nuclear and mtDNA findings suggested that our combined nuclear dataset have the resolving power comparable to mtDNA dataset for the phylogenetic interpretation of the bear family. As can be seen from present study, the unanimous phylogeny for this recently derived family was still not produced and additional independent genetic markers were in need.     

Ancient DNA reveals differences in behaviour and sociality between brown bears and extinct cave bears

Ancient DNA studies have revolutionized the study of extinct species and populations, providing insights on phylogeny, phylogeography, admixture and demographic history. However, inferences on behaviour and sociality have been far less frequent. Here, we investigate the complete mitochondrial genomes of extinct Late Pleistocene cave bears and middle Holocene brown bears that each inhabited multiple geographically proximate caves in northern Spain. In cave bears, we find that, although most caves were occupied simultaneously, each cave almost exclusively contains a unique lineage of closely related haplotypes. This remarkable pattern suggests extreme fidelity to their birth site in cave bears, best described as homing behaviour, and that cave bears formed stable maternal social groups at least for hibernation. In contrast, brown bears do not show any strong association of mitochondrial lineage and cave, suggesting that these two closely related species differed in aspects of their behaviour and sociality. This difference is likely to have contributed to cave bear extinction, which occurred at a time in which competition for caves between bears and humans was likely intense and the ability to rapidly colonize new hibernation sites would have been crucial for the survival of a species so dependent on caves for hibernation as cave bears. Our study demonstrates the potential of ancient DNA to uncover patterns of behaviour and sociality in ancient species and populations, even those that went extinct many tens of thousands of years ago.     

Dietary adjustability of grizzly bears and American black bears in Yellowstone National Park

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.     

Mercury in polar bears from Alaska

Alaskan polar bear (Ursus maritimus) muscle and liver samples collected in 1972 were analyzed for total mercury. Bears north of Alaska had more mercury than bears west of Alaska. The only difference between young and adult animals was in the northern area where adults had more mercury in liver tissue than young animals. Levels were probably not high enough to be a serious threat to bears.     

Implications of the Circumpolar Genetic Structure of Polar Bears for Their Conservation in a Rapidly Warming Arctic

We provide an expansive analysis of polar bear (Ursus maritimus) circumpolar genetic variation during the last two decades of decline in their sea-ice habitat. We sought to evaluate whether their genetic diversity and structure have changed over this period of habitat decline, how their current genetic patterns compare with past patterns, and how genetic demography changed with ancient fluctuations in climate. Characterizing their circumpolar genetic structure using microsatellite data, we defined four clusters that largely correspond to current ecological and oceanographic factors: Eastern Polar Basin, Western Polar Basin, Canadian Archipelago and Southern Canada. We document evidence for recent (ca. last 1–3 generations) directional gene flow from Southern Canada and the Eastern Polar Basin towards the Canadian Archipelago, an area hypothesized to be a future refugium for polar bears as climate-induced habitat decline continues. Our data provide empirical evidence in support of this hypothesis. The direction of current gene flow differs from earlier patterns of gene flow in the Holocene. From analyses of mitochondrial DNA, the Canadian Archipelago cluster and the Barents Sea subpopulation within the Eastern Polar Basin cluster did not show signals of population expansion, suggesting these areas may have served also as past interglacial refugia. Mismatch analyses of mitochondrial DNA data from polar and the paraphyletic brown bear (U. arctos) uncovered offset signals in timing of population expansion between the two species, that are attributed to differential demographic responses to past climate cycling. Mitogenomic structure of polar bears was shallow and developed recently, in contrast to the multiple clades of brown bears. We found no genetic signatures of recent hybridization between the species in our large, circumpolar sample, suggesting that recently observed hybrids represent localized events. Documenting changes in subpopulation connectivity will allow polar nations to proactively adjust conservation actions to continuing decline in sea-ice habitat.     

Phylogeography and mitochondrial diversity of extirpated brown bear (Ursus arctos) populations in the contiguous United States and Mexico

The fossil record indicates that the brown bear (Ursus arctos) colonized North America from Asia over 50 000 years ago. The species historically occupied the western United States and northern Mexico but has been extirpated from over 99% of this range in the last two centuries. To evaluate colonization hypotheses, subspecific classifications, and historical patterns and levels of genetic diversity in this region, we sequenced 229 nucleotides of the mitochondrial DNA control region in 108 museum specimens. The work was set in a global context by synthesizing all previous brown bear control region sequences from around the world. In mid-latitude North America a single moderately diverse clade is observed, represented by 23 haplotypes with up to 3.5% divergence. Only eight of 23 haplotypes (35%) are observed in the extensively sampled extant populations suggesting a substantial loss of genetic variability. The restriction of all haplotypes from mid-latitude North America to a single clade suggests that this region was founded by bears with a similar maternal ancestry. However, the levels and distributions of diversity also suggest that the colonizing population was not a small founder event, and that expansion occurred long enough ago for local mutations to accrue. Our data are consistent with recent genetic evidence that brown bears were south of the ice prior to the last glacial maximum. There is no support for previous subspecies designations, although bears of the southwestern United States may have had a distinctive, but recent, pattern of ancestry.     

Factors associated with fecal glucocorticoids in Alaskan brown bears (Ursus arctos horribilis)

The aims of this study were to validate a radioimmunoassay (RIA) for quantifying glucocorticoid metabolite concentrations in the feces of Alaskan brown bears (Ursus arctos horribilis) and to investigate whether any of the following factors are associated with those concentrations: the presence of humans or other bears, fishing difficulty, sex-age class, diet, and season. We tested an established corticosterone RIA for assay sensitivity, similarity, precision, and sample matrix effects of brown bear feces, and it proved satisfactory. We collected fecal samples from brown bears along salmon-spawning streams and assessed fecal glucocorticoid (FG) concentrations. We observed that the factors explaining the most variation in measured concentrations were date and diet type and that there was a significant interaction between the two. We did not observe a significant effect of human and bear activities or sex-age class on FG concentrations. This study demonstrates that although FG concentrations may be assessed in brown bears, complex dietary patterns and seasonal variations must be taken into consideration in the study design in order to make inferences regarding stress.     

Predation of Svalbard reindeer by polar bears

Polar bears (Ursus maritimus) are believed to be obligate predators on marine mammals, and particularly, on two species of seals. This paper reports on observations of polar bears preying (n=7) and scavenging (n=6) on Svalbard reindeer (Rangifer tarandus platyrhyncus). Similar to their closest evolutionary ancestor, the brown bear (U. arctos), polar bears are opportunistic and will prey on ungulates. Reindeer are likely of minor importance to the foraging ecology of polar bears in Svalbard, but the observations suggest behavioural plasticity in response to a novel prey item. Accepted: 26 March 2000     

Staying out in the cold: glacial refugia and mitochondrial DNA phylogeography in ancient European brown bears

Models for the development of species distribution in Europe typically invoke restriction in three temperate Mediterranean refugia during glaciations, from where recolonization of central and northern Europe occurred. The brown bear, Ursus arctos, is one of the taxa from which this model is derived. Sequence data generated from brown bear fossils show a complex phylogeographical history for western European populations. Long-term isolation in separate refugia is not required to explain our data when considering the palaeontological distribution of brown bears. We propose continuous gene flow across southern Europe, from which brown bear populations expanded after the last glaciation.