Publications by authors named "Jon E Swenson"

85 Publications

Specific shifts in the endocannabinoid system in hibernating brown bears.

Front Zool 2020 Nov 23;17(1):35. Epub 2020 Nov 23.

Université Clermont Auvergne, INRAE, UNH, Clermont-Ferrand, France.

In small hibernators, global downregulation of the endocannabinoid system (ECS), which is involved in modulating neuronal signaling, feeding behavior, energy metabolism, and circannual rhythms, has been reported to possibly drive physiological adaptation to the hibernating state. In hibernating brown bears (Ursus arctos), we hypothesized that beyond an overall suppression of the ECS, seasonal shift in endocannabinoids compounds could be linked to bear's peculiar features that include hibernation without arousal episodes and capacity to react to external disturbance. We explored circulating lipids in serum and the ECS in plasma and metabolically active tissues in free-ranging subadult Scandinavian brown bears when both active and hibernating. In winter bear serum, in addition to a 2-fold increase in total fatty acid concentration, we found significant changes in relative proportions of circulating fatty acids, such as a 2-fold increase in docosahexaenoic acid C22:6 n-3 and a decrease in arachidonic acid C20:4 n-6. In adipose and muscle tissues of hibernating bears, we found significant lower concentrations of 2-arachidonoylglycerol (2-AG), a major ligand of cannabinoid receptors 1 (CB1) and 2 (CB2). Lower mRNA level for genes encoding CB1 and CB2 were also found in winter muscle and adipose tissue, respectively. The observed reduction in ECS tone may promote fatty acid mobilization from body fat stores, and favor carbohydrate metabolism in skeletal muscle of hibernating bears. Additionally, high circulating level of the endocannabinoid-like compound N-oleoylethanolamide (OEA) in winter could favor lipolysis and fatty acid oxidation in peripheral tissues. We also speculated on a role of OEA in the conservation of an anorexigenic signal and in the maintenance of torpor during hibernation, while sustaining the capacity of bears to sense stimuli from the environment.
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http://dx.doi.org/10.1186/s12983-020-00380-yDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7681968PMC
November 2020

Disentangling direct and indirect determinants of the duration of maternal care in brown bears: Environmental context matters.

J Anim Ecol 2021 02 27;90(2):376-386. Epub 2020 Oct 27.

Département de biologie & Centre for Northern Studies, Université de Sherbrooke, Sherbrooke, QC, Canada.

The duration of maternal care, an important life-history trait affecting population dynamics, varies greatly within species. Yet, our understanding of its predictors is limited, mostly correlative and subject to misinterpretations, due to difficulties to disentangle the role of maternal- and offspring-related characteristics. We conducted path analysis on a dataset including 217 brown bear litters captured over a 29-year period in two populations in Sweden ('North' and 'South') facing contrasting environmental conditions to identify and quantify the causes of variation in the duration of maternal care (1.5 or 2.5 years). We showed that the causal determinants of the duration of maternal care were context-dependent. Contrary to their expected central role in the determination of the duration of maternal care, yearling mass and its direct determinants (i.e. litter size and maternal mass) were only important in the North population, where environmental conditions are harsher and the cost of extended maternal care presumably higher. In the South, the duration of maternal care was not caused by yearling mass nor any maternal or litter characteristics. Extension of maternal care may thus result from factors independent from maternal and offspring condition in the South, such as an artificial hunting-induced selection for longer maternal care through the legal protection of family groups. Our results provide an important contribution to our very limited knowledge of the direct and indirect determinants of the duration of maternal care and highlight the importance of accounting for the environmental context when assessing maternal reproductive tactics.
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http://dx.doi.org/10.1111/1365-2656.13371DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7894530PMC
February 2021

Room without a view-Den excavation in relation to body size in brown bears.

Ecol Evol 2020 Aug 20;10(15):8044-8054. Epub 2020 Jul 20.

Department of Natural Sciences and Environmental Health University of South-Eastern Norway Telemark Norway.

Hibernation is an adaptive strategy to survive harsh winter conditions and food shortage. The use of well-insulated winter dens helps animals minimize energy loss during hibernation. Brown bears () commonly use excavated dens for hibernation. Physical attributes of excavated dens are expected to impact the bear's heat retention and energy conservation. The objective of this study was to examine the determinants of cavity size of excavated dens and the impact of physical attributes of excavated dens on energy conservation in hibernating bears, hypothesizing that bears excavate dens in a way to minimize heat loss and optimize energy conservation during hibernation. We predicted that den cavity size would be determined by the bear's body size and that older bears would excavate better-fitting cavities to minimize heat loss, due to their previous experience. We further predicted that physical attributes of excavated dens would affect the bears' posthibernation body condition. Our results revealed that bears excavated a den cavity in relation to their body size, regardless of sex, and that older bears tended to excavate better-fitting den cavities compared to young bears, as we expected. Older bears excavated better-fitting den cavities, suggesting a potentially experience-based shift with age in den-excavation behavior and an optimum cavity size relative to a bear's body size. Our key finding is that insulation of excavated dens provided by wall/rood thickness and bedding materials had a significant positive effect on bears' posthibernation body condition. We believe that our study provides new insight into how not only the quality of denning habitat, but also the quality of dens may affect hibernating animals, by presenting a potential adaptive aspect of den preparation (age effect on efficiency in den excavation) and effect of den attributes on the posthibernation body condition of brown bears.
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http://dx.doi.org/10.1002/ece3.6371DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7417226PMC
August 2020

Wolf habitat selection when sympatric or allopatric with brown bears in Scandinavia.

Sci Rep 2020 06 18;10(1):9941. Epub 2020 Jun 18.

Grimsӧ Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, SE-730 91, Riddarhyttan, Sweden.

Habitat selection of animals depends on factors such as food availability, landscape features, and intra- and interspecific interactions. Individuals can show several behavioral responses to reduce competition for habitat, yet the mechanisms that drive them are poorly understood. This is particularly true for large carnivores, whose fine-scale monitoring is logistically complex and expensive. In Scandinavia, the home-range establishment and kill rates of gray wolves (Canis lupus) are affected by the coexistence with brown bears (Ursus arctos). Here, we applied resource selection functions and a multivariate approach to compare wolf habitat selection within home ranges of wolves that were either sympatric or allopatric with bears. Wolves selected for lower altitudes in winter, particularly in the area where bears and wolves are sympatric, where altitude is generally higher than where they are allopatric. Wolves may follow the winter migration of their staple prey, moose (Alces alces), to lower altitudes. Otherwise, we did not find any effect of bear presence on wolf habitat selection, in contrast with our previous studies. Our new results indicate that the manifestation of a specific driver of habitat selection, namely interspecific competition, can vary at different spatial-temporal scales. This is important to understand the structure of ecological communities and the varying mechanisms underlying interspecific interactions.
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http://dx.doi.org/10.1038/s41598-020-66626-1DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7303184PMC
June 2020

Do follicles matter? Testing the effect of follicles on hair cortisol levels.

Conserv Physiol 2020 1;8(1):coaa003. Epub 2020 Feb 1.

Department of Natural Sciences and Environmental Health, University of South-Eastern Norway, Gullbringvegen 36, 3800 Bø, Norway.

Cortisol concentrations in hair are used increasingly as a biomarker of long-term stress in free-ranging wildlife. Cortisol is believed to be integrated into hair primarily during its active growth phase, typically occurring over weeks to months or longer periods, depending on latitude. Cortisol concentrations in hair thus reflect the activity of the hypothalamic-pituitary-adrenal axis over this time. However, local, independent cortisol secretion within the skin, which includes hair follicles, may also contribute to cortisol levels in growing hair. Methodological differences between studies include the measurement of cortisol in only the hair shaft (i.e. follicle absent, as with shaved hair) versus the whole hair (i.e. follicle present, as with plucked hair). If the concentration of cortisol in the follicle is high enough to influence the overall hair cortisol concentration (HCC), this could confound comparisons between studies using different types of hair samples (hair shafts vs. whole hair) and collection methods. Here, we test the hypothesis that cortisol present in follicles influences HCC. We compared HCC in paired subsamples of hair with and without follicles from 30 free-ranging Scandinavian brown bears () and observed significantly greater HCC in samples with follicles present. The effect of follicles remained significant also with sex and age of sampled bears taken into account in a linear mixed model. Finally, we provide an overview of collection methods and types of hair samples used for HCC analysis in 77 studies dealing with stress in wild mammal species. Our findings highlight the need to unify methods of hair collection and preparation to allow for valid comparisons, and to optimize labour input in ecophysiological studies.
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http://dx.doi.org/10.1093/conphys/coaa003DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6994724PMC
February 2020

Diet of the brown bear in Himalaya: Combining classical and molecular genetic techniques.

PLoS One 2019 26;14(12):e0225698. Epub 2019 Dec 26.

Department of Ecology and Natural Resource Management, Norwegian University of Life Sciences, Norway.

The ecological requirements of brown bears are poorly known in the Himalaya region, which complicates conservation efforts. We documented the diet of the Himalayan brown bear (Ursus arctos isabellinus) by combining classical scat analysis and a newly developed molecular genetic technique (the trnL approach), in Deosai National Park, Pakistan. Brown bears consumed over 50 plant species, invertebrates, ungulates, and several rodents. Eight plant families; Poaceae, Polygonaceae, Cyperaceae, Apiaceae, Asteraceae, Caryophyllaceae, Lamiaceae, and Rubiaceae were commonly eaten with graminoids comprising the bulk of the diet. Golden marmots comprised the major mammalian biomass in the park, and were also the main meat source for bears. Animal matter, making 36% of dietary content, contributed half of the digestible energy, due to its higher nutritious value. We did not find a significant temporal pattern in diet, perhaps because the availability of the major diet (graminoids) did not change over the foraging period. Male brown bears were more carnivorous than females, probably because of their larger size, which requires higher energy and also makes them more efficient in capturing marmots. Frequencies of three plant species were also significantly higher in male brown bears; Bistorta affinis, Carex diluta, and Carex sp. Diet of the brown bear differed significantly between the park and surrounding valleys. In valleys, diet consisted predominantly of graminoids and crops, whereas the park provided more nutritious and diverse foodThe estimated digestible energy available to brown bears in Deosai was the lowest documented among brown bear populations, due to the lack of fruits and a relatively lower meat content. The low nutritious diet and high cost of metabolism in a high-altitude environment, probably explains the very low reproductive potential of this population.
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http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0225698PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6932756PMC
April 2020

MicroRNAs facilitate skeletal muscle maintenance and metabolic suppression in hibernating brown bears.

J Cell Physiol 2020 04 23;235(4):3984-3993. Epub 2019 Oct 23.

Department of Biology, Carleton University, Ottawa, Canada.

Hibernating brown bears, Ursus arctos, undergo extended periods of inactivity and yet these large hibernators are resilient to muscle disuse atrophy. Physiological characteristics associated with atrophy resistance in bear muscle have been examined (e.g., muscle mechanics, neural activity) but roles for molecular signaling/regulatory mechanisms in the resistance to muscle wasting in bears still require investigation. Using quantitative reverse transcription PCR (RT-qPCR), the present study characterized the responses of 36 microRNAs linked with development, metabolism, and regeneration of skeletal muscle, in the vastus lateralis of brown bears comparing winter hibernating and summer active animals. Relative levels of mRNA of selected genes (mef2a, pax7, id2, prkaa1, and mstn) implicated upstream and downstream of the microRNAs were examined. Results indicated that hibernation elicited a myogenic microRNA, or "myomiR", response via MEF2A-mediated signaling. Upregulation of MEF2A-controlled miR-1 and miR-206 and respective downregulation of pax7 and id2 mRNA are suggestive of responses that promote skeletal muscle maintenance. Increased levels of metabolic microRNAs, such as miR-27, miR-29, and miR-33, may facilitate metabolic suppression during hibernation via mechanisms that decrease glucose uptake and fatty acid oxidation. This study identified myomiR-mediated mechanisms for the promotion of muscle regeneration, suppression of ubiquitin ligases, and resistance to muscle atrophy during hibernation mediated by observed increases in miR-206, miR-221, miR-31, miR-23a, and miR-29b. This was further supported by the downregulation of myomiRs associated with a muscle injury and inflammation (miR-199a and miR-223) during hibernation. The present study provides evidence of myomiR-mediated signaling pathways that are activated during hibernation to maintain skeletal muscle functionality in brown bears.
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http://dx.doi.org/10.1002/jcp.29294DOI Listing
April 2020

Limited Oxidative Stress Favors Resistance to Skeletal Muscle Atrophy in Hibernating Brown Bears ().

Antioxidants (Basel) 2019 Aug 22;8(9). Epub 2019 Aug 22.

Université de Strasbourg, CNRS, IPHC UMR 7178, F-670000 Strasbourg, France.

Oxidative stress, which is believed to promote muscle atrophy, has been reported to occur in a few hibernators. However, hibernating bears exhibit efficient energy savings and muscle protein sparing, despite long-term physical inactivity and fasting. We hypothesized that the regulation of the oxidant/antioxidant balance and oxidative stress could favor skeletal muscle maintenance in hibernating brown bears. We showed that increased expressions of cold-inducible proteins CIRBP and RBM3 could favor muscle mass maintenance and alleviate oxidative stress during hibernation. Downregulation of the subunits of the mitochondrial electron transfer chain complexes I, II, and III, and antioxidant enzymes, possibly due to the reduced mitochondrial content, indicated a possible reduction of the production of reactive oxygen species in the hibernating muscle. Concomitantly, the upregulation of cytosolic antioxidant systems, under the control of the transcription factor NRF2, and the maintenance of the GSH/GSSG ratio suggested that bear skeletal muscle is not under a significant oxidative insult during hibernation. Accordingly, lower levels of oxidative damage were recorded in hibernating bear skeletal muscles. These results identify mechanisms by which limited oxidative stress may underlie the resistance to skeletal muscle atrophy in hibernating brown bears. They may constitute therapeutic targets for the treatment of human muscle atrophy.
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http://dx.doi.org/10.3390/antiox8090334DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6770786PMC
August 2019

Phylogeography, genetic diversity, and connectivity of brown bear populations in Central Asia.

PLoS One 2019 13;14(8):e0220746. Epub 2019 Aug 13.

Department of Fish and Wildlife Sciences, University of Idaho, Moscow, Idaho, United States of America.

Knowledge of genetic diversity and population structure is critical for conservation and management planning at the population level within a species' range. Many brown bear populations in Central Asia are small and geographically isolated, yet their phylogeographic relationships, genetic diversity, and contemporary connectivity are poorly understood. To address this knowledge gap, we collected brown bear samples from the Gobi Desert (n = 2360), Altai, Sayan, Khentii, and Ikh Khyangan mountains of Mongolia (n = 79), and Deosai National Park in the Himalayan Mountain Range of Pakistan (n = 5) and generated 927 base pairs of mitochondrial DNA (mtDNA) sequence data and genotypes at 13 nuclear DNA microsatellite loci. We documented high levels of mtDNA and nDNA diversity in the brown bear populations of northern Mongolia (Altai, Sayan, Buteeliin nuruu and Khentii), but substantially lower diversity in brown bear populations in the Gobi Desert and Himalayas of Pakistan. We detected 3 brown bear mtDNA phylogeographic groups among bears of the region, with clade 3a1 in Sayan, Khentii, and Buteeliin nuruu mountains, clade 3b in Altai, Sayan, Buteeliin nuruu, Khentii, and Ikh Khyangan, and clade 6 in Gobi and Pakistan. Our results also clarified the phylogenetic relationships and divergence times with other brown bear mtDNA clades around the world. The nDNA genetic structure analyses revealed distinctiveness of Gobi bears and different population subdivisions compared to mtDNA results. For example, genetic distance for nDNA microsatellite loci between the bears in Gobi and Altai (FST = 0.147) was less than that of the Gobi and Pakistan (FST = 0.308) suggesting more recent male-mediated nuclear gene flow between Gobi and Altai than between Gobi and the Pakistan bears. Our results provide valuable information for conservation and management of bears in this understudied region of Central Asia and highlight the need for special protection and additional research on Gobi brown bears.
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http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0220746PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6692007PMC
February 2020

Heritability of head size in a hunted large carnivore, the brown bear ().

Evol Appl 2019 Jun 21;12(6):1124-1135. Epub 2019 Mar 21.

Department of Natural Sciences and Environmental Health, Faculty of Technology, Natural Sciences and Maritime Sciences University of South-Eastern Norway Bø i Telemark Norway.

Wild animal populations experience selection pressures from both natural and anthropogenic sources. The availability of extensive pedigrees is increasing along with our ability to quantify the heritability and evolvability of phenotypic traits and thus the speed and potential for evolutionary change in wild populations. The environment may also affect gene expressions in individuals, which may in turn affect the potential of phenotypic traits to respond to selection. Knowledge about the relationship between the genetic and environmental components of phenotypic variation is particularly relevant, given ongoing anthropogenically driven global change. Using a quantitative genetic mixed model, we disentangled the genetic and environmental components of phenotypic variance in a large carnivore, the brown bear (). We combined a pedigree covering ~1,500 individual bears over seven generations with location data from 413 bears, as well as data on bear density, habitat characteristics, and climatic conditions. We found a narrow-sense heritability of 0.24 (95% CrI: 0.06-0.38) for brown bear head size, showing that the trait can respond to selection at a moderate speed. The environment contributed substantially to phenotypic variation, and we partitioned this into birth year (5.9%), nonadditive among-individual genetic (15.0%), and residual (50.4%) environmental effects. Brown bear head circumference showed an evolvability of 0.2%, which can generate large changes in the trait mean over some hundreds of generations. Our study is among the first to quantify heritability of a trait in a hunted large carnivore population. Such knowledge about the degree to which species experiencing hunting can respond to selection is crucial for conservation and to make informed management decisions. We show that including important environmental variables when analyzing heritability is key to understanding the dynamics of the evolutionary potential of phenotypic traits.
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http://dx.doi.org/10.1111/eva.12786DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6597896PMC
June 2019

Metabolic reprogramming involving glycolysis in the hibernating brown bear skeletal muscle.

Front Zool 2019 6;16:12. Epub 2019 May 6.

1Université de Strasbourg, CNRS, IPHC UMR 7178, F-67000 Strasbourg, France.

Background: In mammals, the hibernating state is characterized by biochemical adjustments, which include metabolic rate depression and a shift in the primary fuel oxidized from carbohydrates to lipids. A number of studies of hibernating species report an upregulation of the levels and/or activity of lipid oxidizing enzymes in muscles during torpor, with a concomitant downregulation for glycolytic enzymes. However, other studies provide contrasting data about the regulation of fuel utilization in skeletal muscles during hibernation. Bears hibernate with only moderate hypothermia but with a drop in metabolic rate down to ~ 25% of basal metabolism. To gain insights into how fuel metabolism is regulated in hibernating bear skeletal muscles, we examined the vastus lateralis proteome and other changes elicited in brown bears during hibernation.

Results: We show that bear muscle metabolic reorganization is in line with a suppression of ATP turnover. Regulation of muscle enzyme expression and activity, as well as of circulating metabolite profiles, highlighted a preference for lipid substrates during hibernation, although the data suggested that muscular lipid oxidation levels decreased due to metabolic rate depression. Our data also supported maintenance of muscle glycolysis that could be fuelled from liver gluconeogenesis and mobilization of muscle glycogen stores. During hibernation, our data also suggest that carbohydrate metabolism in bear muscle, as well as protein sparing, could be controlled, in part, by actions of n-3 polyunsaturated fatty acids like docosahexaenoic acid.

Conclusions: Our work shows that molecular mechanisms in hibernating bear skeletal muscle, which appear consistent with a hypometabolic state, likely contribute to energy and protein savings. Maintenance of glycolysis could help to sustain muscle functionality for situations such as an unexpected exit from hibernation that would require a rapid increase in ATP production for muscle contraction. The molecular data we report here for skeletal muscles of bears hibernating at near normal body temperature represent a signature of muscle preservation despite atrophying conditions.
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http://dx.doi.org/10.1186/s12983-019-0312-2DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6503430PMC
May 2019

Lipidomics Reveals Seasonal Shifts in a Large-Bodied Hibernator, the Brown Bear.

Front Physiol 2019 12;10:389. Epub 2019 Apr 12.

IPHC, University of Strasbourg, Strasbourg, France.

Prior to winter, heterotherms retain polyunsaturated fatty acids ("PUFA"), resulting in enhanced energy savings during hibernation, through deeper and longer torpor bouts. Hibernating bears exhibit a less dramatic reduction (2-5°C) in body temperature, but lower their metabolism to a degree close to that of small hibernators. We determined the lipid composition, via lipidomics, in skeletal muscle and white adipose tissues ("WAT"), to assess lipid retention, and in blood plasma, to reflect lipid trafficking, of winter hibernating and summer active wild Scandinavian brown bears (). We found that the proportion of monounsaturated fatty acids in muscle of bears was significantly higher during winter. During hibernation, omega-3 PUFAs were retained in WAT and short-length fatty acids were released into the plasma. The analysis of individual lipid moieties indicated significant changes of specific fatty acids, which are in line with the observed seasonal shift in the major lipid categories and can be involved in specific regulations of metabolisms. These results strongly suggest that the shift in lipid composition is well conserved among hibernators, independent of body mass and of the animals' body temperature.
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http://dx.doi.org/10.3389/fphys.2019.00389DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6474398PMC
April 2019

No place like home? A test of the natal habitat-biased dispersal hypothesis in Scandinavian wolves.

R Soc Open Sci 2018 Dec 12;5(12):181379. Epub 2018 Dec 12.

Faculty of Applied Ecology and Agricultural Sciences, Inland Norway University of Applied Sciences, Evenstad, 2480 Koppang, Norway.

Natal dispersal is an important mechanism for the viability of populations. The influence of local conditions or experience gained in the natal habitat could improve fitness if dispersing individuals settle in an area with similar habitat characteristics. This process, defined as 'natal habitat-biased dispersal' (NHBD), has been used to explain distribution patterns in large carnivores, but actual studies evaluating it are rare. We tested whether grey wolf territory establishment was influenced by the habitat characteristics of the natal territory using the long-term monitoring of the Scandinavian wolf population. We paired the locations of natal and established territories, accounted for available habitats along the dispersing route, and compared their habitat characteristics for 271 wolves during 1998-2012. Wolves with the shortest dispersal distances established in natal-like habitat types more than expected by chance, whereas wolves that dispersed longer distances did not show NHBD. The pattern was consistent for male and female wolves, with females showing more NHBD than males. Chances to detect NHBD increased with the size of habitat defined as available. This highlights the importance of considering the biological characteristics of the studied species when defining habitat availability. Our methodological approach can prove useful to inform conservation and management to identify habitats to be selected by reintroduced or naturally expanding populations.
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http://dx.doi.org/10.1098/rsos.181379DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6304128PMC
December 2018

Habitat segregation between brown bears and gray wolves in a human-dominated landscape.

Ecol Evol 2018 Dec 11;8(23):11450-11466. Epub 2018 Nov 11.

Grimsӧ Wildlife Research Station Department of Ecology Swedish University of Agricultural Sciences Riddarhyttan Sweden.

Identifying how sympatric species belonging to the same guild coexist is a major question of community ecology and conservation. Habitat segregation between two species might help reduce the effects of interspecific competition and apex predators are of special interest in this context, because their interactions can have consequences for lower trophic levels. However, habitat segregation between sympatric large carnivores has seldom been studied. Based on monitoring of 53 brown bears () and seven sympatric adult gray wolves () equipped with GPS collars in Sweden, we analyzed the degree of interspecific segregation in habitat selection within their home ranges in both late winter and spring, when their diets overlap the most. We used the K-select method, a multivariate approach that relies on the concept of ecological niche, and randomization methods to quantify habitat segregation between bears and wolves. Habitat segregation between bears and wolves was greater than expected by chance. Wolves tended to select for moose occurrence, young forests, and rugged terrain more than bears, which likely reflects the different requirements of an omnivore (bear) and an obligate carnivore (wolf). However, both species generally avoided human-related habitats during daytime. Disentangling the mechanisms that can drive interspecific interactions at different spatial scales is essential for understanding how sympatric large carnivores occur and coexist in human-dominated landscapes, and how coexistence may affect lower trophic levels. The individual variation in habitat selection detected in our study may be a relevant mechanism to overcome intraguild competition and facilitate coexistence.
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http://dx.doi.org/10.1002/ece3.4572DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6303696PMC
December 2018

Long-Term Safety of Intraperitoneal Radio Transmitter Implants in Brown Bears ().

Front Vet Sci 2018 15;5:252. Epub 2018 Oct 15.

Norwegian Institute for Nature Research, Trondheim, Norway.

Intraperitoneal radio transmitters have been widely used in free-ranging wild mammals, but there are no long-term studies on their biocompatibility or technical stability within the abdominal cavity of animals. Possible negative health effects may bias results from ecological studies on instrumented animals and raise concerns over animal welfare issues. The aim of this study was to evaluate the long-term technical stability and pathological effects of Telonics intraperitoneal very high frequency (VHF) radio transmitters in brown bears (). We instrumented 305 individual bears with intraperitoneal VHF radio transmitters during a 19-year period. We surgically removed devices that had been in bears for 1-9 years and collected transmitters from animals that died 1-13 years after implantation. We took biopsies for histopathology from tissue encapsulating implants in live bears. Retrieved transmitters underwent a technical inspection. Of the 125 transmitters removed from live bears, 66 were free-floating in the peritoneal cavity [a mean (SD) of 3.8 (1.5) years after implantation], whereas 59 were encapsulated in the greater omentum [4.0 (1.8) years after implantation]. Histopathology of biopsies of the 1-15 mm thick capsules in 33 individuals showed that it consisted of organized layers of connective tissue. In one third of the bears, the inner part of the capsule was characterized by a foreign body reaction. We inspected 68 implants that had been in bears for 3.9 (2.4) years. The batteries had short-circuited four (5.9%) of these devices. This resulted in the death of two animals 10 and 13 years after implantation. In two other bears that underwent surgery, we found the short-circuited devices to be fully encapsulated within the peritoneal cavity 5 and 6 years after implantation. A significant proportion of the other 64 inspected implants showed serious technical problems, such as corrosion of metal parts or the batteries (50%), detachment of the end cap (11.8%), and erosion (7.4%) or melting (5.9%) of the wax coating. We conclude that the wax coating of the transmitters was not biocompatible, that the technical quality of the devices was poor, and that these implants should not be used in brown bears.
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http://dx.doi.org/10.3389/fvets.2018.00252DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6196346PMC
October 2018

Seasonal changes in eicosanoid metabolism in the brown bear.

Naturwissenschaften 2018 Sep 17;105(9-10):58. Epub 2018 Sep 17.

CARMEN, INSERM U1060 / University of Lyon / INRA U1235, Oullins, France.

Polyunsaturated fatty acids (PUFAs) exert several important functions across organ systems. During winter, hibernators divert PUFAs from oxidation, retaining them in their tissues and membranes, to ensure proper body functions at low body temperature. PUFAs are also precursors of eicosanoids with pro- and anti-inflammatory properties. This study investigated seasonal changes in eicosanoid metabolism of free-ranging brown bears (Ursus arctos). By using a lipidomic approach, we assessed (1) levels of specific omega-3 and omega-6 fatty acids involved in the eicosanoid cascade and (2) concentrations of eicosanoids in skeletal muscle and blood plasma of winter hibernating and summer active bears. We observed significant seasonal changes in the specific omega-3 and omega-6 precursors. We also found significant seasonal alterations of eicosanoid levels in both tissues. Concentrations of pro-inflammatory eicosanoids, such as thromboxane B2, 5-hydroxyeicosatetraenoic acid (HETE), and 15-HETE and 18-HETE, were significantly lower in muscle and/or plasma of hibernating bears compared to summer-active animals. Further, plasma and muscle levels of 5,6-epoxyeicosatrienoic acid (EET), as well as muscle concentration of 8,9-EET, tended to be lower in bears during winter hibernation vs. summer. We also found lower plasma levels of anti-inflammatory eicosanoids, such as 15dPGJ and PGE, in bears during winter hibernation. Despite of the limited changes in omega-3 and omega-6 precursors, plasma and muscle concentrations of the products of all pathways decreased significantly, or remained unchanged, independent of their pro- or anti-inflammatory properties. These findings suggest that hibernation in bears is associated with a depressed state of the eicosanoid cascade.
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http://dx.doi.org/10.1007/s00114-018-1583-8DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6182652PMC
September 2018

Brown bear (Ursus arctos) attacks resulting in human casualties in Scandinavia 1977-2016; management implications and recommendations.

PLoS One 2018 23;13(5):e0196876. Epub 2018 May 23.

Norwegian Institute for Nature Research, Trondheim, Norway.

Human persecution and habitat loss have endangered large carnivore populations worldwide, but some are recovering, exacerbating old conflicts. Carnivores can injure and kill people; the most dramatic form of wildlife-human conflict. In Scandinavia, the brown bear (Ursus arctos) population increased from ~500 bears in 1977 to ~3300 in 2008, with an increase in injuries, fatalities, and public fear of bear attacks. We reviewed media coverage and interviewed victims to explore how bear population trends, hunter education, and other factors may have influenced the number of injuries and fatalities in Scandinavia from 1977 to 2016. We found 42 incidents with 42 injuries and 2 fatalities; 42 were adult men, one was an adult woman conducting forestry work, and one was a boy skiing off-piste. Thirty-three adult men were hunting bears, moose, or small game, often with a hunting dog, and 26 had shot at the bear at 8±11 m before injury. Eleven nonhunters were conducting forestry work, inspecting a hunting area, picking berries, tending livestock, hiking, harassing a denned bear, and one person was killed outside his house at night. Eight of the 11 incidents of nonhunters involved female bears with cubs; three of these family groups were in dens and two were on carcasses. The annual number of hunters injured/killed was mostly influenced by the increase in the bear population size. The pattern was similar regarding injuries/fatalities to other outdoor users, but the relation with the bear population size was weaker than for hunters, and the null model was equally supported. Bear physiology at denning may make encounters with bears more risky in the fall, when bears show prehibernation behavior. Awareness and education efforts, especially among hunters, seem important to ensure human safety. Recreationists and forestry workers should avoid dense vegetation or make noise to warn bears of their presence.
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http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0196876PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5965840PMC
August 2018

Can only poorer European countries afford large carnivores?

PLoS One 2018 27;13(4):e0194711. Epub 2018 Apr 27.

Faculty of Environmental Sciences and Natural Resources Management, Norwegian University of Life Sciences, Norway, and Norwegian Institute for Nature Research, Trondheim, Norway.

Background: One of the classic approaches in environmental economics is the environmental Kuznets curve, which predicts that when a national economy grows from low to medium levels, threats to biodiversity conservation increase, but they decrease when the economy moves from medium to high. We evaluated this approach by examining how population densities of the brown bear (Ursus arctos), gray wolf (Canis lupus), and Eurasian lynx (Lynx lynx) were related to the national economy in 24 European countries.

Methodology/principal Findings: We used forest proportions, the existence of a compensation system, and country group (former socialist countries, Nordic countries, other countries) as covariates in a linear model with the first- and the second-order polynomial terms of per capita gross domestic product (GDP). Country group was treated as a random factor, but remained insignificant and was ignored. All models concerning brown bear and wolf provided evidence that population densities decreased with increasing GDP, but densities of lynx were virtually independent of GDP. Models for the wolf explained >80% of the variation in densities, without a difference between the models with all independent variables and the model with only GDP. For the bear, the model with GDP alone accounted for 10%, and all three variables 33%, of the variation in densities.

Conclusions: Wolves exhibit a higher capacity for dispersal and reproduction than bear or lynx, but still exists at the lowest densities in wealthy European countries. We are aware that several other factors, not available for our models, influenced large carnivore densities. Based on the pronounced differences among large carnivore species in their countrywide relationships between densities and GDP, and a strikingly high relationship for the gray wolf, we suggest that our results reflected differences in political history and public acceptance of these species among countries. The compensation paid for the damages caused by the carnivores is not a key to higher carnivore densities, but might be necessity for the presence of large carnivores to be accepted in countries with high GDP.
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http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0194711PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5922549PMC
July 2018

A case for considering individual variation in diel activity patterns.

Behav Ecol 2017 Nov-Dec;28(6):1524-1531. Epub 2017 Sep 11.

Faculty of Environmental Sciences and Natural Resource Management, Norwegian University of Life Sciences, NO-1430 Ås, Norway.

There is a growing recognition of the role of individual variation in patterns emerging at higher levels of biological organization. Despite the importance of the temporal configuration of ecological processes and patterns, intraspecific individual variation in diel activity patterns is almost never accounted for in behavioral studies at the population level. We used individual-based monitoring data from 98 GPS-collared brown bears in Scandinavia to estimate diel activity patterns before the fall hunting season. We extracted 7 activity measures related to timing and regularity of activity from individual activity profiles. We then used multivariate analysis to test for the existence of distinct activity tactics and their environmental determinants, followed by generalized linear regression to estimate the extent of within-individual repeatability of activity tactics. We detected 4 distinct activity tactics, with a high degree of individual fidelity to a given tactic. Demographic factors, availability of key foraging habitat, and human disturbance were important determinants of activity tactics. Younger individuals and those with higher bear and road densities within their home range were more nocturnal and more likely to rest during the day. Good foraging habitat and increasing age led to more diurnal activity patterns and nocturnal resting periods. We did not find evidence of diel activity tactics influencing survival during the subsequent hunting season. We conclude that individual variation in activity deserves greater attention than it currently receives, as it may help account for individual heterogeneity in fitness and could facilitate within-population niche partitioning that can have population- or community-level consequences.
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http://dx.doi.org/10.1093/beheco/arx122DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5873257PMC
September 2017

Proteolysis inhibition by hibernating bear serum leads to increased protein content in human muscle cells.

Sci Rep 2018 04 3;8(1):5525. Epub 2018 Apr 3.

CarMeN Laboratory, INSERM, INRA, University of Lyon, Pierre-Benite, France.

Muscle atrophy is one of the main characteristics of human ageing and physical inactivity, with resulting adverse health outcomes. To date, there are still no efficient therapeutic strategies for its prevention and/or treatment. However, during hibernation, bears exhibit a unique ability for preserving muscle in conditions where muscle atrophy would be expected in humans. Therefore, our objective was to determine whether there are components of bear serum which can control protein balance in human muscles. In this study, we exposed cultured human differentiated muscle cells to bear serum collected during winter and summer periods, and measured the impact on cell protein content and turnover. In addition, we explored the signalling pathways that control rates of protein synthesis and degradation. We show that the protein turnover of human myotubes is reduced when incubated with winter bear serum, with a dramatic inhibition of proteolysis involving both proteasomal and lysosomal systems, and resulting in an increase in muscle cell protein content. By modulating intracellular signalling pathways and inducing a protein sparing phenotype in human muscle cells, winter bear serum therefore holds potential for developing new tools to fight human muscle atrophy and related metabolic disorders.
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http://dx.doi.org/10.1038/s41598-018-23891-5DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5883044PMC
April 2018

Hunting regulation favors slow life histories in a large carnivore.

Nat Commun 2018 03 27;9(1):1100. Epub 2018 Mar 27.

Département de biologie, Université de Sherbrooke, Sherbrooke, QC, J1K 2R1, Canada.

As an important extrinsic source of mortality, harvest should select for fast reproduction and accelerated life histories. However, if vulnerability to harvest depends upon female reproductive status, patterns of selectivity could diverge and favor alternative reproductive behaviors. Here, using more than 20 years of detailed data on survival and reproduction in a hunted large carnivore population, we show that protecting females with dependent young, a widespread hunting regulation, provides a survival benefit to females providing longer maternal care. This survival gain compensates for the females' reduced reproductive output, especially at high hunting pressure, where the fitness benefit of prolonged periods of maternal care outweighs that of shorter maternal care. Our study shows that hunting regulation can indirectly promote slower life histories by modulating the fitness benefit of maternal care tactics. We provide empirical evidence that harvest regulation can induce artificial selection on female life history traits and affect demographic processes.
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http://dx.doi.org/10.1038/s41467-018-03506-3DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5871616PMC
March 2018

Can concentrations of steroid hormones in brown bear hair reveal age class?

Conserv Physiol 2018 29;6(1):coy001. Epub 2018 Jan 29.

Department of Natural Sciences and Environmental Health, Telemark University College of Southeast Norway, NO-3800 Bø i Telemark, Norway.

Although combining genetic and endocrine data from non-invasively collected hair samples has potential to improve the conservation of threatened mammals, few studies have evaluated this opportunity. In this study, we determined if steroid hormone (testosterone, progesterone, estradiol and cortisol) concentration profiles in 169 hair samples collected from free-ranging brown bears () could be used to accurately discriminate between immature and adult bears within each sex. Because hair samples were acquired opportunistically, we also needed to establish if interactions between hormones and several non-hormone factors (ordinal day, year, contact method, study area) were associated with age class. For each sex, we first compared a suite of candidate models by Akaike Information Criteria model selection, using different adult-age thresholds (3, 4 and 5 years), to determine the most supported adult age. Because hair hormone levels better reflect the endocrine state at an earlier time, possibly during the previous year, then at the time of sampling, we re-analysed the data, excluding the records for bears at the adult-age threshold, to establish if classification accuracy improved. For both sexes, candidate models were most supported based on a 3-year-old adult-age threshold. Classification accuracy did not improve with the 3-year-old bear data excluded. Male age class was predicted with a high degree of accuracy (88.4%) based on the concomitant concentrations of all four hormones. Female age class was predicted with less accuracy (77.1%) based only on testosterone and cortisol. Accuracy was reduced for females, primarily because we had poor success in correctly classifying immature bears (60%) whereas classification success for adult females was similar to that for males (84.5%). Given the small and unbalanced sample used in this study, our findings should be viewed as preliminary, but they should also provide a basis for more comprehensive future studies.
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http://dx.doi.org/10.1093/conphys/coy001DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5788069PMC
January 2018

Regulated hunting re-shapes the life history of brown bears.

Nat Ecol Evol 2018 Jan 11;2(1):116-123. Epub 2017 Dec 11.

Faculty of Environmental Sciences and Natural Resource Management, Norwegian University of Life Sciences, Ås, Norway.

Management of large carnivores is among the most controversial topics in natural resource administration. Regulated hunting is a centrepiece of many carnivore management programmes and, although a number of hunting effects on population dynamics, body-size distributions and life history in other wildlife have been observed, its effects on life history and demography of large carnivores remain poorly documented. We report results from a 30-year study of brown bears (Ursus arctos) analysed using an integrated hierarchical approach. Our study revealed that regulated hunting has severely disrupted the interplay between age-specific survival and environmental factors, altered the consequences of reproductive strategies, and changed reproductive values and life expectancy in a population of the world's largest terrestrial carnivore. Protection and sustainable management have led to numerical recovery of several populations of large carnivores, but managers and policymakers should be aware of the extent to which regulated hunting may be influencing vital rates, thereby reshaping the life history of apex predators.
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http://dx.doi.org/10.1038/s41559-017-0400-7DOI Listing
January 2018

Sociodemographic factors modulate the spatial response of brown bears to vacancies created by hunting.

J Anim Ecol 2018 Jan 13;87(1):247-258. Epub 2017 Nov 13.

Department of Natural Sciences and Environmental Health, Faculty of Technology, Natural Sciences, and Maritime Sciences, University College of Southeast Norway, Telemark, Norway.

There is a growing recognition of the importance of indirect effects from hunting on wildlife populations, e.g. social and behavioural changes due to harvest, which occur after the initial offtake. Nonetheless, little is known about how the removal of members of a population influences the spatial configuration of the survivors. We studied how surviving brown bears (Ursus arctos) used former home ranges that had belonged to casualties of the annual bear hunting season in southcentral Sweden (2007-2015). We used resource selection functions to explore the effects of the casualty's and survivor's sex, age and their pairwise genetic relatedness, population density and hunting intensity on survivors' spatial responses to vacated home ranges. We tested the competitive release hypothesis, whereby survivors that increase their use of a killed bear's home range are presumed to have been released from intraspecific competition. We found strong support for this hypothesis, as survivors of the same sex as the casualty consistently increased their use of its vacant home range. Patterns were less pronounced or absent when the survivor and casualty were of opposite sex. Genetic relatedness between the survivor and the casualty emerged as the most important factor explaining increased use of vacated male home ranges by males, with a stronger response from survivors of lower relatedness. Relatedness was also important for females, but it did not influence use following removal; female survivors used home ranges of higher related female casualties more, both before and after death. Spatial responses by survivors were further influenced by bear age, population density and hunting intensity. We have shown that survivors exhibit a spatial response to vacated home ranges caused by hunting casualties, even in nonterritorial species such as the brown bear. This spatial reorganization can have unintended consequences for population dynamics and interfere with management goals. Altogether, our results underscore the need to better understand the short- and long-term indirect effects of hunting on animal social structure and their resulting distribution in space.
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http://dx.doi.org/10.1111/1365-2656.12767DOI Listing
January 2018

Staying cool or staying safe in a human-dominated landscape: which is more relevant for brown bears?

Oecologia 2017 Oct 8;185(2):191-194. Epub 2017 Sep 8.

Faculty of Environmental Sciences and Natural Resource Management, Norwegian University of Life Sciences, Postbox 5003, NO-1432, Ås, Norway.

Pigeon et al. (2016) Staying cool in a changing landscape: the influence of maximum daily ambient temperature on grizzly bear habitat selection. Oecologia 181:1101. doi: 10.1007/s00442-016-3630-5 analyzed the effect of ambient temperature on the habitat selection of grizzly bears (Ursus arctos) in Alberta, Canada. They concluded that temperature played a significant role in bear habitat selection and that it was unlikely that human activity introduced biases to the habitat selection of bears. However, Pigeon et al. did not consider variables related to human activities in their analyses. They also misinterpreted previous research that has accounted for temperature in the habitat selection of brown bears. There is much literature published on the negative effects of human disturbance on wildlife in general and on bears in particular. Downplaying the role of human disturbance could have important negative consequences if, in fact, human disturbance were a more important factor than thermoregulation. Indeed, dismissing the importance of human influence, in the face of contradictory evidence, could tempt managers to disregard an important factor that is difficult and often unpopular to deal with in their conservation plans.
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http://dx.doi.org/10.1007/s00442-017-3948-7DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5617871PMC
October 2017

Humans and climate change drove the Holocene decline of the brown bear.

Sci Rep 2017 09 4;7(1):10399. Epub 2017 Sep 4.

Faculty of Environmental Sciences and Natural Resource Management, Norwegian University of Life Sciences, PO Box 5003, NO-1432, Ås, Norway.

The current debate about megafaunal extinctions during the Quaternary focuses on the extent to which they were driven by humans, climate change, or both. These two factors may have interacted in a complex and unexpected manner, leaving the exact pathways to prehistoric extinctions unresolved. Here we quantify, with unprecedented detail, the contribution of humans and climate change to the Holocene decline of the largest living terrestrial carnivore, the brown bear (Ursus arctos), on a continental scale. We inform a spatially explicit metapopulation model for the species by combining life-history data and an extensive archaeofaunal record from excavations across Europe with reconstructed climate and land-use data reaching back 12,000 years. The model reveals that, despite the broad climatic niche of the brown bear, increasing winter temperatures contributed substantially to its Holocene decline - both directly by reducing the species' reproductive rate and indirectly by facilitating human land use. The first local extinctions occurred during the Mid-Holocene warming period, but the rise of the Roman Empire 2,000 years ago marked the onset of large-scale extinctions, followed by increasingly rapid range loss and fragmentation. These findings strongly support the hypothesis that complex interactions between climate and humans may have accelerated megafaunal extinctions.
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http://dx.doi.org/10.1038/s41598-017-10772-6DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5583342PMC
September 2017

Sex-specific genetic analysis indicates low correlation between demographic and genetic connectivity in the Scandinavian brown bear (Ursus arctos).

PLoS One 2017 3;12(7):e0180701. Epub 2017 Jul 3.

Norwegian Institute of Bioeconomy Research, NIBIO - Svanhovd, Svanvik, Norway.

The degree of gene flow within and among populations, i.e. genetic population connectivity, may closely track demographic population connectivity. Alternatively, the rate of gene flow may change relative to the rate of dispersal. In this study, we explored the relationship between genetic and demographic population connectivity using the Scandinavian brown bear as model species, due to its pronounced male dispersal and female philopatry. Thus, we expected that females would shape genetic structure locally, whereas males would act as genetic mediators among regions. To test this, we used eight validated microsatellite markers on 1531 individuals sampled noninvasively during country-wide genetic population monitoring in Sweden and Norway from 2006 to 2013. First, we determined sex-specific genetic structure and substructure across the study area. Second, we compared genetic differentiation, migration/gene flow patterns, and spatial autocorrelation results between the sexes both within and among genetic clusters and geographic regions. Our results indicated that demographic connectivity was not a reliable indicator of genetic connectivity. Among regions, we found no consistent difference in long-term gene flow and estimated current migration rates between males and females. Within regions/genetic clusters, only females consistently displayed significant positive spatial autocorrelation, indicating male-biased small-scale dispersal. In one cluster, however, males showed a dispersal pattern similar to females. The Scandinavian brown bear population has experienced substantial recovery over the last decades; however, our results did not show any changes in its large-scale population structure compared to previous studies, suggesting that an increase in population size and dispersal of individuals does not necessary lead to increased genetic connectivity. Thus, we conclude that both genetic and demographic connectivity should be estimated, so as not to make false assumptions about the reality of wildlife populations.
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http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0180701PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5495496PMC
October 2017

Don't forget to look down - collaborative approaches to predator conservation.

Biol Rev Camb Philos Soc 2017 Nov 24;92(4):2157-2163. Epub 2017 Mar 24.

Department of Zoology and Merton College, Tasso Leventis Professor of Biodiversity, University of Oxford, Oxford OX1 3PS, U.K.

Finding effective ways of conserving large carnivores is widely recognised as a priority in conservation. However, there is disagreement about the most effective way to do this, with some favouring top-down 'command and control' approaches and others favouring collaboration. Arguments for coercive top-down approaches have been presented elsewhere; here we present arguments for collaboration. In many parts of the developed world, flexibility of approach is built into the legislation, so that conservation objectives are balanced with other legitimate goals. In the developing world, limited resources, poverty and weak governance mean that collaborative approaches are likely to play a particularly important part in carnivore conservation. In general, coercive policies may lead to the deterioration of political legitimacy and potentially to non-compliance issues such as illegal killing, whereas collaborative approaches may lead to psychological ownership, enhanced trust, learning, and better social outcomes. Sustainable hunting/trapping plays a crucial part in the conservation and management of many large carnivores. There are many different models for how to conserve carnivores effectively across the world, research is now required to reduce uncertainty and examine the effectiveness of these approaches in different contexts.
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http://dx.doi.org/10.1111/brv.12326DOI Listing
November 2017