When fed ad libitum, ursids can maximize mass gain by selecting mixed diets wherein protein provides 17 ± 4% of digestible energy, relative to carbohydrates or lipids. In the wild, this ability is ...likely constrained by seasonal food availability, limits of intake rate as body size increases, and competition. By visiting locations of 37 individuals during 274 bear-days, we documented foods consumed by grizzly (Ursus arctos) and black bears (Ursus americanus) in Grand Teton National Park during 2004-2006. Based on published nutritional data, we estimated foods and macronutrients as percentages of daily energy intake. Using principal components and cluster analyses, we identified 14 daily diet types. Only 4 diets, accounting for 21% of days, provided protein levels within the optimal range. Nine diets (75% of days) led to over-consumption of protein, and 1 diet (3% of days) led to under-consumption. Highest protein levels were associated with animal matter (i.e., insects, vertebrates), which accounted for 46-47% of daily energy for both species. As predicted: 1) daily diets dominated by high-energy vertebrates were positively associated with grizzly bears and mean percent protein intake was positively associated with body mass; 2) diets dominated by low-protein fruits were positively associated with smaller-bodied black bears; and 3) mean protein was highest during spring, when high-energy plant foods were scarce, however it was also higher than optimal during summer and fall. Contrary to our prediction: 4) allopatric black bears did not exhibit food selection for high-energy foods similar to grizzly bears. Although optimal gain of body mass was typically constrained, bears usually opted for the energetically superior trade-off of consuming high-energy, high-protein foods. Given protein digestion efficiency similar to obligate carnivores, this choice likely supported mass gain, consistent with studies showing monthly increases in percent body fat among bears in this region.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Animal movements are major determinants of energy expenditure and ultimately the cost-benefit of landscape use. Thus, we sought to understand those costs and how grizzly bears (
) move in mountainous ...landscapes. We trained captive grizzly bears to walk on a horizontal treadmill and up and down 10% and 20% slopes. The cost of moving upslope increased linearly with speed and slope angle, and this was more costly than moving horizontally. The cost of downslope travel at slower speeds was greater than the cost of traveling horizontally but appeared to decrease at higher speeds. The most efficient walking speed that minimized cost per unit distance was 1.19±0.11 m s
However, grizzly bears fitted with GPS collars in the Greater Yellowstone Ecosystem moved at an average velocity of 0.61±0.28 m s
and preferred to travel on near-horizontal slopes at twice their occurrence. When traveling uphill or downhill, grizzly bears chose paths across all slopes that were ∼54% less steep and costly than the maximum available slope. The net costs (J kg
m
) of moving horizontally and uphill were the same for grizzly bears, humans and digitigrade carnivores, but those costs were 46% higher than movement costs for ungulates. These movement costs and characteristics of landscape use determined using captive and wild grizzly bears were used to understand the strategies that grizzly bears use for preying on large ungulates and the similarities in travel between people and grizzly bears that might affect the risk of encountering each other on shared landscapes.
Avoiding humans will be more difficult and energetically costly for animals as outdoor recreation increases and people venture farther into wildland areas that provide high‐quality habitat for ...wildlife. Restricting human access can be an attractive management tool to mitigate effects of human recreation activities on wildlife; however, the efficacy of such measures is rarely assessed. In 1982, Yellowstone National Park identified areas important to grizzly bears (Ursus arctos) to help protect critical grizzly bear habitat and reduce the likelihood of human injuries by bears. Referred to as bear management areas (BMAs), human access is restricted in these areas for 2–8 months each year, with timing and type of restrictions varying by area. We examined 2 datasets to evaluate grizzly bear selection of BMAs and differences of bear density in BMAs and non‐BMAs. First, we used 17 years of recent global positioning system telemetry data for grizzly bears to assess their selection of BMAs during periods when human access was allowed, and when access was restricted. We used step‐selection functions to test the hypothesis that bears spend time in places that allow them to avoid people and select quality food sources. There was support that grizzly bears differentially select for BMAs regardless of whether human access was restricted at the time, compared with areas outside BMAs, and that selection changed with sex and season. Only males during the summer and hyperphagic seasons changed their selection of BMAs based on whether access restrictions were in place, and overall, male bears preferred unrestricted BMAs (BMAs without restrictions in place). Females preferentially selected BMAs regardless of whether the area had access restrictions in place only during the mating season. Individuals varied widely in their preference for BMAs and access restrictions. Bears likely choose to spend time in BMAs based on available food resources rather than restrictions to human access. Supporting this interpretation, our analyses indicated that a greater proportion of BMA in an area was associated with higher densities of grizzly bear. Thus, restrictions to human access likely help reduce the potential for human–bear interactions, accomplishing one of the original objectives for establishing the BMAs.
In 1982, Yellowstone National Park instituted seasonal (from 2 to 8 months) restrictions to human access in areas thought to be important to grizzly bears (Bear Management Areas, BMAs), to help protect critical habitat for grizzly bears and reduce the likelihood of human injuries by bears. We evaluated selection of these areas by grizzly bears and found support that they selected BMAs, but that bears likely chose to spend time in these areas for the resources available rather than the restrictions to human access. We also found higher densities of grizzly bears inside BMAs, compared with outside these areas, suggesting that restrictions of human access likely helped to reduce the potential for human‐bear interactions.
Understanding the density‐dependent processes that drive population demography in a changing world is critical in ecology, yet measuring performance–density relationships in long‐lived mammalian ...species demands long‐term data, limiting scientists' ability to observe such mechanisms. We tested performance–density relationships for an opportunistic omnivore, grizzly bears (Ursus arctos, Linnaeus, 1758) in the Greater Yellowstone Ecosystem, with estimates of body composition (lean body mass and percent body fat) serving as indicators of individual performance over two decades (2000–2020) during which time pronounced environmental changes have occurred. Several high‐calorie foods for grizzly bears have mostly declined in recent decades (e.g., whitebark pine Pinus albicaulis, Engelm, 1863), while increasing human impacts from recreation, development, and long‐term shifts in temperatures and precipitation are altering the ecosystem. We hypothesized that individual lean body mass declines as population density increases (H1), and that this effect would be more pronounced among growing individuals (H2). We also hypothesized that omnivory helps grizzly bears buffer energy intake from changing foods, with body fat levels being independent from population density and environmental changes (H3). Our analyses showed that individual lean body mass was negatively related to population density, particularly among growing‐age females, supporting H1 and partially H2. In contrast, population density or sex had little effect on body fat levels and rate of accumulation, indicating that sufficient food resources were available on the landscape to accommodate successful use of shifting food sources, supporting H3. Our results offer important insights into ecological feedback mechanisms driving individual performances within a population undergoing demographic and ecosystem‐level changes. However, synergistic effects of continued climate change and increased human impacts could lead to more extreme changes in food availability and affect observed population resilience mechanisms. Our findings underscore the importance of long‐term studies in protected areas when investigating complex ecological relationships in an increasingly anthropogenic world.
Revealing mechanisms underlying demography of long‐lived omnivores is of critical in a changing world. Using long‐term data on grizzly bears from the Greater Yellowstone Ecosystem, we show that variation in individual performance (measured as lean body mass) is primarily a density‐dependent process, particularly among younger females. Conversely, large omnivores exhibit plastic tactics in the face of current landscape‐level perturbations maintaining seasonal threshold of body fat required for hibernation and reproduction. However, synergistic effects of continued climate change and other forms of human impact could in the future affect food availability, and consequently, the observed population resilience mechanisms.
Changes in life history traits of species can be an important indicator of potential factors influencing populations. For grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem (GYE), ...recent decline of whitebark pine (WBP; Pinus albicaulis), an important fall food resource, has been paired with a slowing of population growth following two decades of robust population increase. These observations have raised questions whether resource decline or density-dependent processes may be associated with changes in population growth. Distinguishing these effects based on changes in demographic rates can be difficult. However, unlike the parallel demographic responses expected from both decreasing food availability and increasing population density, we hypothesized opposing behavioral responses of grizzly bears with regard to changes in home-range size. We used the dynamic changes in food resources and population density of grizzly bears as a natural experiment to examine hypotheses regarding these potentially competing influences on grizzly bear home-range size. We found that home-range size did not increase during the period of whitebark pine decline and was not related to proportion of whitebark pine in home ranges. However, female home-range size was negatively associated with an index of population density. Our data indicate that home-range size of grizzly bears in the GYE is not associated with availability of WBP, and, for female grizzly bears, increasing population density may constrain home-range size.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
In 2011, 2 hikers were killed by grizzly bears (Ursus arctos) in separate incidents on backcountry trails in Hayden Valley, Yellowstone National Park, USA (YNP). Hayden Valley provides prime habitat ...for grizzly bears and is known to have high densities of bears. During 1970–2017, 23% (10 of 44) of all backcountry grizzly bear–inflicted human injuries and fatalities in YNP occurred in the valley even though it comprises only 1% of the park. In addition, 3 of the last 5 fatal bear attacks in the park occurred in the valley. We evaluated retrospectively whether restrictions and closures on visitor recreational activity would have prevented many of these injuries. We considered prohibitions on recreational activity during seasons when bears forage for specific high-quality foods; potential closures that coincided with the times of day and year bears were most active in the valley; and visitor use restrictions that would have prevented the most common human behaviors associated with grizzly bear–caused human injuries. The food-based closure that may have prevented the most human injuries occurred during middle to late summer when bears scavenge bison (Bison bison) carcasses that result from annual rutting behavior of bison in the valley. However, safety precautions such as hiking in groups of ≥3, remaining on maintained trails, and carrying bear spray would likely reduce the frequency of bear-inflicted human injuries more than most food-based seasonal closures. Our analyses provide broadly applicable findings regarding use of visitor behavior restrictions and seasonal closures to reduce the risk of bear-inflicted human injuries.
Heat dissipation limit theory posits that energy available for growth and reproduction in endotherms is limited by their ability to dissipate heat. In mammals, endogenous heat production increases ...markedly during gestation and lactation, and thus female mammals may be subject to greater thermal constraints on energy expenditure than males. Such constraints likely have important implications for behaviour and population performance in a warming climate.
We used a mechanistic simulation model based on the first principles of heat and mass transfer to study thermal constraints on activity (both timing and intensity) of captive female grizzly bears Ursus arctos in current and future climate scenarios. We then quantified the relative importance of regulatory behaviours for maintaining heat balance using GPS telemetry locations of lactating versus non‐lactating female bears from Yellowstone National Park, and assessed the degree to which costs of thermoregulation constrained the distribution of sampled bears in space and time.
Lactating female bears benefitted considerably more from behavioural cooling mechanisms (e.g. partial submersion in cool water or bedding on cool substrate) than non‐lactating females in our simulations; the availability of water for thermoregulation increased the number of hours during which lactating females could be active by up to 60% under current climatic conditions and by up to 43% in the future climate scenario. Moreover, even in the future climate scenario, lactating bears were able to achieve heat balance 24 hr/day by thermoregulating behaviourally when water was available to facilitate cooling.
The most important predictor of female grizzly bear distribution in Yellowstone, regardless of reproductive status, was elevation. However, variables associated with the thermal environment were relatively more important for predicting the distribution of lactating than non‐lactating female bears.
Our results suggest that the costs of heat dissipation, which are modulated by climate, may impose constraints on the behaviour and energetics of large endotherms like grizzly bears, and that access to water for cooling will likely be an increasingly important driver of grizzly bear distribution in Yellowstone as the climate continues to warm.
A free Plain Language Summary can be found within the Supporting Information of this article.
A free Plain Language Summary can be found within the Supporting Information of this article.
Effective population size (Nₑ) is a key parameter for monitoring the genetic health of threatened populations because it reflects a population's evolutionary potential and risk of extinction due to ...genetic stochasticity. However, its application to wildlife monitoring has been limited because it is difficult to measure in natural populations. The isolated and well‐studied population of grizzly bears (Ursus arctos) in the Greater Yellowstone Ecosystem provides a rare opportunity to examine the usefulness of different Nₑ estimators for monitoring. We genotyped 729 Yellowstone grizzly bears using 20 microsatellites and applied three single‐sample estimators to examine contemporary trends in generation interval (GI), effective number of breeders (Nb) and Nₑ during 1982–2007. We also used multisample methods to estimate variance (NₑV) and inbreeding Nₑ (NₑI). Single‐sample estimates revealed positive trajectories, with over a fourfold increase in Nₑ (≈100 to 450) and near doubling of the GI (≈8 to 14) from the 1980s to 2000s. NₑV (240–319) and NₑI (256) were comparable with the harmonic mean single‐sample Nₑ (213) over the time period. Reanalysing historical data, we found NₑV increased from ≈80 in the 1910s–1960s to ≈280 in the contemporary population. The estimated ratio of effective to total census size (Nₑ/Nc) was stable and high (0.42–0.66) compared to previous brown bear studies. These results support independent demographic evidence for Yellowstone grizzly bear population growth since the 1980s. They further demonstrate how genetic monitoring of Nₑ can complement demographic‐based monitoring of Nc and vital rates, providing a valuable tool for wildlife managers.
The Greater Yellowstone Ecosystem (GYE) has experienced changes in the distribution and availability of grizzly bear (Ursus arctos) food resources in recent decades. The decline of ungulates, fish, ...and whitebark pine seeds (Pinus albicaulis) has prompted questions regarding their ability to adapt. We examined body composition and diet of grizzly bears using bioelectrical impedance and stable isotopes to determine if 1) we can detect a change in diet quality associated with the decline in either ungulates or whitebark pine, and 2) the combined decline in ungulates, fish, and pine seeds resulted in a change in grizzly bear carrying capacity in the GYE. We contrasted body fat and mass in grizzly bears with a potential competitor, the American black bear (Ursus americanus) , to address these questions. Grizzly bears assimilated more meat into their diet and were in better body condition than black bears throughout the study period, indicating the decline in ungulate resources did not affect grizzly bears more than black bears. We also found no difference in autumn fat levels in grizzly bears in years of good or poor pine seed production, and stable isotope analyses revealed this was primarily a function of switching to meat resources during poor seedproducing years. This dietary plasticity was consistent over the course of our study. We did not detect an overall downward trend in either body mass or the fraction of meat assimilated into the diet by grizzly bears over the past decade, but we did detect a downward trend in percent body fat in adult female grizzly bears after 2006. Whether this decline is an artifact of small sample size or due to the population reaching the ecological carrying capacity of the Yellowstone ecosystem warrants further investigation.