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Summary 1. We explored multiple linkages among grey wolves (Canis lupus), elk (Cervus elaphus), berry-producing shrubs and grizzly bears (Ursus arctos) in Yellowstone National Park. 2. We hypothesized competition between elk and grizzly bears whereby, in the absence of wolves, increases in elk numbers would increase browsing on berry-producing shrubs and decrease fruit availability to grizzly bears. After wolves were reintroduced and with a reduced elk population, we hypothesized there would be an increase in the establishment of berry-producing shrubs, such as serviceberry (Amelanchier alnifolia), which is a major berry-producing plant. We also hypothesized that the percentage fruit in the grizzly bear diet would be greater after than before wolf reintroduction. 3. We compared the frequency of fruit in grizzly bear scats to elk densities prior to wolf reintroduction during a time of increasing elk densities (1968–1987). For a period after wolf reintroduction, we calculated the percentage fruit in grizzly bear scat by month based on scats collected in 2007–2009 (n = 778 scats) and compared these results to scat data collected before wolf reintroduction. Additionally, we developed an age structure for serviceberry showing the origination year of stems in a northern range study area. 4. We found that over a 19-year period, the percentage frequency of fruit in the grizzly diet (6231 scats) was inversely correlated (P

Grizzly bears (Ursus arctos) have been reported as either nocturnal or diurnal in various studies, but have not been known to switch between the 2 times unless disturbed by humans. Black bears (Ursus americanus) are almost solely diurnal in studies unless human influences occur. Because human disturbance is often difficult to control, the relative temporal niche of both species remains ill-defined. Thus, the present study examined bears in Yellowstone National Park (Wyoming) where hunting does not occur, human activities are relatively benign, and bear species are sympatric to determine if niche occupancy was a stable feature of the species. Onset of activity was anticipatory of both sunrise or morning civil twilight (illumination sufficient for human vision) for individuals of either species. The peak hour of activity in black bears was consistently midday, but fluctuated in grizzly bears from midday during early spring, late summer, and fall to evening during late spring and early summer. Black bears did not temporally avoid the times when the more dominant grizzly bears were active. Mean activity levels were higher for male black bears than for both male and female grizzly bears. Together, results suggest that the foraging needs of black bears necessitate ingestion of less-digestible, lower-quality foods requiring longer foraging time during daytime hours, whereas grizzly bears adapt their diet to seasonally available food sources, necessitating greater temporal flexibility.

The proposed dietary pattern of extinct Late Pleistocene cave bears (Ursus spelaeus Rosenmüller, 1794) has become controversial, as some authors have suggested that they were strictly vegetarian, whereas others maintain they were omnivores that at times ate large amounts of animal protein. We evaluated these alternatives by compiling stable isotope data of carbon (δ13C) and nitrogen (δ15N) from the bone collagen of adult European cave bears from the Late Pleistocene (Marine Isotopic Stage 3). The data include previously published analyses and additional data from the southeastern European (Carpathian) sites of Cioclovina, Muierii, Oase, and Urşilor. The cave bear isotopic values from bone collagen were compared with those from hair keratin occurring in grizzly bears (Ursus arctos horribilis Ord, 1815) collected from 1989 to 2009 in the western United States (Yellowstone National Park). The Yellowstone bears have access to a wide diversity of plants and animals, such that their diets can range from vegetarian to carnivorous. Thus, there was considerable δ13C and δ15N variation among the grizzly bear isotopic values, and the cave bear isotopic variation was encompassed within the overall grizzly bear isotopic distribution. More importantly, the δ15N distributions, reflecting principally trophic level, were not different between the cave bears and the grizzly bears; the cave bear values are, on average, slightly higher or lower than those of the grizzly bears, depending on the criteria for inclusion in the comparisons. It is therefore no longer appropriate to view Late Pleistocene cave bears as strictly or even predominantly vegetarian but as flexible omnivores within their diverse communities.

Global warming has the potential to reduce arctic sea ice and thereby increase the length of summer–fall fasting when polar bears (Ursus maritimus) lose access to most marine mammals. To evaluate the consequences of such changes, we compared the cost of fasting by polar bears with hibernation by brown bears (U. arctos), American black bears (U. americanus), and polar bears and made projections about tissue reserves polar bears will need to survive and reproduce as fasts become longer. Hibernating polar bears expend energy at the same rate per unit mass as do brown bears and black bears. However, daily mass losses, energy expenditures, and the losses of lean mass are much higher in fasting, active polar bears than in hibernating bears. The average pregnant polar bear living around Hudson Bay during the 1980s and 1990s could fast for 10.0 6 2.3 months (¯ X 6 SD), and the average lactating female with cubs born during the preceding winter could fast for 4.2 6 1.9 months. Thus, some pregnant or lactating females with lower levels of body fat content were already approaching or beyond the constraint of being able to produce cubs and survive the required 8 months of fasting if producing new offspring or 4 months if accompanied by older offspring. Pregnant or lactating females and their dependent offspring have the most tenuous future as global warming occurs. Thus, we predict a significant reduction in productivity with even modest increases in global warming for polar bears living in the very southern part of their range and are concerned about more northern populations depending on their ability to accumulate increasing amounts of fat.

Understanding behavioral responses of species to environmental change is critical to forecasting population-level effects. Although climate change is significantly impacting species’ distributions, few studies have examined associated changes in behavior. Polar bear (Ursus maritimus) subpopulations have varied in their near-term responses to sea ice decline. We examined behavioral responses of two adjacent subpopulations to changes in habitat availability during the annual sea ice minimum using activity data. Location and activity sensor data collected from 1989 to 2014 for 202 adult female polar bears in the Southern Beaufort Sea (SB) and Chukchi Sea (CS) subpopulations were used to compare activity in three habitat types varying in prey availability: (1) land; (2) ice over shallow, biologically productive waters; and (3) ice over deeper, less productive waters. Bears varied activity across and within habitats with the highest activity at 50–75% sea ice concentration over shallow waters. On land, SB bears exhibited variable but relatively high activity associated with the use of subsistence-harvested bowhead whale carcasses, whereas CS bears exhibited low activity consistent with minimal feeding. Both subpopulations had fewer observations in their preferred shallow-water sea ice habitats in recent years, corresponding with declines in availability of this substrate. The substantially higher use of marginal habitats by SB bears is an additional mechanism potentially explaining why this subpopulation has experienced negative effects of sea ice loss compared to the still-productive CS subpopulation. Variability in activity among, and within, habitats suggests that bears alter their behavior in response to habitat conditions, presumably in an attempt to balance prey availability with energy costs.

The majority of terrestrial locomotion studies have focused on parasagittal motion and paid less attention to forces or movement in the frontal plane. Our previous research has shown that grizzly bears produce higher medial ground reaction forces (lateral pushing from the animal) than would be expected for an upright mammal, suggesting frontal plane movement may be an important aspect of their locomotion. To examine this, we conducted an inverse dynamics analysis in the sagittal and frontal planes, using ground reaction forces and position data from three high-speed cameras of four adult female grizzly bears. Over the speed range collected, the bears used walks, running walks and canters. The scapulohumeral joint, wrist and the limb overall absorb energy (average total net work of the forelimb joints, -0.97 W kg

Grizzly bears (Ursus arctos horribilis) have evolved remarkable metabolic adaptations including enormous fat accumulation during the active season followed by fasting during hibernation. However, these fluctuations in body mass do not cause the same harmful effects associated with obesity in humans. To better understand these seasonal transitions, we performed insulin and glucose tolerance tests in captive grizzly bears, characterized the annual profiles of circulating adipokines, and tested the anorectic effects of centrally administered leptin at different times of the year. We also used bear gluteal adipocyte cultures to test insulin and beta-adrenergic sensitivity in vitro. Bears were insulin resistant during hibernation but were sensitive during the spring and fall active periods. Hibernating bears remained euglycemic, possibly due to hyperinsulinemia and hyperglucagonemia. Adipokine concentrations were relatively low throughout the active season but peaked in mid-October prior to hibernation when fat content was greatest. Serum glycerol was highest during hibernation, indicating ongoing lipolysis. Centrally administered leptin reduced food intake in October, but not in August, revealing seasonal variation in the brain’s sensitivity to its anorectic effects. This was supported by strong phosphorylated signal transducer and activator of transcription 3 labeling within the hypothalamus of hibernating bears; labeling virtually disappeared in active bears. Adipocytes collected during hibernation were insulin resistant when cultured with hibernation serum but became sensitive when cultured with active season serum. Heat treatment of active serum blocked much of this action. Clarifying the cellular mechanisms responsible for the physiology of hibernating bears may inform new treatments for metabolic disorders.

SUMMARY Hibernating bears retain most of their skeletal muscle strength despite drastically reduced weight-bearing activity. Regular neural activation of muscles is a potential mechanism by which muscle atrophy could be limited. However, both mechanical loading and neural activity are usually necessary to maintain muscle size. An alternative mechanism is that the signaling pathways related to the regulation of muscle size could be altered so that neither mechanical nor neural inputs are needed for retaining strength. More specifically, we hypothesized that muscles in hibernating bears are resistant to a severe reduction in neural activation. To test this hypothesis, we unilaterally transected the common peroneal nerve, which innervates ankle flexor muscles, in hibernating and summer-active brown bears (Ursus arctos). In hibernating bears, the long digital extensor (LDE) and cranial tibial (CT) musculotendon masses on the denervated side decreased after 11 weeks post-surgery by 18±11 and 25±10%, respectively, compared with those in the intact side. In contrast, decreases in musculotendon masses of summer-active bears after denervation were 61±4 and 58±5% in the LDE and CT, respectively, and significantly different from those of hibernating bears. The decrease due to denervation in summer-active bears was comparable to that occurring in other mammals. Whole-muscle cross-sectional areas (CSAs) measured from ultrasound images and myofiber CSAs measured from biopsies decreased similarly to musculotendon mass. Thus, hibernating bears alter skeletal muscle catabolic pathways regulated by neural activity, and exploration of these pathways may offer potential solutions for disuse atrophy of muscles.

Grizzly bears (Ursus arctos horribilis) tolerate extended periods of extremely low heart rate during hibernation without developing congestive heart failure or cardiac chamber dilation. Left ventricular atrophy and decreased left ventricular compliance have been reported in this species during hibernation. We evaluated the myocardial response to significantly reduced heart rate during hibernation by measuring relative myosin heavy-chain (MyHC) isoform expression and expression of a set of genes important to muscle plasticity and mass regulation in the left atria and left ventricles of active and hibernating bears. We supplemented these data with measurements of systolic and diastolic function via echocardiography in unanesthetized grizzly bears. Atrial strain imaging revealed decreased atrial contractility, decreased expansion/reservoir function (increased atrial stiffness), and decreased passive-filling function (increased ventricular stiffness) in hibernating bears. Relative MyHC-α protein expression increased significantly in the atrium during hibernation. The left ventricle expressed 100% MyHC-β protein in both groups. Insulin-like growth factor (IGF-I) mRNA expression was reduced by ∼50% in both chambers during hibernation, consistent with the ventricular atrophy observed in these bears. Interestingly, mRNA expression of the atrophy-related ubiquitin ligases Muscle Atrophy F-box (MAFBx) and Muscle Ring Finger 1 did not increase, nor did expression of myostatin or hypoxia-inducible factor 1α (HIF-1α). We report atrium-specific decreases of 40% and 50%, respectively, in MAFBx and creatine kinase mRNA expression during hibernation. Decreased creatine kinase expression is consistent with lowered energy requirements and could relate to reduced atrial emptying function during hibernation. Taken together with our hemodynamic assessment, these data suggest a potential downregulation of atrial chamber function during hibernation to prevent fatigue and dilation due to excessive work against an optimally filled ventricle, a response unpredicted by the Frank-Starling mechanism.

Summary 1. Accurately predicting isotope ratio discrimination is central to using mixing models to estimate assimilated diets of wild animals. This process is complicated when omnivores consume mixed diets because their discrimination is unlikely to be the weighted average of the various dietary constituents as occurs in current models. 2. We sought a basic understanding of how protein quality and quantity determine Δ15N and Δ34S in mammals and birds. Dietary protein is the primary source of both elements in many plants and animals. Low protein quality and high protein content have the potential to increase Δ15N by increasing protein turnover. 3. Protein quality, defined as the relative amount of the most limiting amino acid, accounted for 87–90% of the variation in Δ15N when mammals and birds consumed plant matter and mixed diets of plants and animals with protein of intermediate quality and quantity. However, foods containing relatively large amounts of high quality protein (i.e. vertebrate flesh) and foods with exceptionally low quality protein (e.g. lentils, Lens culinaris) had disparate nitrogen discriminations relative to what would be predicted from protein quality alone. 4. Supplementation of plant and animal diets with nitrogen-free carbohydrates and fats to reduce protein quantity did not reduce Δ15N in three plant-based diets fed to laboratory rats, but reduced Δ15N in two of three meat diets with >50% protein. 5. Δ34S was weakly correlated with Δ15N (R2 = 0·48) but was highly correlated with dietary δ34S (R2 = 0·89). Because methionine, a sulphur amino acid, was the most limiting amino acid in all diets, sulphur should be highly conserved as indicated by the lack of any change in Δ34S when diets were supplemented with carbohydrates and fat to both provide additional energy and reduce protein content. 6. Predictive equations incorporating both protein quality and quantity accounted for 81% of the variation in Δ15N and offer the opportunity to create more realistic mixing models to accurately estimate assimilated diets for omnivores.

Disuse typically causes an imbalance in bone formation and bone resorption, leading to losses of cortical and trabecular bone. In contrast, bears maintain balanced intracortical remodeling and prevent cortical bone loss during disuse (hibernation). Trabecular bone, however, is more detrimentally affected than cortical bone in other animal models of disuse. Here we investigated the effects of hibernation on bone remodeling, architectural properties, and mineral density of grizzly bear (Ursus arctos horribilis) and black bear (Ursus americanus) trabecular bone in several skeletal locations. There were no differences in bone volume fraction or tissue mineral density between hibernating and active bears or between pre- and post-hibernation bears in the ilium, distal femur, or calcaneus. Though indices of cellular activity level (mineral apposition rate, osteoid thickness) decreased, trabecular bone resorption and formation indices remained balanced in hibernating grizzly bears. These data suggest that bears prevent bone loss during disuse by maintaining a balance between bone formation and bone resorption, which consequently preserves bone structure and strength. Further investigation of bone metabolism in hibernating bears may lead to the translation of mechanisms preventing disuse induced bone loss in bears into novel treatments for osteoporosis.

Accurately predicting isotopic discrimination is central to estimating assimilated diets of wild animals when using stable isotopes. Current mixing models assume that the stable N isotope ratio (delta(15)N) discrimination (Delta(15)N) for each food in a mixed diet is constant and independent of other foods being consumed. Thus, the discrimination value for the mixed diet is the combined, weighted average for each food when consumed as the sole diet. However, if protein quality is a major determinant of Delta(15)N, discrimination values for mixed diets may be higher or lower than the weighted average and will reflect the protein quality of the entire diet and not that of the individual foods. This potential difference occurs because the protein quality of a mixed diet depends on whether, and to what extent, the profiles and amounts of essential amino acids in the individual foods are complementary or non-complementary to each other in meeting the animal's requirement. We tested these ideas by determining the Delta(15)N of several common foods (corn, wheat, alfalfa, soybean, and fish meal) with known amino acid profiles when fed singly and in combination to laboratory rats. Discrimination values for the mixed diets often differed from the weighted averages for the individual foods and depended on the degree of complementation. Delta(15)N for mixed diets ranged from 1.1 per thousand lower than the weighted average for foods with complementary amino acid profiles to 0.4 per thousand higher for foods with non-complementary amino acid profiles. These differences led to underestimates as high as 44% and overestimates as high as 36% of the relative proportions of fish meal and soybean meal N, respectively, in the assimilated mixed diets. We conclude that using isotopes to estimate assimilated diets is more complex than often appreciated and will require developing more biologically based, time-sensitive models.

Energy maximization, time minimization, and linear programming models subject to various constraints have dominated foraging ecology ideas and methods for decades. However, animals must use very complex physiological processes and foraging decisions to ensure fitness that in many cases may not be adequately described by these approaches. An example of this problem occurs when brown bears, Ursus arctos, have access to both abundant salmon and fruit. Salmon are one of the most energy and nutrient dense foods available to bears. Fruits are often high in soluble carbohydrates, low to deficient in many required nutrients, and more difficult to efficiently exploit. Therefore, wild brown bears that fatten primarily on fruits without access to salmon are 50% smaller than salmon-feeding bears. Thus, we predicted based on a linear, energy-maximizing model without dietary interaction effects that wild brown bears with access to both abundant salmon and fruit would feed almost exclusively on salmon. However, wild adult females with or without accompanying offspring foraged three times longer per day on fruit than on salmon. Similarly, the relative dry matter intake of ad libitum apples and salmon by captive, adult brown bears averaged 76 ±5% fruit and 24±5% salmon. Captive brown bears consuming mixed diets with intermediate dietary protein levels had 60% lower maintenance energy costs, 37% to 139% higher efficiencies of mass gain, and 72% to 520% higher maximum rates of gain than when they consumed either salmon or fruit alone. These relationships were nonlinear functions of dietary protein content in which salmon and fruit provided complementary nutritional resources. Both wild and captive bears attempted to regulate total protein, energy, and carbohydrate intake within a multidimensional intake target that both maximized energy intake and mass gain.

Although numerous studies have documented behavioral effects of nature-based tourism on wildlife populations, few studies have determined whether behavioral changes translate to effects on individual condition and population health. This issue is currently a concern for wildlife managers in Alaska, USA, and Canada where bear viewing is a rapidly growing industry expanding into previously undisturbed bear habitats. Rather than record observations at long established tourism sites, we experimentally introduced bear viewing into 2 relatively undisturbed brown bear (Ursus arctos) populations in south-central Alaska. We examined the nutritional consequences of behavioral changes induced by the presence and activity of bear viewers for bears feeding on early summer vegetation and late-summer salmon (Oncorhynchus kisutch and O. nerka). We used Global Positioning System collars, monitored food resource availability, and quantified individual resource use and condition for a year prior to and during the introduction of bear viewing. Though bear viewing altered spatiotemporal resource use in all treatments, total resource use declined only when we exposed bears to 24-hour daily human activity. Energy expenditure, indexed as daily travel distances, was significantly higher when bears responded by altering spatial rather than temporal resource use. However, body weight and composition were unaffected by all treatments as bears shifted their foraging to other locations or times. Managers can minimize nutritional impacts of bear-viewing programs by avoiding spatial displacement and providing predictable time periods when bears can access food resources free of human activity. Bears in this study exhibited a high degree of behavioral plasticity, which may be an important factor in identifying flagship species for sustainable ecotourism programs.

We hypothesized that there would be minimal dietary overlap between sympatric brown bears (Ursus arctos) and American black bears (U. americanus) relative to salmon (Oncorhynchus spp.) utilization when alternative foods (e.g., fruits) are abundant. To maximize the chance that we would reject this hypothesis, we examined the diets of brown and black bears known to have visited salmon streams. Species, sex, and individual identification of bears visiting salmon streams were determined by DNA analysis of hair and feces collected in 2002–2004 along those streams. Diets were estimated from fecal residues and stable isotope analyses of hair. Assimilated diets of brown bears were 66.0% (SD = 16.7%) salmon, 13.9% (SD = 7.5%) terrestrial animal matter, and 20.1% (SD = 17.2%) plant matter. Assimilated diets of black bears were 8.0% (SD = 5.4%) salmon, 8.4% (SD = 9.7%) terrestrial animal matter, and 83.6% (SD = 7.7%) plant matter. Male and female brown bears did not differ in either the proportion of dietary salmon, terrestrial animal matter, or plant matter. The relative amounts of fruit residues in the feces of brown bears (87.0%, SD = 15.2%) and black bears (91.8%, SD = 7.2%) did not differ. Both sexes of brown bears visited salmon streams and consumed significant amounts of salmon, but only male American black bears visited streams and then consumed minimal amounts of salmon. Thus, brown bears were largely carnivorous and black bears were largely herbivorous and frugivorous. This reduced dietary overlap relative to salmon and fruit use is understandable in light of the concentrated, defendable nature of salmon in small streams, the widely dispersed, non-defendable nature of abundant fruits, the dominance of brown over black bears, the higher energy requirement of the larger brown bear, and, therefore, the differing ability of the species to efficiently exploit different food resources.

Brown bears (Ursus arctos) exhibit hyperphagia each fall and can become obese in preparation for hibernation. They do this without displaying the physiological problems typically seen in obese humans, such as Type 2 diabetes and heart disease. The study of brown bear hibernation biology could therefore aid in the development of novel methods for combating metabolic diseases. To this end, we isolated mesenchymal stem cells from subcutaneous fat biopsies, and culture methods were developed to differentiate these into the adipogenic lineage. Biopsies were taken from 8 captive male (N = 6) and female (N = 2) brown bears, ages 2-12 years. Plastic adherent, fibroblast-like cells were proliferated and subsequently cryopreserved or differentiated. Differentiation conditions were optimized with respect to fetal bovine serum content and time spent in differentiation medium. Cultures were characterized through immunostaining, RT-qPCR, and Oil red O staining to quantify lipid accumulation. Adiponectin, leptin, and glycerol medium concentrations were also determined over the course of differentiation. The culturing protocol succeeded in generating hormone-sensitive lipase-expressing, lipid-producing white-type adipocytes (UCP1 negative). Serum concentration and time of exposure to differentiation medium were both positively related to lipid production. Cells cultured to low passage numbers retained similar lipid production and expression of lipid markers PLIN2 and FABP4. Ultimately, the protocols described here may be useful to biologists in the field investigating the health of wild bear populations and could potentially increase our understanding of metabolic disorders in humans.

Food habit studies are among the first steps used to understand wildlife-habitat relationships. However, these studies are in themselves insufficient to understand differences in population productivity and life histories, because they do not provide a direct measure of the energetic value or nutritional composition of the complete diet. Here, we developed a dynamic model integrating food habits and nutritional information to assess nutritional parameters of brown bear (Ursus arctos) diets among three interior ecosystems of North America. Specifically, we estimate the average amount of digestible energy and protein (per kilogram fresh diet) content in the diet and across the active season by bears living in western Alberta, the Flathead River (FR) drainage of southeast British Columbia, and the Greater Yellowstone Ecosystem (GYE). As well, we estimate the proportion of energy and protein in the diet contributed by different food items, thereby highlighting important food resources in each ecosystem. Bear diets in Alberta had the lowest levels of digestible protein and energy through all seasons, which might help explain the low reproductive rates of this population. The FR diet had protein levels similar to the recent male diet in the GYE during spring, but energy levels were lower during late summer and fall. Historic and recent diets in GYE had the most energy and protein, which is consistent with their larger body sizes and higher population productivity. However, a recent decrease in consumption of trout (Oncorhynchus clarki), whitebark pine nuts (Pinus albicaulis), and ungulates, particularly elk (Cervus elaphus), in GYE bears has decreased the energy and protein content of their diet. The patterns observed suggest that bear body size and population densities are influenced by seasonal availability of protein an energy, likely due in part to nutritional influences on mass gain and reproductive success.

Locomotion of plantigrade generalists has been relatively little studied compared with more specialised postures even though plantigrady is ancestral among quadrupeds. Bears (Ursidae) are a representative family for plantigrade carnivorans, they have the majority of the morphological characteristics identified for plantigrade species, and they have the full range of generalist behaviours. This study compared the locomotion of adult grizzly bears (Ursus arctos horribilis Linnaeus 1758), including stride parameters, gaits and analysis of three-dimensional ground reaction forces, with that of previously studied quadrupeds. At slow to moderate speeds, grizzly bears use walks, running walks and canters. Vertical ground reaction forces demonstrated the typical M-shaped curve for walks; however, this was significantly more pronounced in the hindlimb. The rate of force development was also significantly higher for the hindlimbs than for the forelimbs at all speeds. Mediolateral forces were significantly higher than would be expected for a large erect mammal, almost to the extent of a sprawling crocodilian. There may be morphological or energetic explanations for the use of the running walk rather than the trot. The high medial forces (produced from a lateral push by the animal) could be caused by frontal plane movement of the carpus and elbow by bears. Overall, while grizzly bears share some similarities with large cursorial species, their locomotor kinetics have unique characteristics. Additional studies are needed to determine whether these characters are a feature of all bears or plantigrade species.

Despite significant sexual dimorphism and differing reproductive strategies in carnivores, sexual segregation is rarely studied and is often overlooked in the management of wild populations. Potential nutritional constraints imposed by sexual dimorphism and differing reproductive strategies between the sexes have important implications, particularly when combined with differential effects of human activities on sex and age classes. We examined the effects of sexual dimorphism, reproductive strategies, and human activities (bear-viewing and hunting) on resource use by different sex and age classes of brown bears (Ursus arctos). Sexual segregation of habitat use and effects of experimental bear-viewing were quantified at a single site in south-central Alaska, U.S.A., by capturing, collaring, and observing brown bears at a salt marsh and salmon stream. Effects of salmon capture rate, availability of alternative salmon runs, harvest pressure, and numbers of annual visitors on sex and age class use were examined from data collected or previously published from 13 other sites. Bear-viewing sites on salmon streams where salmon capture rates were low (4 salmon/hour) resulted in low use by adult males (10% of all bears), except for sites with falls. However, maximum male use of viewing areas also depended on the availability of alternative salmon streams and harvest pressure. Use of habitats by females with dependent young was significantly related to the prevalence of adult males at the site. Thus, both sexual dimorphism and differing reproductive strategies led to sexual segregation in habitat use by bears. As a result of infanticide, females with young appear to prioritize avoidance of male bears over avoidance of humans when choosing habitats, in contrast to responses documented in herbivores. Because carnivores often exhibit both sexual dimorphism and infanticide, selection for sexual segregation is likely to be high. In these cases, the nutritional demands of large adult males, balanced with responses to human activity, drive dynamic temporal and spatial distributions of individuals in the population.

The hibernating bear is an excellent model for disuse osteoporosis in humans because it is a naturally occurring large animal model. Furthermore, bears and humans have similar lower limb skeletal morphology, and bears walk plantigrade like humans. Black bears (Ursus americanus) may not develop disuse osteoporosis during long periods of disuse (i.e. hibernation) because they maintain osteoblastic bone formation during hibernation. As a consequence, bone volume, mineral content, porosity, and strength are not adversely affected by annual periods of disuse. In fact, cortical bone bending strength has been shown to increase with age in hibernating black bears without a significant change in porosity. Other animals require remobilization periods 2-3 times longer than the immobilization period to recover the bone lost during disuse. Our findings support the hypothesis that black bears, which hibernate for as long as 5-7 months annually, have evolved biological mechanisms to mitigate the adverse effects of disuse on bone porosity and strength.

TROY N. TOLLEFSON, School of Biological Sciences, Washington State University, Pullman, WA 99164-4236, USA COLLEEN MATT, Division of Wildlife Conservation, Alaska Department of Fish and Game, 333 Raspberry Road, Anchorage, AK 99518, USA JOE MEEHAN, Division of Wildlife Conservation, Alaska Department of Fish and Game, 333 Raspberry Road, Anchorage, AK 99518, USA CHARLES T. ROBBINS,1 Department of Natural Resource Sciences and School of Biological Sciences, Washington State University, Pullman, WA 99164-4236, USA

The accuracy of bioptome placement for endomyocardial biopsy was evaluated using standard fluoroscopic guidance and compared simultaneously with two-dimensional echocardiography in six healthy subadult grizzly bears (Ursus arctos horribilis). Despite perceived adequate positioning of the bioptome toward the septum when judged using fluoroscopy, only one out of the six placements were actually positioned in the right ventricular apex toward the septum when evaluated using two-dimensional echocardiography. The bioptome was readily relocated to the right ventricular apex/septal location by echocardiography. The forceps were easily visualized at all times and the open cups were readily directed into the desired location for biopsy. Based on a subjective score for tissue quantity from 1 to 4 (poor to excellent), biopsies collected by echocardiographic-guided technique had a mean score of 3.4, while biopsies collected by fluoroscopic-guided technique had a mean score of 1.8. In summary, this study suggests that endomyocardial biopsy locations can be accurately positioned using two-dimensional echocardiography with the advantages of less radiation, lower procedural costs, and direct visualization of the sampling site. Clear definition of endocardial surface as afforded by echocardiography may enhance tissue sample quality and lower potential complications from this procedure.

The capability to understand the nutritional ecology of free-ranging bears has increased dramatically in the last 20 years. Advancements have occurred because (1) managers and biologists recognized the need to link habitat quality, productivity, and variability with bear movements, home ranges, and demographic parameters like reproductive output, survival, and population growth, and (2) several research teams are using new methods to build on the results of earlier field studies. Our ability to couple new field methods and empirical field research with controlled experiments using captive bears has been central to our increased understanding of bear nutrition. Newer methods include the use of stable isotopes to quantify assimilated diet and nutrient flows within ecosystems, bioelectrical impedance to measure body composition, and naturally occurring mercury to estimate fish intake. Controlled experiments using captive bears have been integral to developing methods, isolating specific variables by...

Salmon (Oncorhynchus spp.) are an important resource for terrestrial wildlife. However, the salmon requirements of wildlife populations and the role wildlife play in nutrient transport across ecosystems are largely ignored in salmon and habitat management. Any activity that reduces the availability of or access to salmon by wildlife may adversely affect wildlife populations and, potentially, ecosystem-level processes. Thus, when the conservation of specific wildlife populations or healthy ecosystems is the management objective, allocation of salmon to wildlife should be considered. We provide an example of how such allocations could be calculated for a hypothetical bear population. Ultimately, salmon allocation for wildlife calls for integrated management of natural resources across agencies, across species, and across ecosystems. We summarize the current state of knowledge relative to the interaction between Pacific salmon and the terrestrial ecosystem, with special emphasis on the import of salmon to terrestrial wildlife and the import of wildlife to terrestrial and aquatic ecosystems.

Objective—To evaluate cardiac function parameters in a group of active and hibernating grizzly bears. Design—Prospective study. Animals—6 subadult grizzly bears. Procedure—Indirect blood pressure, a 12-lead ECG, and a routine echocardiogram were obtained in each bear during the summer active phase and during hibernation. Results—All measurements of myocardial contractility were significantly lower in all bears during hibernation, compared with the active period. Mean rate of circumferential left ventricular shortening, percentage fractional shortening, and percentage left ventricular ejection fraction were significantly lower in bears during hibernation, compared with the active period. Certain indices of diastolic function appeared to indicate enhanced ventricular compliance during the hibernation period. Mean mitral inflow ratio and isovolumic relaxation time were greater during hibernation. Heart rate was significantly lower for hibernating bears, and mean cardiac index was lower but not significantly different from cardiac index during the active phase. Contrary to results obtained in hibernating rodent species, cardiac index was not significantly correlated with heart rate. Conclusions and Clinical Relevance—Cardiac function parameters in hibernating bears are opposite to the chronic bradycardic effects detected in nonhibernating species, likely because of intrinsic cardiac muscle adaptations during hibernation. Understanding mechanisms and responses of the myocardium during hibernation could yield insight into mechanisms of cardiac function regulation in various disease states in nonhibernating species. (J Am Vet Med Assoc 2003;223:1170–1175)

Whitebark pine (Pinus albicaulis) is a masting species that produces relatively large, fat- and protein-rich nuts that are consumed by grizzly bears (Ursus arctos horribilis). Trees produce abundant nut crops in some years and poor crops in other years. Grizzly bear survival in the Greater Yellowstone Ecosystem is strongly linked to variation in pine-nut availability. Because whitebark pine trees are infected with blister rust (Cronartium ribicola), an exotic fungus that has killed the species throughout much of its range in the northern Rocky Mountains, we used stable isotopes to quantify the importance of this food resource to Yellowstone grizzly bears while healthy populations of the trees still exist. Whitebark pine nuts have a sulfur-isotope signature (9.2 ± 1.3‰ (mean ± 1 SD)) that is distinctly different from those of all other grizzly bear foods (ranging from 1.9 ± 1.7‰ for all other plants to 3.1 ± 2.6‰ for ungulates). Feeding trials with captive grizzly bears were used to develop relationships between dietary sulfur-, carbon-, and nitrogen-isotope signatures and those of bear plasma. The sulfur and nitrogen relationships were used to estimate the importance of pine nuts to free-ranging grizzly bears from blood and hair samples collected between 1994 and 2001. During years of poor pine-nut availability, 72% of the bears made minimal use of pine nuts. During years of abundant cone availability, 8 ± 10% of the bears made minimal use of pine nuts, while 67 ± 19% derived over 51% of their assimilated sulfur and nitrogen (i.e., protein) from pine nuts. Pine nuts and meat are two critically important food resources for Yellowstone grizzly bears.

Phillips and Koch (2002) suggested that the standard, linear mixing model used to estimate assimilated diets may produce inaccurate results when animals consume foods that vary markedly in their C:N ratios and elemental concentrations. They presented a concentration-dependent model and used published values on captive and freeranging bears to illustrate theoretical shortcomings of earlier, concentration-independent models. The assimilated diet estimates for Alaskan bears on the Kenai Peninsula using their concentration-dependent model differed markedly from previously published results (Jacoby et al. 1999). For example, the importance of salmon to brown bears (Ursus arctos) decreased from 50% to 26% of the assimilated diet and for black bears (Ursus americanus) allopatric to brown bears from 53% to zero. Estimating the importance of salmon to bears has profound conservation implications as salmon stocks become increasingly depleted by human activities. Brown bear populations with significant access to salmon are up to 50 times denser, adult females are twice as heavy, and litter sizes are 25% larger than in salmon-deprived populations (Hilderbrand et al. 1999c). Recently, brown bears have been identified as major conveyors of the marinederived nutrients in salmon to Alaskan riparian forests (Hilderbrand et al. 1999a; Helfield and Naiman 2001). Thus, the validity of the Phillips and Koch (2002) estimates must be evaluated as they note that incorrect assumptions in mixing models "may produce misleading results with major ramifications on assessments of feeding ecology and management strategies". The original dietary estimates of the importance of salmon by Jacoby et al. (1999) and Hilderbrand et al. (1999b, c) used a bear-specific, carbon-enrichment, linear mixing model (Hilderbrand et al. 1996) to separate marine (salmon) from terrestrial carbon sources. This approach is the most direct of all the models, has the fewest assumptions as fractionation effects related to digestion and substrate routing are part of the feeding trial results used to generate the original regression, and has been used previously to estimate the importance of salmon to earlier humans (Chisholm et al. 1982; Taylor et al. 1998). Phillips and Koch (2002) also used a concentration-independent mixing model to simultaneously estimate assimilated dietary salmon, terrestrial meat, and terrestrial plant content and "reached roughly the same conclusions" as Jacoby et al. (1999) and Hilderbrand et al. (1999b, c). Even though the above two estimates are in agreement and are derived from markedly different models and assumptions, Phillips and Koch (2002) insisted that the very different values generated by their concentration-dependent model that requires more inputs and assumptions are the correct values. The estimates from the Phillips and Koch (2002) concentration-dependent model are driven by the composition and C:N concentrations of the hypothesized assimilated diet, particularly the plant portion of the diet that was composed of equal parts apples (Malus sylvestris), blackberries (Rubus sp.), domestic cranberries (Vaccinium macrocarpon) and raspberries (Rubus sp.), and rhubarb stems (Rheum rhabarbarum). Despite extensive literature on the diets of bears, Phillips and Koch (2002) clearly made no effort to mimic a realistic plant diet in their modeling efforts as no Alaskan bear consumes their hypothetical diet. Although the actual plant names are irrelevant in illustrating their model, outputs of the model, comparisons to previous estimates, and management implications do depend on the diet chosen. The lack of realistic diet inputs in their example poses serious scientific and conservation concerns. The plant component of their hypothetical diet has a gross C:N ratio of 44:1 as compared to 4.6:1 and 3.7:1 for salmon and terrestrial meat (Table 1 in Phillips and Koch C.T. Robbins (X) School of Biological Sciences and Department of Natural Resource Sciences, Washington State University, Pullman, WA 99164-4236, USA e-mail: ctrobbins@wsu.edu Tel.: +1-509-3351119, Fax: +1-509-3353184

Adiponectin is the most abundant plasma adipokine, and is well known for its role in energy homeostasis and cardiac protection. In humans with dilated cardiomyopathy, myocardial adiponectin protein expression is reduced compared to normal hearts and has been implicated in the pathology of cardiomyopathy. Serum adiponectin levels are often conflicting, with higher levels associated with poor survival in humans with congestive heart failure (CHF). We evaluated adiponectin serum concentrations and myocardial protein expression in dogs with naturally occurring myxomatous mitral valve disease and CHF. We compared the findings to active and hibernating brown bears as bears are adapted to endure an extreme period of low cardiac output during their annual hibernation. Bears exhibited largely the active high-molecular weight (HMW) versus the low-molecular weight isoforms of myocardial adiponectin (HMW:LMW = 6.3) during both the active period and hibernation, while healthy dogs exhibited a more balanced mix of isoforms. Dogs with CHF expressed predominately HMW isoforms of adiponectin (HMW:LMW = 12.5), appearing more similar to bears. In contrast to humans, serum adiponectin was significantly lower in dogs with CHF and lowest levels in the severest CHF class. In both dogs and bears, myocardial adiponectin was expressed independent of circulating adiponectin concentrations, suggesting a local regulatory mechanism within the heart.

Grizzly bears (Ursus arctos horribilis) are inactive for up to 6 months during hibernation. They undergo profound seasonal changes in food intake, body mass, and energy expenditure. The circa-annual regulation of metabolism is poorly understood. In the present study, we measured plasma ghrelin, leptin, obestatin, and neuropeptide-Y (NPY) levels, hormones known to be involved in the regulation of energy homeostasis, in ten grizzly bears. Blood samples were collected during the active summer period, early hibernation and late hibernation. Plasma levels of leptin, obestatin, and NPY did not change between the active and the hibernation periods. Plasma total ghrelin and desacyl-ghrelin concentrations significantly decreased during the inactive winter period compared to summer levels. The elevated ghrelin levels may help enhance body mass during pre-hibernation, while the low plasma ghrelin concentrations during hibernation season may contribute to the maintenance of hypophagia, low energy utilization and behavioral inactivity. Our results suggest that ghrelin plays a potential role in the regulation of metabolic changes and energy homeostasis during hibernation in grizzly bears.

This is a Forum article in response to: Barber-Meyer, S. (2015) Trophic cascades from wolves to grizzly bears or changing abundance of bears and alternate foods? Journal of Animal Ecology, 83, doi: 10.1111/1365-2656.12338. We used multiple data sets and study areas as well as several lines of evidence to investigate potential trophic linkages in Yellowstone National Park. Our results suggest that a trophic cascade from wolves to elk to berry production to berry consumption by grizzly bears may now be underway in the Park.

Stable isotope signatures of lactating females and their nursing offspring were measured on 11 species, including herbivores, carnivores, hibernators, and non-hibernators. We hypothesized that: (1) nursing offspring would have stable isotope signatures that were a trophic level higher than their mothers, and (2) this pattern would be species-independent. The plasma of adult females had a δ15N enrichment over their diets of 4.1±0.7‰, but offspring plasma had a mean δ15N enrichment over maternal plasma of 0.9±0.8‰ and no C enrichment (0.0±0.6‰). The trophic level enrichment did not occur between mother and offspring because milk was depleted in both δ15N (1.0±0.5‰) and δ13C (2.1±0.9‰) relative to maternal plasma. Milk to offspring plasma enrichment was relatively small (δ15N enrichment of 1.9±0.7‰ and δ13C enrichment of 1.9±0.8‰) compared to the trophic level enrichment between the adults and their diets. While some species did have significant differences between the isotope signatures of mother and offspring, the differences were not related to whether they were hibernators or non-hibernators, carnivores or herbivores. Investigators wanting to use stable isotopes to quantify weaning or other lactation processes or diets of predators when both adults and nursing offspring are consumed must first establish the parameters that apply to a particular species/environment/diet combination.

Although well known as carnivores and not capable of digesting plant fiber, grizzly bears (Ursus arctos horribilis) consume over 200 species of plants and are entirely vegetarian in some ecosystems. Even in ecosystems with abundant meat resources, green vegetation can be an important seasonal food resource. Therefore, we examined the morphological, physiological, and environmental constraints that determine the nutritional value of herbaceous vegetation to grizzly bears. Short-term, board foraging trials were used with captive grizzly bears to determine constraints on intake rate including bite size, bite rate, bear size, plant species, plant height, and plant distribution. Feeding trials were conducted to determine the effect of protein level (12-35%) and digestible dry matter intake on weight gain. Finally, maximum daily intake, daily foraging time, and weight change were measured for captive bears foraging on highly abundant and nutritious forbs and grasses during 12-day trials. Intake during short-term board trials overestimated the intake of freely foraging bears from two- to seven-fold depending on bear size. Because of their relatively larger bite sizes, smaller absolute energy requirements, and relatively larger intake capacity, smaller bears (120 kg) made greater weight gains than very large bears on herbaceous vegetation. Smaller bears with ad libitum access to palatable, nutritious forbs gained weight at rates equal to wild bears. However, depending upon plant characteristics, bite sizes and available daily foraging time increasingly prevented large bears (120 kg) from gaining weight on herbaceous vegetation. Both captive and wild bears select forbs over grasses at similar growth stages because forbs are generally higher in protein and more digestible than grasses. Therefore, the nutritional well-being of wild grizzly bears could be improved in areas where bears are largely herbivorous and, thus, relatively small by purposefully managing for nutritious forbs.

The influence of seasonal dietary meat intake on changes in body mass and composition in wild and captive brown bears (Ursus arctos) was investigated because the importance and availability of meat to brown bear populations is currently an important management consideration in several North American ecosystems. Adult female brown bears on the Kenai Peninsula, Alaska, utilized meat heavily in both spring and fall. Meat accounted for 76.2 ± 26.0% (mean ± 1 SD; primarily moose carrion and calves) of assimilated carbon and nitrogen in the spring and 80.4 ± 22.2% (primarily salmon) in the fall. Mass increases in the spring (71.8 ± 28.2%) were mostly lean body mass, but increases in the fall (81.0 ± 19.5%) were primarily fat. Daily intake by captive brown bears fed meat ad libitum during 12-day trials was positively related to body mass. Mass change was positively related to intake in both seasons, but the composition of the gain varied by season, with spring gains primarily lean body mass (64.2 ± 9.4%), while fall gains were 78.8 ± 19.6% lipid. Absolute rates of gain by wild bears occasionally equaled, but were usually much less than, those of captive bears. This was likely due to a combination of factors, which included the time required to locate and handle meat resources, the limited availability of or access to meat resources, and (or) the duration of meat resource availability. Estimated intake by bears not feeding selectively on high-energy components of moose and salmon were 8.5 ± 1.5 kg/day and 541 ± 156 kg/year and 10.8 ± 4.6 kg/day and 1003 ± 489 kg/year, respectively. Intake would drop by as much as 58% for bears feeding exclusively on salmon roe. Management strategies for areas with brown bears that consume significant amounts of meat should address the perpetuation and availability of these meat resources.

We hypothesized that the relative availability of meat, indicated by contribution to the diet, would be positively related to body size and population productivity of North American brown, or grizzly, bears (Ursus arctos). Dietary contributions of plant matter and meat derived from both terrestrial and marine sources were quantified by stable-isotope analysis (δ13C and δ15N) of hair samples from 13 brown bear populations. Estimates of adult female body mass, mean litter size, and population density were obtained from two field studies of ours and from other published reports. The populations ranged from largely vegetarian to largely carnivorous, and food resources ranged from mostly terrestrial to mostly marine (salmon, Oncorhynchus spp.). The proportion of meat in the diet was significantly correlated with mean adult female body mass (r = 0.87, P < 0.01), mean litter size (r = 0.72, P < 0.01), and mean population density (r = 0.91, P < 0.01). Salmon was the most important source of meat for the largest, most carnivorous bears and most productive populations. We conclude that availability of meat, particularly salmon, greatly influences habitat quality for brown bears at both the individual level and the population level.

Subadult bears were studied during their autumn hyperphagia (n = 3) and winter dormancy (n = 6). Urea kinetics were measured with14C- and15N-urea, protein turnover was estimated with15N-glycine, and body composition was assessed with3H-water. Reduced amino acid degradation in winter was indicated by declines in plasma urea and aminotransferase activities, and lower urea production than in autumn (4.7 vs. 27.5 mmol urea-N∙kg−0.75∙d−1). Only 7.5% of urea produced in hyperphagic bears was degraded and just 1.1% of the degraded N reutilized as amino-N. Dormant bears reutilized 99.7% of urea produced, indicating thorough microbial ureolysis and urea-N resorption. Low rates of body N loss during dormancy suggested losses of non-urea N as creatinine. Protein turnover rates (15.2–21.5 g∙kg−0.75∙d−1) were similar between seasons and reflected the apparent maintenance of hepatic, intestinal, and muscular functions through dormancy. Protein synthesis accounted for 32% of energy expended in dormancy, which was mainly (91.5%) derived from fat oxidation. Consistent organ function and body temperature in dormant bears enables recycling of urea-N, which minimizes body protein loss and conserves mobility. In comparison with heterothermic hibernation, ursid dormancy would provide greater flexibility during winter and facilitate rapid resumption of foraging and growth in spring.

Bears consuming wild fruits for fall energy accumulation are constrained by several factors, including intake rate, the physiological capacity of the gastrointestinal tract, and the metabolic efficiency of gain in body mass. We measured these relationships through foraging and feeding trials using captive and wild black bears (Ursus americanus) and grizzly bears (Ursus arctos). Four fruit types covering a range of sizes and clustering were offered to captive bears to determine the effect of density, size, and presentation on intake rate. Intake rate (in grams per minute) and bite rates (in bites per minute) increased cur- vilinearly with increasing fruit density in singly spaced fruits. Maximum intakes ranged from 30 g/min for 0.5-g berries to >200 g/min for 4.2-g fruits. The highest bite rates were obtained during the initial encounter with each patch as bears consumed all visually apparent fruits on the surface. Bite rates quickly dropped by 15-20% as foraging continued within the patch. Maximum bite rates were not depressed until initial fruit density fell to 100 kg. We concluded that large bears, such as grizzlies, must depend on plants that permit large bite sizes or high bite rates through fruit clustering and bush configuration that reduce leaf-to-fruit ratios.

Historically, moose have been difficult to maintain in captivity when on diets of grass or legume hays and grain due to enteritis that frequently leads to chronic diarrhea/wasting disease. The development of wood-fiber diets has increased the lifespan of moose in captivity, but these diets do not completely prevent chronic wasting. Purina Mills (St. Louis, MO) hypothesized that captive moose are unable to digest starch that escapes the rumen, and therefore abnormal bacterial fermentation in the hindgut causes chronic diarrhea. An earlier study found no evidence of a digestive problem, so we tested the hypothesis that moose have difficulty metabolizing excess propionate produced from the fermentation of starch found in traditional cervid rations and high-grain wood-fiber diets. When challenged with an i.v. propionate load, moose metabolized propionate similar to healthy mule deer and domestic livestock. We then tested the hypothesis that grass forage is an initiating factor to chronic diarrhea/wasting and further hypothesized that grass, alfalfa, and other agriculture-based forages in association with an anaerobic bacteria produce inflammatory bowel disease (IBD) in moose. Captive moose that had ad libitum access to a wood-fiber pelleted moose diet and grazed in grass pastures developed chronic wasting symptoms at 2–4 years of age and died at 4.7 ± 0.3 years unless restricted from grass before the development of advanced symptoms. We isolated Bacteroides vulgatus in the feces and successfully treated a moose with chronic diarrhea/wasting disease with long-term metronidazole therapy, suggesting that the chronic enteritis causing wasting disease arises from a bacteria-associated defective immunosuppressive response similar to IBD in other species. Further support for the IBD cause of wasting in moose is that this animal will relapse within hours if the metronidazole treatment is discontinued even after many months. We developed a highly palatable high-fiber, low-starch moose ration that can be fed as the sole source of nourishment, although additional research and dietary improvements are required. Zoo Biol 16:479–494, 1997. © 1997 Wiley-Liss, Inc.

Safe and effective immobilization of grizzly bears (Ursus arctos) is essential for research and management. Fast induction of anesthesia, maintenance of healthy vital rates, and predictable recoveries are priorities. From September 2010 to May 2012, we investigated these attributes in captive and wild grizzly bears anesthetized with a combination of a reversible α2 agonist (dexmedetomidine [dexM], the dextrorotatory enantiomer of medetomidine) and a nonreversible N-methyl-d-aspartate (NMDA) agonist and tranquilizer (tiletamine and zolazepam [TZ], respectively). A smaller-than-expected dose of the combination (1.23 mg tiletamine, 1.23 mg zolazepam, and 6.04 µg dexmedetomidine per kg bear) produced reliable, fast ataxia (3.7 ± 0.5 min, x̄±SE) and workable anesthesia (8.1 ± 0.6 min) in captive adult grizzly bears. For wild bears darted from a helicopter, a dose of 2.06 mg tiletamine, 2.06 mg zolazepam, and 10.1 µg dexmedetomidine/kg produced ataxia in 2.5 ± 0.3 min and anesthesia in 5.5 ± 1.0 min. Contrary to published accounts of bear anesthesia with medetomidine, tiletamine, and zolazepam, this combination did not cause hypoxemia or hypoventilation, although mild bradycardia (50 beats per min) occurred in most bears during the active season. With captive bears, effective dose rates during hibernation were approximately half those during the active season. The time to first signs of recovery after the initial injection of dexMTZ was influenced by heart rate (P0.001) and drug dose (P0.001). Intravenous (IV) injection of the reversal agent, atipamezole, significantly decreased recovery time (i.e., standing on all four feet and reacting to the surrounding environment) relative to intramuscular injection. Recovery times (25 ± 8 min) following IV injections of the recommended dose of atipamezole (10 µg/µg of dexmedetomidine) and half that dose (5 µg/µg) did not differ. However, we recommend use of the full dose based on the appearance of a more complete recovery. Field trials confirmed that the dexMTZ + atipamezole protocol is safe, reliable, and predictable when administered to wild grizzly bears, especially during helicopter capture operations.

Many temperate zone animals adapt to seasonal changes by altering their physiology. This is mediated in large part by endocrine signals that encode day length and regulate energy balance and metabolism. The objectives of this study were to determine if the daily patterns of two important hormones, melatonin and cortisol, varied with day length in captive brown bears (Ursus arctos) under anesthetized and nonanesthetized conditions during the active (March-October) and hibernation periods. Melatonin concentrations varied with time of day and season in nonanesthetized female bears despite exceedingly low nocturnal concentrations (1-4 pg/mL) in the active season. In contrast, melatonin concentrations during hibernation were 7.5-fold greater than those during the summer in anesthetized male bears. Functional assessment of the pineal gland revealed a slight but significant reduction in melatonin following nocturnal light application during hibernation, but no response to beta-adrenergic stimulation was detected in either season. Examination of pineal size in two bear species bears combined with a phylogenetically corrected analysis of pineal glands in 47 other species revealed a strong relationship to brain size. However, pineal gland size of both bear species deviated significantly from the expected pattern. Robust daily plasma cortisol rhythms were observed during the active season but not during hibernation. Cortisol was potently suppressed following injection with a synthetic glucocorticoid. The results suggest that melatonin and cortisol both retain their ability to reflect seasonal changes in day length in brown bears. The exceptionally small pineal gland in bears may be the result of direct or indirect selection.

Cognitive bias tasks purport to assess affective states via responses to ambiguous stimuli. We hypothesized that a novel cognitive bias task based on positive reinforcement using quantity differences would detect changes in affect in captive grizzly bears (Ursus arctos horribilis). We trained bears (n = 8) to respond differently (nose or paw touch) to two stimuli (light or dark gray cue cards), with responses counterbalanced across bears. The two cues signaled a small or large food reward, respectively. Responses to ambiguous probe stimuli (i.e., shades of gray) intermediate to the trained stimuli were classified as either ‘optimistic,’ appropriate for the larger reward, or ‘pessimistic,’ appropriate for the smaller reward. In Experiment 1, we explored the contrast in reward size necessary to detect a change in response across probe stimuli (large reward, 3 or 6 apple slices: small reward, 1 slice). We observed a change in response across probe stimuli, with no difference in response between reward-value groups, indicating that a contrast of 3:1 apple slices was sufficient to affect responses. In Experiment 2, we investigated cognitive bias after 2.1 h of exposure to enrichment items varying in attractiveness. Results were unaffected by enrichment type or time spent interacting with enrichments, indicating that the task failed to demonstrate criterion validity for comparing mood following exposure to different enrichment items. However, greater time spent pacing prior to testing was associated with ‘optimistic’ judgments. The data provide some support for use of cognitive bias tasks based on quantity differences in animal welfare assessments involving captive wildlife.

The potential use of stable-isotope analyses (δ13C and δ15N) to estimate bear diets was assessed in 40-day feeding trials using American black bears (Ursus americanus). Bear plasma and red blood cells have half-lives of ~4 days and ~28 days, respectively. The isotopic signature of bear plasma is linearly related to that of the diet, and with the exception of adipose tissue, there is no isotopic fractionation across bear tissues. Isotopic analyses were used to estimate the diets of three bear populations: Pleistocene cave bears (U. speleaus) in Europe, grizzly bears (Ursus arctos horribilis) inhabiting the Columbia River drainage prior to 1931, and brown bears (U. arctos) of Chichagof and Admiralty islands, Alaska. Cave bears were omnivores with terrestrially produced meat contributing from 41 to 78% (58 ± 14%) of their metabolized carbon and nitrogen. Salmon contributed from 33 to 90% (58 ± 23%) of the metabolized carbon and nitrogen in grizzly bears from the Columbia River drainage. Finally, most brown bears on Chichagof and Admiralty islands feed upon salmon during the late summer and fall; however, a subpopulation of bears exists that does not utilize salmon.

Foraging efficiency (metabolizable energy intake/energy expenditure when foraging) was determined over a 2-year period in nine free-ranging Sitka black-tailed deer (Odocoileus hemionus sitkensis) in Alaska, and related to foraging-bout duration, distances travelled, and average speeds of travel. We calculated the energy-intake component from seasonal dry matter and energy content, dry matter digestibility, and a metabolizable energy coefficient for each plant species ingested. We estimated energy expenditures when foraging as the sum of energy costs of standing, horizontal and vertical locomotion, sinking depths in snow, and supplementary expenditures associated with temperatures outside thermoneutrality. Energy intake per minute averaged 4.0 times more in summer than winter; energy expenditure was 1.2 times greater in summer. Animals obtained higher amounts of metabolizable energy with higher amounts of energy invested. Energy intake during foraging bouts in summer was 2.5 times the energy invested; in contrast, energy intake during winter was only 0.7 times the energy expended. Changes in body mass of deer throughout the year increased asymptotically with foraging efficiency, driven primarily by the rate of metabolizable energy intake. Within a season, summer intake rates and winter rates of energy expediture had the greatest effects on the relation between foraging efficiency and mass status. Seasonal changes in foraging efficiency result in seasonal cycles in body mass and condition in black-tailed deer. Body reserves accumulated during summer, however, are essential for over-winter survival of north-temperate ungulates because energy demands cannot be met by foraging alone.

Milk composition and intake, cub growth, and mass loss of hibernating lactating and nonlactating American black bears (Ursus americanus) and grizzly bears (Ursus arctos horribilis) were investigated. Except for ash content, milk composition was similar between species. Lipid content varied only slightly throughout lactation, whereas carbohydrate content increased from 1 to 3% during hibernation before decreasing to less than 0.5% at the end of lactation. Protein and dry matter content increased from 6.6 ± 0.4 and 29.8 ± 3.9% during hibernation to 13.7 ± 1.1 and 34.4 ± 3.7% post hibernation, respectively. The ash content of black bear milk increased from 1% during hibernation to 2% after den emergence, but the ash content of grizzly bear milk (1.3 ± 0.1%) did not fluctuate. Mean milk intake and growth during hibernation were 185 ± 89 and 49 ± 9 g/day for black bear cubs and 353 ± 54 and 98 ± 22 g/day for grizzly bear cubs, respectively, which accounted for about 9% of the estimated yearly milk consumption by the cubs. Milk intake peaked during the summer at levels approximately 4 times higher than those occurring in the winter den. The mass lost by older, hibernating, nonlactating bears was proportional to their metabolic body mass and was almost exclusively lipid. The rate of mass loss by denning, lactating females relative to nonlactating bears was 45% higher for black bears and 95% higher for grizzly bears. The less costly black bear cub may be at an important competitive advantage when both species occupy nutritionally limited habitat.

Milk composition, neonatal milk intake, and growth of captive, maternally raised, singleton moose calves (Alces alces gigas) were measured during the first 4 months post partum. Milk intake rose to a peak of 4760 ± 1017 g/day at day 23 and then decreased with increasing calf age. Milk fat and crude protein levels declined until the time of peak milk output, and then increased. Ash content remained relatively constant whereas carbohydrate content diminished as lactation progressed. At peak output, milk averaged 20.5 ± 1.5% dry matter, 7.9 ± 1.5% fat, 7.2 ± 0.4% protein, 1.4 ± 0.1% ash, 3.7 ± 0.2% carbohydrate, and 1.29 ± 0.13 kcal/g wet mass (1 kcal = 4.18 kJ). The growth rates of maternally raised calves during the first 30 days (785 ± 115 g/day) were higher than those reported for bottle-raised calves.

The measurement of body composition in black bears (Ursus americanus), brown bears (U. arctos), and polar bears (U. maritimus) was investigated by means of isotopic water dilution and bioelectrical impedance analysis (BIA). The basic relationships between body lipid, water, protein, and ash were determined by direct chemical measurement of 13 black and 6 brown bears. Body water and lipid content as a percentage are highly correlated (r2 = 0.98, standard error of the estimate (SEE) = 1.1%) and inversely proportional. The dry, lipid-free mass averaged 83.5 ± 1.6% protein and 16.5 ± 1.6% ash. Either isotopic water dilution or BIA can be used to estimate body lipid content of healthy, uninjured bears (r2 = 0.93, SEE = 2.7% and r2 = 0.96, SEE = 2.2%, respectively). Isotopic water equilibrated with body water by 150 min. Abscesses and recent injuries (i.e., gunshot or snare wounds) produced erroneous body composition estimates when BIA was used, but only when the injury was in the conductor path between the BIA electrodes. Dilution estimates were not affected by injuries. Currently, neither method can be used on dead bears.

Reproductive physiology in North American ursids is characterized by mating from spring to early summer, delayed implantation, and birth during hibernation. During spring 2008, a captive adult female brown bear was mated with two adult males. Pregnancy was determined by elevated progesterone concentrations during late fall before hibernation. Two male cubs were born on December 31, 2008, and a third female cub was born 17 days later on January 16. All were successfully raised and all were confirmed to have identical paternity. When normalized to age, cub growth rates did not differ. To our knowledge, this is the first documented case of markedly different birth dates in a single litter of brown bear cubs.

Captive mule deer (Odocoileus hemionus hemionus) and black-tailed deer (O. h. sitkensis) were used in cafeteria-type, two choice feeding trials to test the hypothesis that digestible dry matter (energy) and nontannin phenolics of tree, shrub, and forb leaves are major determinants of diet preference. Deer selected plants in relation to a trade-off between the benefit derived from digestible dry matter and the cost of nontannin phenolics presumably associated with toxicity when absorbed. When one of the forages contained both the highest digestible dry matter and lowest nontannin phenolics, the deer always preferred that plant. When one forage had the highest digestible dry matter but the other plant had the lowest nontannin phenolics, the deer selected the high-energy plant when the difference in nontannin phenolics was relatively small, but they preferred the low-energy plant when the other forage had much higher levels of nontannin phenolics. Tannins influenced diet choice only as one of the factors reducing digestible dry matter in these forages. Apparently total dry-matter intake was constrained by the nontannin phenolic fraction but not by tannins. Tannins and nontannin phenolics both contribute to defending plants against browsers.

Body mass, body composition, and water transfer rates were determined over a continuous 2-year period in nine free-ranging Sitka black-tailed deer (Odocoileus hemionus sitkensis). Body masses showed a cyclical pattern, declined 14 – 31% between October and March, and were best described by a five-parameter, biologically based regression model. The amount of mass lost by black-tailed deer over winter depended on the peak body mass attained during fall. During winter, animals used 70 – 82% of their body fat and 10 – 15% of their protein reserves. Body fat was preferentially mobilized at rates 2.3 – 11.6 times higher than protein. Because of the higher energy content of fat, fat accounted for 83 – 92% of the catabolized energy. Water transfer rates varied seasonally with average summer values approximately four times those of winter; lactating deer had significantly higher rates of water transfer than nonlactating animals. Seasonal changes in all of the above parameters for wild deer were greater than those for eight deer of the same age in captivity.