Field Metabolic Rate and PCB Adipose Tissue Deposition Efficiency in East Greenland Polar Bears Derived from Contaminant Monitoring Data

Climate change will increasingly affect the natural habitat and diet of polar bears (Ursus maritimus). Understanding the energetic needs of polar bears is therefore important. We developed a theoretical method for estimating polar bear food consumption based on using the highly recalcitrant polychlorinated biphenyl (PCB) congener, 2,2′,4,4′,55-hexaCB (CB153) in bear adipose tissue as an indicator of food intake. By comparing the CB153 tissue concentrations in wild polar bears with estimates from a purposely designed individual-based model, we identified the possible combinations of field metabolic rates (FMR) and CB153 deposition efficiencies in East Greenland polar bears. Our simulations indicate that if 30% of the CB153 consumed by polar bear individuals were deposited into their adipose tissue, the corresponding FMR would be only two times the basal metabolic rate. In contrast, if the modelled CB153 deposition efficiency were 10%, adult polar bears would require six times more energy than that needed to cover basal metabolism. This is considerably higher than what has been assumed for polar bears in previous studies though it is similar to FMRs found in other marine mammals. An implication of this result is that even relatively small reductions in future feeding opportunities could impact the survival of East Greenland polar bears.     

The effect of lower limb length on the energetic cost of locomotion: implications for fossil hominins

The consequences of the relatively short lower limbs characteristic of AL 288-1 have been widely discussed, as have the causes and consequences of the short limbs of Neanderthals. Previous studies of the effect of limb length on the energetic cost of locomotion have reported no relationship; however, limb length could have accounted for as much as 19% of the variation in cost and gone undetected (Steudel and Beattie, 1995; Steudel, 1994, 1996). Kramer (1999) and Kramer and Eck (2000) have recently used a theoretical model to predict the effect of the shorter limbs of early hominids, concluding that the shorter limbs may actually have been energetically advantageous. Here, we took an experimental approach. Twenty-one human subjects, of varying limb lengths, walked on a treadmill at 2.6, 2.8, 3.0 and 3.2m.p.h., while their expired gases were analyzed. The subjects walked for 12 minutes at each speed and their rates of oxygen consumption (VO2) over four minutes were averaged to estimate VO2. We also measured each subject's height, weight and lower limb length. Lean body mass and % fat were determined using dual-energy x-ray absorptiometry. ANCOVA with total VO2 at either speed as the dependent variable and total lean mass, % fat and lower limb length as covariates resulted in all three covariates having a significant positive effect on VO2 at p<0.01. Subjects with relatively longer lower limbs had lower locomotor costs. Thus the short lower limbs characteristic of some hominid taxa would have resulted in more costly locomotion, barring some physiological anomaly. The magnitude of this effect is substantial; Neanderthals are estimated to have had locomotor costs 30% greater than those of contemporary anatomically modern humans. By contrast the increase in lower limb length seen in H. erectus would have mitigated the increase in locomotor costs produced by the increase in body size.     

The energetic cost of locomotion: humans and primates compared to generalized endotherms

A wide range of selective pressures have been advanced as possible causes for the adoption of bipedalism in the hominin lineage. One suggestion has been that because modern human walking is relatively efficient compared to that of a typical quadruped, the ancestral quadruped may have reaped an energetic advantage when it walked on two legs. While it has become clear that human walking is relatively efficient and human running inefficient compared to "generalized endotherms", workers differ in their opinion of how the cost of human bipedal locomotion compares to that of a generalized primate walking quadrupedally. One view is that human walking is particularly efficient in comparison to other primates. The present study addresses this by comparing the cost of human walking and running to that of the eight primate species for which data are available and by comparing cost in primates to that of a "generalized endotherm". There is no evidence that primate locomotion is more costly than that of a generalized endotherm, although more data on adult Old World monkeys and apes would be useful. Further, human locomotion does not appear to be particularly efficient relative to that of other primates.     

The evolution of human running: effects of changes in lower-limb length on locomotor economy

Previous studies have differed in expectations about whether long limbs should increase or decrease the energetic cost of locomotion. It has recently been shown that relatively longer lower limbs (relative to body mass) reduce the energetic cost of human walking. Here we report on whether a relationship exists between limb length and cost of human running. Subjects whose measured lower-limb lengths were relatively long or short for their mass (as judged by deviations from predicted values based on a regression of lower-limb length on body mass) were selected. Eighteen human subjects rested in a seated position and ran on a treadmill at 2.68 ms(-1) while their expired gases were collected and analyzed; stride length was determined from videotapes. We found significant negative relationships between relative lower-limb length and two measures of cost. The partial correlation between net cost of transport and lower-limb length controlling for body mass was r=-0.69 (p=0.002). The partial correlation between the gross cost of locomotion at 2.68 ms(-1) and lower-limb length controlling for body mass was r=-0.61 (p=0.009). Thus, subjects with relatively longer lower limbs tend to have lower locomotor costs than those with relatively shorter lower limbs, similar to the results found for human walking. Contrary to general expectation, a linear relationship between stride length and lower-limb length was not found.     

Brown bears selectively kill salmon with higher energy content but only in habitats that facilitate choice

Pacific salmon return to spawn in thousands of streams across the Pacific Rim, from large rivers to tiny headwater streams. Once on the spawning grounds, salmon undergo dramatic biochemical changes as they metabolize stored lipid and protein reserves; at stream entrance, they will contain up to 85% more lipid and 40% more protein than at their senescent death a week or two later. Foraging brown and black bears that congregate at spawning streams thus encounter salmon that vary dramatically in their energy content and thus energetic reward. We hypothesized that bears would selectively kill salmon that are highest in energy content (fewest number of days on the spawning grounds) when they pursue salmon at small shallow streams where little effort is necessary to capture salmon, i.e. habitats that facilitate choice. In contrast, bears in environments where foraging is difficult (deeper, more complex streams) should be less selective and should capture salmon that are most available. We tested these ideas by examining predation rates on fish of different in-stream ages (i.e. energy content) at three different streams that varied in physical habitat attributes. At a very shallow, simple stream, bears preferentially killed salmon that had spent the fewest days in the stream. At two streams where deeper water and woody debris provided refuges for salmon, predation rates increased with in-stream age. At the shallowest streams encounter rates and capture success are likely equal among the high- and low-energy salmon and thus predation rates reflect active choice by bears. In contrast capture success probably increases on the older salmon at the larger streams (due to a loss of vigor), and thus 'preference' for these fish increases due to decreasing effort necessary to capture them.     

Does limb length predict the relative energetic cost of locomotion in mammals?

Do relatively longer limbs result in a lower energetic cost of locomotion? To determine whether or not cost is correlated with limb length in some way other than that due to their respective relationships to body mass, we have removed the effects of size by calculating the residuals of the relationship between each character and body size. We then regressed the pairs of residuals on one another. Because biological variables do not occur in a series of units that have evolved independently, the degree of divergence of two species is likely to be influenced by the length of time since they last shared a common ancestor. We therefore corrected for the phylogenetic relatedness of species. Data on the energetic cost of locomotion of a wide variety of species were taken from published sources. Data on limb lengths were taken from specimens in various museum collections which were similar in body mass (± 12%) to the specimens on which the cost measurements were made. None of the correlations between the residuals of either fore- or hindlimb length and neither of two estimates of the cost of locomotion was significant at P = 0.05. It is concluded that limb length does not importantly influence an animal's locomotor efficiency. These results do not imply the lack of a close relationship between cost and stride length.     

The energetic cost of various behaviors in the laboratory mouse

The continuous 12-hr observation of a mouse placed in a respiratory chamber has made it possible to record the succession of behaviors of the animal, as well as the associated variations of CO2 concentrations in the chamber. Ten behaviors have been considered; these included rest, locomotion, sniffing, feeding, drinking, nest building and various types of grooming. These data have been used for the estimation, by means of Kalman filtering, of the energetic cost of each behavior. Thus, at 20 degrees C, these costs vary between 1.51 ml CO2/g/hr (rest) and 4.90 ml CO2/g/hr (locomotion). These costs seem independent of any underlying biological rhythm; they yield basal metabolic estimates and average daily metabolic rates similar to those found in the literature. The low energetic cost of each behavioral bout, as compared to the daily energy intake of the mouse, leads one to believe that the behavioral transitions of this animal are not to be ascribed to energetical reasons. These results have been validated with data obtained from two other mice under similar conditions.     

Energetics of walking and running: insights from simulated reduced-gravity experiments.

On Earth, a person uses about one-half as much energy to walk a mile as to run a mile. On another planet with lower gravity, would walking still be more economical than running? When people carry weights while they walk or run, energetic cost increases in proportion to the added load. It would seem to follow that if gravity were reduced, energetic cost would decrease in proportion to body weight in both gaits. However, we find that under simulated reduced gravity, the rate of energy consumption decreases in proportion to body weight during running but not during walking. When gravity is reduced by 75%, the rate of energy consumption is reduced by 72% during running but only by 33% during walking. Because reducing gravity decreases the energetic cost much more for running than for walking, walking is not the cheapest way to travel a mile at low levels of gravity. These results suggest that the link between the mechanics of locomotion and energetic cost is fundamentally different for walking and for running.     

Competition for space: Effects of disturbance and indeterminate competitive success

Organisms occupying and competing for space appear to coexist indefinitely on a relatively homogeneous and limited resource. Armstrong (1976) developed a mathematical model showing how fugitive species, with high recruitment rates and low interference competitive ability, can coexist with dominant species, those with high competitive ability but lower rates of recruitment. This and similar models of coexistence have not considered the case where competitive success is indeterminate, that is, where the poorer competitor can sometimes win competitive encounters.Armstrong's model and others considering coexistence of more than two species (e.g., Hastings, 1980) predict coexistence of multiple species at intermediate rates of disturbance and monopolization of space by single species at either very low or very high disturbance rates. This result is not true for a three-species modification of Armstrong's model. Under certain parameter values, a single species can monopolize space at intermediate disturbance rates as well.     

Limb morphology,bipedal gait,and the energetics of hominid locomotion

How viable is the argument that increased locomotor efficiency was an important agent in the origin of hominid bipedalism? This study reviews data from the literature on the cost of human bipedal walking and running and compares it to data on quadrupedal mammals including several non-human primate species. Literature data comparing the cost of bipedal and quadrupedal locomotion in trained capuchin monkeys and chimpanzees are also considered. It is concluded that increased energetic efficiency would not have accrued to early bipeds. Presumably, however, selection for improved efficiency in the bipedal stance would have occurred once the transition was made. Would such a process have included selection for increased limb length? Data on the cost of locomotion vs. limb length reveal no significant relationship between these variables in 21 species of mammals or in human walking or running. © 1996 Wiley-Liss, Inc.     

Activity patterns of brown bears (Ursus arctos) in Slovenia and Croatia

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

Locomotor energetics in primates: gait mechanics and their relationship to the energetics of vertical and horizontal locomotion

All primates regularly move within three-dimensional arboreal environments and must often climb, but little is known about the energetic costs of this critical activity. Limited previous work on the energetics of incline locomotion suggests that there may be differential selective pressures for large compared to small primates in choosing to exploit a complex arboreal environment. Necessary metabolic and gait data have never been collected to examine this possibility and biomechanical mechanisms that might explain size-based differences in the cost of arboreal movement. Energetics and kinematics were collected for five species of primate during climbing and horizontal locomotion. Subjects moved on a treadmill with a narrow vertical substrate and one with a narrow horizontal substrate at their maximum sustainable speed for 10–20 min while oxygen consumption was monitored. Data during climbing were compared to those during horizontal locomotion and across size. Results show that climbing energetic costs were similar to horizontal costs for small primates (<0.5 kg) but were nearly double for larger species. Spatio-temporal gait characteristics suggest that the relationship between the cost of locomotion and the rate of force production changes between the two locomotor modes. Thus, the main determinants of climbing costs are fundamentally different from those during horizontal locomotion. These new results combining spatiotemporal and energetic data confirm and expand on our previous argument (Hanna et al.: Science 320 (2008) 898) that similar costs of horizontal and vertical locomotion in small primates facilitated the successful occupation of a fine-branch arboreal milieu by the earliest primates.     

Alaskan brown bears (Ursus arctos) aggregate and display fidelity to foraging neighborhoods while preying on Pacific salmon along small streams

The interaction between brown bears (Ursus arctos) and Pacific salmon (Oncorhynchus spp.) is important to the population dynamics of both species and a celebrated example of consumer‐mediated nutrient transport. Yet, much of the site‐specific information we have about the bears in this relationship comes from observations at a few highly visible but unrepresentative locations and a small number of radio‐telemetry studies. Consequently, our understanding of brown bear abundance and behavior at more cryptic locations where they commonly feed on salmon, including small spawning streams, remains limited. We employed a noninvasive genetic approach (barbed wire hair snares) over four summers (2012–2015) to document patterns of brown bear abundance and movement among six spawning streams for sockeye salmon, O. nerka, in southwestern Alaska. The streams were grouped into two trios on opposite sides of Lake Aleknagik. Thus, we predicted that most bears would forage within only one trio during the spawning season because of the energetic costs associated with swimming between them or traveling around the lake and show fidelity to particular trios across years because of the benefits of familiarity with local salmon dynamics and stream characteristics. Huggins closed‐capture models based on encounter histories from genotyped hair samples revealed that as many as 41 individuals visited single streams during the annual 6‐week sampling season. Bears also moved freely among trios of streams but rarely moved between these putative foraging neighborhoods, either during or between years. By implication, even small salmon spawning streams can serve as important resources for brown bears, and consistent use of stream neighborhoods by certain bears may play an important role in spatially structuring coastal bear populations. Our findings also underscore the efficacy of noninvasive hair snagging and genetic analysis for examining bear abundance and movements at relatively fine spatial and temporal scales.     

Snowpack,fire, and forest disturbance: interactions affect montane invasions by non‐native shrubs

Montane regions worldwide have experienced relatively low plant invasion rates, a trend attributed to increased climatic severity, low rates of disturbance, and reduced propagule pressure relative to lowlands. Manipulative experiments at elevations above the invasive range of non‐native species can clarify the relative contributions of these mechanisms to montane invasion resistance, yet such experiments are rare. Furthermore, global climate change and land use changes are expected to cause decreases in snowpack and increases in disturbance by fire and forest thinning in montane forests. We examined the importance of these factors in limiting montane invasions using a field transplant experiment above the invasive range of two non‐native lowland shrubs, Scotch broom (Cytisus scoparius) and Spanish broom (Spartium junceum), in the rain–snow transition zone of the Sierra Nevada of California. We tested the effects of canopy closure, prescribed fire, and winter snow depth on demographic transitions of each species. Establishment of both species was most likely at intermediate levels of canopy disturbance, but at this intermediate canopy level, snow depth had negative effects on winter survival of seedlings. We used matrix population models to show that an 86% reduction in winter snowfall would cause a 2.8‐fold increase in population growth rates in Scotch broom and a 3.5‐fold increase in Spanish broom. Fall prescribed fire increased germination rates, but decreased overall population growth rates by reducing plant survival. However, at longer fire return intervals, population recovery between fires is likely to keep growth rates high, especially under low snowpack conditions. Many treatment combinations had positive growth rates despite being above the current invasive range, indicating that propagule pressure, disturbance, and climate can all strongly affect plant invasions in montane regions. We conclude that projected reductions in winter snowpack and increases in forest disturbance are likely to increase the risk of invasion from lower elevations.     

Capture, anesthesia, and disturbance of free-ranging brown bears (Ursus arctos) during hibernation

We conducted thirteen immobilizations of previously collared hibernating two- to four-year-old brown bears (Ursus arctos) weighing 21-66 kg in central Sweden in winter 2010 and 2011 for comparative physiology research. Here we report, for the first time, an effective protocol for the capture and anesthesia of free-ranging brown bears during hibernation and an assessment of the disturbance the captures caused. Bears were darted in anthill, soil, or uprooted tree dens on eleven occasions, but two bears in rock dens fled and were darted outside the den. We used medetomidine at 0.02-0.06 mg/kg and zolazepam-tiletamine at 0.9-2.8 mg/kg for anesthesia. In addition, ketamine at 1.5 mg/kg was hand-injected intramuscularly in four bears and in six it was included in the dart at 1.1-3.0 mg/kg. Once anesthetized, bears were removed from the dens. In nine bears, arterial blood samples were analyzed immediately with a portable blood gas analyzer. We corrected hypoxemia in seven bears (PaO(2) 57-74 mmHg) with supplemental oxygen. We placed the bears back into the dens and antagonized the effect of medetomidine with atipamezole. Capturing bears in the den significantly increased the risk of den abandonment. One of twelve collared bears that were captured remained at the original den until spring, and eleven, left their dens (mean ± standard deviation) 3.2±3.6 (range 0.5-10.5) days after capture. They used 1.9±0.9 intermediate resting sites, during 6.2±7.8 days before entering a new permanent den. The eleven new permanent dens were located 730±589 m from the original dens. We documented that it was feasible and safe to capture hibernating brown bears, although they behaved differently than black bears. When doing so, researchers should use 25% of the doses used for helicopter darting during the active period and should consider increased energetic costs associated with den abandonment.     

Mass loss in incubating Eurasian dotterel: adaptation or constraint?

Body mass loss is frequently observed in breeding birds: whether this is an adaptive response to a change in the relative value of body stores and locomotion performance or a consequence of energetic constraint is still in debate. The male alone cares for most nests of the Eurasian dotterel Charadrius morinellus , although females assist at a proportion of nests. Energetic costs are probably high in the dotterel's arctic-alpine environment and uniparental care restricts the foraging time available to meet these costs, so that incubating dotterel may have to fuel themselves partly using body stores. Nesting male dotterel lost 7.8% of their mass through the incubation period but were 6.8% heavier during periods of high food abundance. Males that were assisted in incubation by a female were 6.7% heavier than uniparental males. We conclude that, since dotterel were heavier when energetic constraints were lifted, mass loss through incubation was principally a consequence of energetic constraint, rather than adaptive mass optimisation.     

Bioenergetics and the origin of hominid bipedalism

Compared to most quadrupedal mammals, humans are energetically inefficient when running at high speeds. This fact can be taken to mean that human bipedalism evolved for reasons other than to reduce relative energy cost during locomotion. Recalculation of the energy expended during human walking at normal speeds shows that (1) human bipedalism is at least as efficient as typical mammalian quadrupedalism and (2) human gait is much more efficient than bipedal or quadrupedal locomotion in the chimpanzee. We conclude that bipedalism bestowed an energetic advantage on the Miocene hominoid ancestors of the Hominidae.     

GRIZZLY BEAR DEMOGRAPHICS IN AND AROUND BANFF NATIONAL PARK AND KANANASKIS COUNTRY,ALBERTA

Abstract: The area in and around Banff National Park (BNP) in southwestern Alberta, Canada, is 1 of the most heavily used and developed areas where grizzly bears (Ursus arctos) still exist. During 1994–2002, we radiomarked and monitored 37 female and 34 male bears in this area to estimate rates of survival, reproduction, and population growth. Annual survival rates of bears other than dependent young averaged 95% for females and 81–85% for males. Although this area was largely unhunted, humans caused 75% of female mortality and 86% of male mortality. Females produced their first surviving litter at 6–12 years of age (x̄ = 8.4 years). Litters averaged 1.84 cubs spaced at 4.4-year intervals. Adult (≥6-years-old) females produced 0.24 female cubs per year and were expected to produce an average of 1.7 female cubs in their lifetime, based on rates of reproduction and survival. Cub survival was 79%, yearling survival was 91%, and survival through independence at 2.5–5.5 years of age was 72%, as no dependent young older than yearlings died. Although this is the slowest-reproducing grizzly bear population yet studied, high rates of survival seem to have enabled positive population growth (Λ = 1.04, 95% CI = 0.99–1.09), based on analyses using Leslie matrices. Current management practices, instituted in the late 1980s, focus on alleviating human-caused bear mortality. If the 1970–1980s style of management had continued, we estimated that an average of 1 more radiomarked female would have been killed each year, reducing female survival to the point that the population would have declined.     

Time budgets of Snow Geese Chen caerulescens and Ross's Geese Chen rossii in mixed flocks: implications of body size, ambient temperature and family associations

Body size affects foraging and forage intake rates directly via energetic processes and indirectly through interactions with social status and social behaviour. Ambient temperature has a relatively greater effect on the energetics of smaller species, which also generally are more vulnerable to predator attacks than are larger species. We examined variability in an index of intake rates and an index of alertness in Lesser Snow Geese Chen caerulescens caerulescens and Ross's Geese Chen rossii wintering in southwest Louisiana. Specifically we examined variation in these response variables that could be attributed to species, age, family size and ambient temperature. We hypothesized that the smaller Ross's Geese would spend relatively more time feeding, exhibit relatively higher peck rates, spend more time alert or raise their heads up from feeding more frequently, and would respond to declining temperatures by increasing their proportion of time spent feeding. As predicted, we found that Ross's Geese spent more time feeding than did Snow Geese and had slightly higher peck rates than Snow Geese in one of two winters. Ross's Geese spent more time alert than did Snow Geese in one winter, but alert rates differed by family size, independent of species, in contrast to our prediction. In one winter, time spent foraging and walking was inversely related to average daily temperature, but both varied independently of species. Effects of age and family size on time budgets were generally independent of species and in accordance with previous studies. We conclude that body size is a key variable influencing time spent feeding in Ross's Geese, which may require a high time spent feeding at the expense of other activities.     

Grizzly Bear Behavior and Global Positioning System Collar Fix Rates

Abstract: Animal locations collected by Global Positioning System (GPS) collars will represent a biased sample of the sites an animal used if some position fixes fail and if those missed locations do not occur randomly. Probability of a GPS receiver obtaining a position fix is known to decline as canopy cover increases, but the impact of forest canopy cover was insufficient to account for the low fix rates we observed for GPS collars on grizzly bears (Ursus arctos). We tested the hypothesis that GPS fix rates were related to the interaction between animal activity (active vs. resting) and canopy cover by evaluating the following predictions: 1) grizzly bear activity should follow a circadian pattern similar to the circadian fix-rate pattern, 2) grizzly bear use of canopy cover should follow a circadian pattern similar to the circadian fix rates, 3) grizzly bear activity should be related to canopy cover (i.e., bears should rest in areas with relatively high canopy covers and feed and move in relatively open areas), and 4) collar orientation and canopy cover should interact to affect the fix rates of test collars. The GPS fix rates traced a bimodal circadian pattern that was directly related to the circadian pattern of grizzly bear activity. Fix rates declined when bears were more likely to be using denser cover, and fix rates of test collars demonstrated that collar orientation interacted with canopy cover, such that fix rates declined much more with increasing canopy cover when the collar was on its side than when the collar was upright. We concluded that inferences made about grizzly bear microhabitat use, based on GPS locations, will underrepresent high canopy cover sites, especially when grizzly bears are resting there. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):596–602; 2008)