Geology topics

Anthony M. Pagano

Publications and source records attributed to Anthony M. Pagano.

At least 19 recordsLinked to original sources

USGS2025

The effects of carnivory and herbivory on the energy balance of Arctic grizzly bears

Omnivores often face tradeoffs between selecting for spatially dispersed energy-dense vertebrate prey versus densely distributed herbivorous resources that have limited energetic value per unit intake. Arctic grizzly bears ( Ursus arctos ) are large omnivores within a resource-limited ecosystem that are known to exhibit smaller body masses and occur at lower densities than grizzly bears in other regions of North America. We evaluated the energy balance of Arctic grizzly bears during a portion of the fall hyperphagic period in two ecologically differing regions on Alaska’s northern Arctic coast by monitoring mass change, food intake, activity, and energy expenditure of 12 individuals over 17–22 days. Bears in coastal areas were more carnivorous than bears in the foothills that were predominantly herbivorous and frugivorous. Carnivory was associated with greater movement, body fat, and energy expenditure and two of four carnivorous bears lost mass. Overall, the mean body fat of the bears in this study was 34% lower than other grizzly bear populations in North America in the fall. Furthermore, the bears in this study exhibited relatively small changes in body mass (x̄ = 3%, range =−2 to 11%) that were 60% lower than other grizzly bear populations which typically gain substantial mass in the fall in preparation for denning. Our results, while representing a snapshot from a small number of bears during the fall hyperphagic period, are consistent with previous studies and indicate limited availability of energy-dense food resources during this time for grizzly bears in this region of the Arctic.

Alaska↗

USGS2025

Assessing the combined influence of biotic and anthropogenic stressors on polar bears to inform conservation planning

Developing conservation strategies for species vulnerable to the effects of climate change, like polar bears ( Ursus maritimus ), can be challenging given the uncertainty of future environmental conditions. Effective conservation planning requires identifying and ranking threats to the persistence of polar bears throughout their circumpolar range and then assessing the ability of mitigative actions to aid in meeting plan objectives. We used a Bayesian network model to (1) characterize the relative importance of multiple biotic and anthropogenic stressors on four ecoregional polar bear populations, at two future decadal time periods, and based on two Intergovernmental Panel on Climate Change (IPCC) greenhouse gas emissions scenarios (Shared Socioeconomic Pathways [SSPs] 2.6 [low] and 8.5 [high]); and (2) identify achievable management actions that may enhance the prospects of long-term persistence. Normative model runs indicated that populations in all four ecoregions incurred increasing probabilities of being decreased or greatly decreased over time. The probabilities of polar bear populations being decreased or greatly decreased from mid- to end of the century ranged from ~55% to 87% for the SSP 2.6 emissions scenario, and 82% to 94% for the SSP 8.5 emissions scenario among ecoregions. Arctic sea ice conditions and marine prey availability had the greatest influences on future polar bear population outcomes and overrode any relative influence from all other stressors. Hunting mortality was the most influential individual anthropogenic stressor in the Archipelago and Seasonal Ice Ecoregions, whereas terrestrial refugia quality grouped with various anthropogenic activities or factors (e.g., resource extraction, oil spill) was most influential for the Polar Basin Divergent Ice and Polar Basin Convergent Ice ecoregions. Our findings indicate that near-term proactive management of multiple anthropogenic stressors could cumulatively reduce the decline in populations such that if future sea ice habitat loss is eventually curtailed, population abundance would be greater than it would have been otherwise. Additionally, our findings suggest that there is value in tailoring management actions to address ecoregion-specific threats, which may prove useful in informing the development of future circumpolar conservation plans.

Ecosphere↗

USGS2025

Variation in energetic balance among free-ranging polar bears during the spring mating and foraging season

Large carnivores are capable of consuming substantial biomasses that can significantly alter their body mass and condition over short periods. Here we examine the intra-seasonal variation of polar bear ( Ursus maritimus Phipps, 1774) body mass, energy intake, and condition in the spring from two subpopulations. We evaluate the biological and temporal factors that may have driven changes in body mass of 31 individuals captured and recaptured over 2–39 days and assess whether these changes influenced their estimated condition. Body mass changed by –61 to 33 kg ( = –2 kg) with bears exhibiting increases in mass with increasing age and decreases with greater initial mass. On average, estimated intake was 57 MJ/day. Estimated daily mass-specific body mass changes exhibited greater variation relative to previous measures in polar bears or brown bears ( U. arctos Linnaeus, 1758). Yet, across all bears, measures of condition remained similar between captures. The marked variation in mass gains or losses highlights the varying behavioral and physiological limitations that influence foraging success within this apex carnivore during a season when two key life history events converge wherein feeding is often reduced during mating activities despite the importance of the spring hyperphagia period to long-term energy balance.

Lancaster Sound, southern Beaufort Sea↗

USGS2025

Reply to the discussion by Pilfold “Polar bear mass change confirms hyperphagia follows ringed seal whelping”

The spring is a critical period when polar bears ( Ursus maritimus Phipps, 1774) are thought to have peak access to seals and acquire the majority of their annual energy requirements during a period of hyperphagia. Pagano et al. (Pagano A.M., Atkinson S.N., and Archer L.C. 2025. Arctic Science.11:1-14. doi:10.1139/as-2024-0051) examined the intra-seasonal changes in body mass of 31 polar bears on the spring sea ice and found polar bears exhibited a feast or famine lifestyle. A lack of a relationship between changes in body mass and recapture date suggested that many bears had not entered their primary period of hyperphagia. Pilfold extended our discussion to conclude that our data show polar bear hyperphagia begins after the period of ringed seal ( Pusa hispida Schreber, 1775) whelping, and discusses this in relation to previous work on the timing of polar bear seal kills. Here, we reassess whether our data provide information on the timing of polar bear hyperphagia. We find no relationships in our data to conclude when polar bear hyperphagia begins. Instead, our data highlight the conflicting pressures individuals face between the spring breeding season, when time spent foraging is often reduced to engage in mating behavior, and the spring hyperphagia period, when the bulk of annual energy requirements are met.

Arctic Science↗

USGS2024

A benchmark for computational analysis of animal behavior, using animal-borne tags

Background Animal-borne sensors (‘bio-loggers’) can record a suite of kinematic and environmental data, which are used to elucidate animal ecophysiology and improve conservation efforts. Machine learning techniques are used for interpreting the large amounts of data recorded by bio-loggers, but there exists no common framework for comparing the different machine learning techniques in this domain. This makes it difficult to, for example, identify patterns in what works well for machine learning-based analysis of bio-logger data. It also makes it difficult to evaluate the effectiveness of novel methods developed by the machine learning community. Methods To address this, we present the Bio-logger Ethogram Benchmark (BEBE), a collection of datasets with behavioral annotations, as well as a modeling task and evaluation metrics. BEBE is to date the largest, most taxonomically diverse, publicly available benchmark of this type, and includes 1654 h of data collected from 149 individuals across nine taxa. Using BEBE, we compare the performance of deep and classical machine learning methods for identifying animal behaviors based on bio-logger data. As an example usage of BEBE, we test an approach based on self-supervised learning. To apply this approach to animal behavior classification, we adapt a deep neural network pre-trained with 700,000 h of data collected from human wrist-worn accelerometers. Results We find that deep neural networks out-perform the classical machine learning methods we tested across all nine datasets in BEBE. We additionally find that the approach based on self-supervised learning out-performs the alternatives we tested, especially in settings when there is a low amount of training data available. Conclusions In light of these results, we are able to make concrete suggestions for designing studies that rely on machine learning to infer behavior from bio-logger data. Therefore, we expect that BEBE will be useful for making similar suggestions in the future, as additional hypotheses about machine learning techniques are tested. Datasets, models, and evaluation code are made publicly available at https://github.com/earthspecies/BEBE , to enable community use of BEBE.

Movement Ecology↗

USGS2024

Ursids evolved dietary diversity without major alterations in metabolic rates

The diets of the eight species of ursids range from carnivory (e.g., polar bears, Ursus maritimus ) to insectivory (e.g., sloth bears, Melursus ursinus ), omnivory (e.g., brown bears, U. arctos ), and herbivory (e.g., giant pandas, Ailuropoda melanoleuca ). Dietary energy availability ranges from the high-fat, highly digestible, calorically dense diet of polar bears (~ 6.4 kcal digestible energy/g fresh weight) to the high-fiber, poorly digestible, calorically restricted diet (~ 0.7) of giant pandas. Thus, ursids provide the opportunity to examine the extent to which dietary energy drives evolution of energy metabolism in a closely related group of animals. We measured the daily energy expenditure (DEE) of captive brown bears in a relatively large, zoo-type enclosure and compared those values to previously published results on captive brown bears, captive and free-ranging polar bears, and captive and free-ranging giant pandas. We found that all three species have similar mass-specific DEE when travel distances and energy intake are normalized even though their diets differ dramatically and phylogenetic lineages are separated by millions of years. For giant pandas, the ability to engage in low-cost stationary foraging relative to more wide-ranging bears likely provided the necessary energy savings to become bamboo specialists without greatly altering their metabolic rate.

Scientific Reports↗

USGS2024

Polar bear energetic and behavioral strategies on land with implications for surviving the ice-free period

Declining Arctic sea ice is increasing polar bear land use. Polar bears on land are thought to minimize activity to conserve energy. Here, we measure the daily energy expenditure (DEE), diet, behavior, movement, and body composition changes of 20 different polar bears on land over 19–23 days from August to September (2019–2022) in Manitoba, Canada. Polar bears on land exhibited a 5.2-fold range in DEE and 19-fold range in activity, from hibernation-like DEEs to levels approaching active bears on the sea ice, including three individuals that made energetically demanding swims totaling 54–175 km. Bears consumed berries, vegetation, birds, bones, antlers, seal, and beluga. Beyond compensating for elevated DEE, there was little benefit from terrestrial foraging toward prolonging the predicted time to starvation, as 19 of 20 bears lost mass (0.4–1.7 kg•day −1 ). Although polar bears on land exhibit remarkable behavioral plasticity, our findings reinforce the risk of starvation, particularly in subadults, with forecasted increases in the onshore period.

Manitoba↗

USGS2023

A body composition model with multiple storage compartments for polar bears (Ursus maritimus)

Climate warming is rapidly altering Arctic ecosystems. Polar bears ( Ursus maritimus ) need sea ice as a platform from which to hunt seals, but increased sea-ice loss is lengthening periods when bears are without access to primary hunting habitat. During periods of food scarcity, survival depends on the energy that a bear has stored in body reserves, termed storage energy, making this a key metric in predictive models assessing climate change impacts on polar bears. Here, we developed a body composition model for polar bears that estimates storage energy while accounting for changes in storage tissue composition. We used data of dissected polar bears ( n = 31) to link routinely collected field measures of total body mass and straight-line body length to the body composition of individual bears, described in terms of structural mass and two storage compartments, adipose and muscle. We then estimated the masses of metabolizable proteins and lipids within these storage compartments, giving total storage energy. We tested this multi-storage model by using it to predict changes in the lipid stores from an independent dataset of wild polar bears ( n = 36) that were recaptured 8–200 days later. Using length and mass measurements, our model successfully predicted direct measurements of lipid changes via isotopic dilutions (root mean squared error of 14.5 kg). Separating storage into two compartments, and allowing the molecular composition of storage to vary, provides new avenues for quantifying energy stores of individuals across their life cycle. The multi-storage body composition model thus provides a basis for further exploring energetic costs of physiological processes that contribute to individual survival and reproductive success. Given bioenergetic models are increasingly used as a tool to predict individual fitness and population dynamics, our approach for estimating individual energy stores could be applicable to a wide range of species.

Article↗

USGS2023

Incremental evolution of modeling a prognosis for polar bears in a rapidly changing Arctic

Updating predictions of the response of high-profile, at-risk species to climate change and anthropogenic stressors is vital for informing effective conservation action. Here, we review two prior generations of Bayesian network probability models predicting changes in global polar bear ( Ursus maritimus ) population status, and provide a contemporary update based on recent research findings and sea-ice projections by newer climate models. We compare predictions of polar bear population response from all 3 models among four circumpolar Arctic ecoregions, using sea ice projections based on three IPCC greenhouse gas emissions scenarios (SSP2.6, 4.5, 8.5). Consistent with the previous two model generations, polar bears will continue to experience increasing probability of declining or greatly declining populations throughout the 21st century, varying by emission scenario. Populations within the Polar Basin Divergent Ice Ecoregion have the highest predicted probability of declines, but predictions were slightly less dire relative to the previous model generation. Most of the influence, denoted by model sensitivity analysis, is from expected degradation and loss of sea ice and reduced access to marine prey. The lack of terrestrial prey adequate to substitute for loss of access to marine prey, as well as human-caused bear morality associated with hunting and defense of life and property encountered when polar bears are increasingly forced ashore also contributed to predicted declines. Although some tidewater glacial fjords and other localized onshore resources may provide local refugia, their benefit is transient. Our findings continue to inform priorities for inventory, monitoring, and research needs, and suggest that similar updates to models of other at-risk species can capitalize on the comparison framework we present here.

Ecological Indicators↗

USGS2023

Lactation performance in polar bears is associated with fasting time and energetic state

Females must continually make resource allocation decisions because of fitness trade-offs between self-maintenance and investment in current offspring, yet factors underpinning these decisions are unresolved. Polar bears Ursus maritimus face considerable allocation challenges when seasonal sea-ice melt precludes access to prey for several months, and females rely solely on energy stores to cover their own energetic needs and provision offspring. We tested how female polar bears regulate lactation during onshore fasting (i.e. capital breeding) and determined the consequences of moderated lactation for females and cubs. Overall, milk energy declined, and lactation was more likely to cease with longer time fasting. Lactation was partially mediated by maternal energetic state and depended on litter characteristics. Milk energy declined more sharply with fasting time (~2.6 times more strongly) in females with 2 offspring compared to those with 1. Females with cubs-of-the-year produced higher energy milk than those with yearlings, and their milk energy also increased more strongly with maternal energy density. Milk energy declines benefited females via reduced depletion of maternal energy reserves, but cub growth decreased. Altered lactation investment likely has consequences for both female survival and the fate of offspring, which could scale up to influence population dynamics. Given that Arctic warming means polar bears across much of their range will experience longer periods without access to primary prey, our results underscore how lactation will likely become increasingly compromised.

Marine Ecology Progress Series↗

USGS2022

Observed and forecasted changes in land use by polar bears in the Beaufort and Chukchi Seas, 1985–2040

Monitoring changes in the distribution of large carnivores is important for managing human safety and supporting conservation. Throughout much of their range, polar bears ( Ursus maritimus ) are increasingly using terrestrial habitats in response to Arctic sea ice decline. Their increased presence in coastal areas has implications for bear-human conflict, inter-species interactions, and polar bear health and survival. We examined observed trends in land use over three decades by polar bears in the southern Beaufort Sea (SB) and Chukchi Sea (CS) where bears have traditionally spent most of the year on the sea ice. Using data from 408 adult females fitted with satellite radio-collars, we examined trends in the annual proportion of bears coming onshore (hereafter referred to as “percent of bears”) during the summer for ≥21 days, arrival and departure dates, duration spent onshore and relationships with sea ice metrics. We then estimated future land use through 2040 by extrapolating trends and by combining observed relationships between land use and sea ice with projections of future sea ice from an ensemble of earth system models. The observed percent of bears summering onshore and their duration onshore was correlated with the percent of open water that occurred within their population’s range between July and October. As sea ice declined, the percent of bears summering onshore increased from ~5 to 30% in the SB and ~10 to 50% in the CS and duration onshore increased by >30 days to 60–70 days in both populations. Using a range of greenhouse gas emission scenarios and adjustments for faster than forecasted sea ice loss we estimated that 50-62% of SB and 79-88% of CS bears will spend 90–108 and 110–126 days onshore during summer in the SB and CS, respectively, by 2040. Sea ice projections varied little between greenhouse gas emission scenarios prior to 2040 but diverged thereafter. Observed and forecasted increases in polar bear land occupancy puts more bears in proximity to human activities and settlements for longer durations while extending the lack of access to their primary prey. Because human conflict is one of the primary factors affecting the conservation of large carnivores worldwide, mitigation of bear-human interactions on land will be an increasingly important component of polar bear conservation.

Alaska↗

USGS2021

Effects of sea ice decline and summer land use on polar bear home range size in the Beaufort Sea

Animals responding to habitat loss and fragmentation may increase their home ranges to offset declines in localized resources or they may decrease their home ranges and switch to alternative resources. In many regions of the Arctic, polar bears ( Ursus maritimus ) exhibit some of the largest home ranges of any quadrupedal mammal. Polar bears are presently experiencing a rapid decline in Arctic sea ice extent and a change in sea ice composition. For the Southern Beaufort Sea subpopulation of polar bears, this has resulted in a divergent movement pattern where most of the subpopulation remains on the sea ice in the summer melt season while the remainder move to land. We evaluated the effects of summer land use and maternal denning on the annual and seasonal utilization distribution size (i.e., home range) of adult female polar bears in the Southern Beaufort Sea subpopulation over 30 yr (1986–2016) during a period of rapid sea ice decline. For bears that remained on the summer sea ice, model-derived mean annual utilization distributions were 64% larger in 1999–2016 ( x ̄ "> 𝑥 ̄ = 176,000 km 2 ) relative to 1986–1998 ( x ̄ "> 𝑥 ̄ = 107,000 km 2 ). This increase was primarily driven by increases in summer utilization distributions that encompassed increased amounts of open water and decreased amounts of preferred sea ice. The mean centroid of summer utilization distributions for bears that remained on the sea ice was 193 km further north-northeast in 1999–2016. In contrast, bears that summered on land during 1999–2016 exhibited 88% smaller mean annual utilization distribution sizes ( x ̄ "> 𝑥 ̄ = 22,000 km 2 ) relative to bears that remained on the summer sea ice during the same period. Our findings highlight the impacts of sea ice declines on polar bear space use and the increasing importance of land as an alternative summer refuge.

Alaska↗

USGS2021

Polar bear foraging behavior

Polar bears forage in the marine environment, primarily on the sea ice over the shallow waters of the continental shelf. They are solitary, ambush hunters that catch ringed and bearded seals when they surface to breathe in ice holes or haul out on the ice to rest and molt. In most parts of their range, polar bears experience dramatic seasonal variability in their ability to catch seals, with foraging success peaking in late spring and early summer when seal pups are weaned. During this time, the body mass of polar bears can nearly double, especially in pregnant females, such that body composition may reach 49% body fat. The accumulation of body fat is vital for these bears to survive through the autumn and winter when seals are less accessible or when pregnant adult female bears enter dens and fast. When the sea ice retreats in summer, some bears exhibit a temporary switch to omnivory, feeding on a variety of terrestrial food. However, the energetic benefit of most terrestrial food is small relative to their marine mammal prey and, in some regions, increased land use has been associated with declines in body condition. Reduced accessibility of seal prey to polar bears as a result of global climate change threatens the long-term sustainability of this Arctic predator.

Book chapter↗

USGS2020

Identifying reliable indicators of fitness in polar bears

Animal structural body size and condition are often measured to evaluate individual health, identify responses to environmental change and food availability, and relate food availability to effects on reproduction and survival. A variety of condition metrics have been developed but relationships between these metrics and vital rates are rarely validated. Identifying an optimal approach to estimate the body condition of polar bears is needed to improve monitoring of their response to decline in sea ice habitat. Therefore, we examined relationships between several commonly used condition indices (CI), body mass, and size with female reproductive success and cub survival among polar bears ( Ursus maritimus ) measured in two subpopulations over three decades. To improve measurement and application of morphometrics and CIs, we also examined whether CIs are independent of age and structural size–an important assumption for monitoring temporal trends—and factors affecting measurement precision and accuracy. Maternal CIs and mass measured the fall prior to denning were related to cub production. Similarly, maternal CIs, mass, and length were related to the mass of cubs or yearlings that accompanied her. However, maternal body mass, but not CIs, measured in the spring was related to cub production and only maternal mass and length were related to the probability of cub survival. These results suggest that CIs may not be better indicators of fitness than body mass in part because CIs remove variation associated with body size that is important in affecting fitness. Further, CIs exhibited variable relationships with age for growing bears and were lower for longer bears despite body length being related to cub survival and female reproductive success. These results are consistent with findings from other species indicating that body mass is a useful metric to link environmental conditions and population dynamics.

PLoS ONE↗

USGS2020

The seasonal energetic landscape of an apex marine carnivore, the polar bear

Divergent movement strategies have enabled wildlife populations to adapt to environmental change. In recent decades, the Southern Beaufort Sea subpopulation of polar bears (Ursus maritimus) has developed a divergent movement strategy in response to diminishing sea ice where the majority of the subpopulation (73–85%) stays on the sea ice in summer and the remaining bears move to land. Although declines in sea ice are generally considered a challenge to energy balance in polar bears residing in some regions of the Arctic, little quantitative data exists concerning the seasonal energy expenditures of this apex marine carnivore. We used GPS satellite collars with tri-axial accelerometers and conductivity sensors to measure the location, behavior, and energy expenditure of five adult female polar bears in the southern Beaufort Sea across seasons of sea ice breakup and minimum extent. Using a Bayesian mixed-effects model, we found that energy expenditure was influenced by month, ocean depth, and habitat type (sea ice or land). Total energy expenditure from May to October ranged from 37.7 – 47.2 mJ kg-1 for individual bears. Bears that moved to land expended 7% more energy on average from May to October than bears that remained on the receding sea ice. In August, when bears were moving from the sea ice to land or moving north with the receding pack ice, bears that moved to land spent 7% more time swimming and expended 22% more energy. Meaning the immediate cost of moving to land exceeded the cost of remaining on the receding summer pack ice. These findings suggest a physiological reason why the majority of the Southern Beaufort Sea subpopulation continues to inhabit a diminishing summer ice platform. However, bears that moved to land spent 29% more time in preferred hunting habitats over the continental shelf than bears that remained on the sea ice. Bears on land also had access to subsistence-harvested bowhead whale carcasses. Hence, our findings indicate there may be a greater overall energetic benefit to move to land in this region, which suggests that the use of the diminishing summer sea ice may be functioning as an ecological trap.

Alaska↗

USGS2019

Overhauling ocean spatial planning to improve marine megafauna conservation

Tracking data have led to evidence-based conservation of marine megafauna, but a disconnect remains between the many thousands of individual animals that have been tracked and the use of these data in conservation and management actions. Furthermore, the focus of most conservation efforts is within Exclusive Economic Zones despite the ability of these species to move thousands of kilometres across multiple national jurisdictions. To assist the goal of the United Nations General Assembly’s recent effort to negotiate a global treaty to conserve biodiversity on the high seas, we propose the development of a new frontier in dynamic marine spatial management. We argue that a global approach combining tracked movements of marine megafauna and human activities at-sea, and using existing and emerging technologies (e.g., through new tracking devices and big data approaches) can be applied to deliver near real-time diagnostics on existing risks and threats to mitigate global risks for marine megafauna. With technology developments over the next decade expected to catalyse the potential to survey marine animals and human activities in ever more detail and at global scales, the development of dynamic predictive tools based on near real-time tracking and environmental data will become crucial to address increasing risks. Such global tools for dynamic spatial and temporal management will, however, require extensive synoptic data updates and will be dependent on a shift to a culture of data sharing and open access. We propose a global mechanism to store and make such data available in near real-time, enabling a holistic view of space use by marine megafauna and humans that would significantly accelerate efforts to mitigate impacts and improve conservation and management of marine megafauna.

Frontiers in Marine Science↗

USGS2019

Estimating the energy expenditure of free‐ranging polar bears using tri‐axial accelerometers: A validation with doubly labeled water

Measures of energy expenditure can be used to inform animal conservation and management, but methods for measuring the energy expenditure of free‐ranging animals have a variety of limitations. Advancements in biologging technologies have enabled the use of dynamic body acceleration derived from accelerometers as a proxy for energy expenditure. Although dynamic body acceleration has been shown to strongly correlate with oxygen consumption in captive animals, it has been validated in only a few studies on free‐ranging animals. Here, we use relationships between oxygen consumption and overall dynamic body acceleration in resting and walking polar bears Ursus maritimus and published values for the costs of swimming in polar bears to estimate the total energy expenditure of 6 free‐ranging polar bears that were primarily using the sea ice of the Beaufort Sea. Energetic models based on accelerometry were compared to models of energy expenditure on the same individuals derived from doubly labeled water methods. Accelerometer‐based estimates of energy expenditure on average predicted total energy expenditure to be 30% less than estimates derived from doubly labeled water. Nevertheless, accelerometer‐based measures of energy expenditure strongly correlated ( r 2 = 0.70) with measures derived from doubly labeled water. Our findings highlight the strengths and limitations in dynamic body acceleration as a measure of total energy expenditure while also further supporting its use as a proxy for instantaneous, detailed energy expenditure in free‐ranging animals.

Alaska, Northwest Territories, Yukon↗

USGS2019

Energetic costs of aquatic locomotion in a subadult polar bear

Most marine mammals rely on swimming as their primary form of locomotion. These animals have evolved specialized morphologies, physiologies, and behaviors that have enabled them to efficiently move through an aquatic environment (Williams 1999 ). Such adaptations include body streamlining, modified plantar surfaces for propulsion, and abilities to remain submerged for extended durations (Williams 1989 ). As a result of these adaptations, many marine mammal species exhibit minimal increases in metabolism at routine swim speeds relative to resting rates (Williams et al . 1992 , 2017 ; Yazdi et al . 1999 ). Contrary to most marine mammals, polar bears ( Ursus maritimus ) rely on walking as their primary form of locomotion. As a consequence, they exhibit little evidence of body streamlining or abilities to remain submerged for extended durations. The longest dive recorded for a polar bear is 3 min and 10 s (Stirling and van Meurs 2015 ), a relatively brief period compared to other marine mammals (Ponganis 2015 ). Nevertheless, polar bears do exhibit large forepaws (DeMaster and Stirling 1981 ), lower and flatter heads (Slater et al . 2010 ), and more dense forelimb bones (Wall 1983 ) relative to other bear species, potentially as adaptations for swimming. Polar bears further exhibit some distinct physiological and behavioral adaptations from other bear species likely as a consequence of their marine existence (Pagano et al . 2018 a ).

Marine Mammal Science↗