During 17 spring and summer field seasons between 1973 and 1999, we documented 220 bouts of nursing by dependent polar bear (Ursus maritimus Phipps, 1774) cubs at Radstock Bay, Nunavut, Canada. The overall mean duration of nursing bouts for cubs-of-the-year (COY) and yearlings (YRLG) litters was 7.1 min (standard deviation (SD) = 3.3, range = 1–23). Mean nursing bout durations of one- and two-cub litters of COY and YRLG in spring and summer seasons ranged from 6.09 to 7.78 min and from 5.00 to 9.18 min, respectively. The overall mean duration of inter-nursing intervals for COY and YRLG litters was 5.7 h (SD = 4.9, range = 0.0–35.0). The mean inter-nursing interval for one-cub litters was 6.4 h (SD = 4.6, range = 0.0–20.2) and for two-cub litters was 5.1 h (SD = 5.1, range = 0.0–35.0). We found no evidence for effects of season or cub age class on nursing behavior. We found weak evidence that two-cub litters nurse slightly longer than one-cub litters, potentially reflecting reduced nursing efficiency due to sibling rivalry. There was neither evidence for diel patterns in nursing behavior nor a detectable relationship between the cessation of nursing and the onset of hunting or sleeping by the adult female.

Conducting assessments to understand the effects of changing environmental conditions on polar bear (Ursus maritimus) demography has become increasingly important to inform management and conservation. Here, we combined physical (2005–2007) and genetic (2017–2018) mark-recapture with harvest recovery data (2005–2018) to estimate demographic rates of the Davis Strait polar bear subpopulation and examine the possible effects of climate, dynamic ice habitat, and prey resources on survival. Large sample sizes (e.g., 2,513 marked animals) allowed us to estimate temporal variation in annual survival rates using multistate mark-recapture-recovery models. We did not detect statistically significant effects of climate, ice habitat, and prey during the 13-year study. Estimated total abundance in 2006 was 2,190, credible interval (CRI) [1,954, 2,454] and 1,944, CRI [1,593, 2,366] in 2018. Geometric mean population growth rate (0.99, 95% CRI [0.97, 1.01]) indicated the subpopulation may have declined slightly between 2006 and 2018. However, we did not detect a declining trend in survival or substantial change in reproductive metrics over this period. Given forecasts of major environmental change we emphasize the need to review monitoring programs for this subpopulation.

Climate change is a long-term threat to polar bears. However, sea-ice loss is hypothesized to provide transient benefits in high latitudes, where thick multiyear ice historically limited biological productivity and seal abundance. We used joint live-recapture and dead-recovery mark-recapture models to analyze data for one of the most northerly polar bear subpopulations, Kane Basin. The data consisted of 277 initial live captures and genetic identifications (1992–1997 = 150, 2012–-2014 = 127), 89 recaptures or re-identifications (1992–1997 = 53, 2012–2014 = 36), and 24 harvest returns of research-marked bears during 1992–2014. We estimated mean annual abundance of 357 bears (95% CI: 221–493) for 2013–2014. This suggests a likely increase relative to our estimate of 224 (95% CI: 145–303) bears in the mid-1990s and relative to a previously published estimate of 164 (95% CI: 94–234) bears in the mid-1990s that used some of the same data. This is also supported by an apparent increase in the density of bears in eastern Kane Basin during 2012–2014. Estimates of total survival for females ≥3 yr old (mean ± SE: 0.95 ± 0.04) and their dependent offspring were similar to previous estimates from the 1990s, and estimates of unharvested survival for females ≥3 yr (0.96 ± 0.04) appear sufficient for positive population growth. Estimates of total survival were lower for males ≥3 yr (0.87 ± 0.06). We documented a reduction in mortality associated with subsistence harvest, likely attributable to implementation of a harvest quota by Greenland in 2006. Our findings, together with evidence for increased range sizes, improved body condition for all sex and age classes, and stable reproductive metrics, show that this small high-Arctic polar bear subpopulation remains productive and healthy. These benefits are likely temporary given predictions for continued climate change.

During 17 field seasons between 1973 and 1999, we conducted a long-term study of the behavior of undisturbed wild polar bears in Radstock Bay, southwest Devon Island, Nunavut. In a subset of 11 seasons (6 spring and 5 summer) between 1975 and 1997, we used three different drug combinations to chemically immobilize a small number of adult and subadult polar bears on an opportunistic basis and applied a temporary dye mark so that individual bears could be visually reidentified. We then used multinomial logistic regression to compare the behavior of 35 previously immobilized bears of five different demographic classes (sex, age, and reproductive status) to the behavior of non-immobilized bears of the same demographic classes in the same years and seasons. During the first two days after immobilization, bears slept significantly more and spent less time hunting than did bears that had not been immobilized. However, previously immobilized bears returned to the same behavioral patterns and proportion of total time spent hunting as non-immobilized bears within two days and no further negative behavioral effects were detected in the following 21 d. We visually confirmed successful hunting by three adult bears within 0.4 to 2.1 d of being immobilized, all of which went on to make additional kills within the following 24 h. The return to normal behavior patterns, including the ability to hunt successfully, within 48 h of immobilization appears consistent with the hypothesis that polar bears do not experience longer-term behavioral effects following brief chemical immobilization for conservation and management purposes.

Animal movement is a fundamental ecological process affecting the survival and reproduction of individuals, the structure of populations, and the dynamics of communities. Methods to quantify animal movement and spatiotemporal abundances, however, are generally separate and therefore omit linkages between individual-level and population-level processes. We describe an integrated spatial capture–recapture (SCR) movement model to jointly estimate (1) the number and distribution of individuals in a defined spatial region and (2) movement of those individuals through time. We applied our model to a study of polar bears (Ursus maritimus) in a 28,125 km² survey area of the eastern Chukchi Sea, USA in 2015 that incorporated capture–recapture and telemetry data. In simulation studies, the model provided unbiased estimates of movement, abundance, and detection parameters using a bivariate normal random walk and correlated random walk movement process. Our case study provided detailed evidence of directional movement persistence for both male and female bears, where individuals regularly traversed areas larger than the survey area during the 36-day study period. Scaling from individual- to population-level inferences, we found that densities varied from <0.75 bears/625 km² grid cell/day in nearshore cells to 1.6–2.5 bears/grid cell/day for cells surrounded by sea ice. Daily abundance estimates ranged from 53 to 69 bears, with no trend across days. The cumulative number of unique bears that used the survey area increased through time due to movements into and out of the area, resulting in an estimated 171 individuals using the survey area during the study (95% credible interval 124–250). Abundance estimates were similar to a previous multiyear integrated population model using capture–recapture and telemetry data (2008–2016; Regehr et al., Scientific Reports 8:16780, 2018). Overall, the SCR–movement model successfully quantified both individual- and population-level space use, including the effects of landscape characteristics on movement, abundance, and detection, while linking the movement and abundance processes to directly estimate density within a prescribed spatial region and temporal period. Integrated SCR–movement models provide a generalizable approach to incorporate greater movement realism into population dynamics and link movement to emergent properties including spatiotemporal densities and abundances.

Polar bears are susceptible to climate warming because of their dependence on sea ice, which is declining rapidly. We present the first evidence for a genetically distinct and functionally isolated group of polar bears in Southeast Greenland. These bears occupy sea-ice conditions resembling those projected for the High Arctic in the late 21st century, with an annual ice-free period that is >100 days longer than the estimated fasting threshold for the species. Whereas polar bears in most of the Arctic depend on annual sea ice to catch seals, Southeast Greenland bears have a year-round hunting platform in the form of freshwater glacial mélange. This suggests that marine-terminating glaciers, although of limited availability, may serve as previously unrecognized climate refugia. Conservation of Southeast Greenland polar bears, which meet criteria for recognition as the world’s 20th polar bear subpopulation, is necessary to preserve the genetic diversity and evolutionary potential of the species.

Climate change threatens global biodiversity. Many species vulnerable to climate change are important to humans for nutritional, cultural, and economic reasons. Polar bears Ursus maritimus are threatened by sea-ice loss and represent a subsistence resource for Indigenous people. We applied a novel population modeling-management framework that is based on species life history and accounts for habitat loss to evaluate subsistence harvest for the Chukchi Sea (CS) polar bear subpopulation. Harvest strategies followed a state-dependent approach under which new data were used to update the harvest on a predetermined management interval. We found that a harvest strategy with a starting total harvest rate of 2.7% (~85 bears/yr at current abundance), a 2:1 male-to-female ratio, and a 10-yr management interval would likely maintain subpopulation abundance above maximum net productivity level for the next 35 yr (approximately three polar bear generations), our primary criterion for sustainability. Plausible bounds on starting total harvest rate were 1.7–3.9%, where the range reflects uncertainty due to sampling variation, environmental variation, model selection, and differing levels of risk tolerance. The risk of undesired demographic outcomes (e.g., overharvest) was positively related to harvest rate, management interval, and projected declines in environmental carrying capacity; and negatively related to precision in population data. Results reflect several lines of evidence that the CS subpopulation has been productive in recent years, although it is uncertain how long this will last as sea-ice loss continues. Our methods provide a template for balancing trade-offs among protection, use, research investment, and other factors. Demographic risk assessment and state-dependent management will become increasingly important for harvested species, like polar bears, that exhibit spatiotemporal variation in their response to climate change.

There is an imminent need to collect information on distribution and abundance of polar bears (Ursus maritimus) to understand how they are affected by the ongoing decrease in Arctic sea ice. The Kane Basin (KB) subpopulation is a group of high-latitude polar bears that ranges between High Arctic Canada and NW Greenland around and north of the North Water polynya (NOW). We conducted a line transect distance sampling aerial survey of KB polar bears during 28 April–12 May 2014. A total of 4160 linear kilometers were flown in a helicopter over fast ice in the fjords and over offshore pack ice between 76° 50' and 80° N'. Using a mark-recapture distance sampling protocol, the estimated abundance was 190 bears (95% lognormal CI: 87–411; CV 39%). This estimate is likely negatively biased to an unknown degree because the offshore sectors of the NOW with much open water were not surveyed because of logistical and safety reasons. Our study demonstrated that aerial surveys may be a feasible method for obtaining abundance estimates for small subpopulations of polar bears.

Changes in sea-ice dynamics are affecting polar bears Ursus maritimus across their circumpolar range, which highlights the importance of periodic demographic assessments to inform management and conservation. We used genetic mark-recapture-recovery to derive estimates of abundance and survival for the Baffin Bay (BB) polar bear subpopulation — the first time this method has been used successfully for this species. Genetic data from tissue samples we collected via biopsy darting were combined with historical physical capture and harvest recovery data. The combined data set consisted of 1410 genetic samples (2011–2013), 914 physical captures (1993–1995, 1997), and 234 harvest returns of marked bears (1993–2013). The estimate of mean subpopulation abundance was 2826 (95% CI = 2284–3367) in 2012–2013. Estimates of annual survival (mean ñ SE) were 0.90 ñ 0.05 and 0.78 ñ 0.06 for females and males age >= 2 yr, respectively. The proportion of total mortality of adult females and males that was attributed to legal harvest was 0.16 ñ 0.05 and 0.26 ñ 0.06, respectively. Remote sensing sea-ice data, telemetry data, and spatial distribution of onshore sampling indicated that polar bears were more likely to use offshore sea-ice habitat during the 1990s sampling period compared to the 2010s. Furthermore, in the 1990s, sampling of deep fjords and inland areas was limited, and no offshore sampling occurred in either time period, which precluded comparisons of abundance between the 1993–1997 and 2011–2013 study periods. Our findings demonstrate that genetic sampling can be a practical method for demographic assessment of polar bears over large spatial and temporal scales.

Understanding the influence of individual attributes on demographic processes is a key objective of wildlife population studies. Capture–recapture and age data are commonly collected to investigate hypotheses about survival, reproduction, and viability. We present a novel age-structured Jolly–Seber model that incorporates age and capture–recapture data to provide comprehensive information on population dynamics, including abundance, age-dependent survival, recruitment, age structure, and population growth rates. We applied our model to a multi-year capture–recapture study of polar bears (Ursus maritimus) in western Hudson Bay, Canada (2012–2018), where management and conservation require a detailed understanding of how polar bears respond to climate change and other factors. In simulation studies, the age-structured Jolly–Seber model improved precision of survival, recruitment, and annual abundance estimates relative to standard Jolly–Seber models that omit age information. Furthermore, incorporating age information improved precision of population growth rates, increased power to detect trends in abundance, and allowed direct estimation of age-dependent survival and changes in annual age structure. Our case study provided detailed evidence for senescence in polar bear survival. Median survival estimates were lower (<0.95) for individuals aged <5 years, remained high (>0.95) for individuals aged 7–22 years, and subsequently declined to near zero for individuals >30 years. We also detected cascading effects of large recruitment classes on population age structure, which created major shifts in age structure when these classes entered the population and then again when they reached prime breeding ages (10–15 years old). Overall, age-structured Jolly–Seber models provide a flexible means to investigate ecological and evolutionary processes that shape populations (e.g., via senescence, life expectancy, and lifetime reproductive success) while improving our ability to investigate population dynamics and forecast population changes from capture–recapture data.

Arctic marine mammals are harvested by Indigenous people for subsistence and are socially and culturally important. For ice-dependent species like the polar bear Ursus maritimus, management and conservation require understanding interactions between harvest and sea-ice loss due to climate change. We developed a demographic model to evaluate harvest risk for polar bears in Southern Hudson Bay, Canada, where the annual ice-free season has increased by approximately one month in recent decades. The model was based on the theta-logistic equation and allowed for density-dependent changes (through carrying capacity [K]) and density-independent changes (through population growth rate [r]). Model parameters were estimated using a Bayesian Monte Carlo method that included capture-recapture, aerial survey, and harvest data. Harvest management followed a state-dependent approach under which new estimates of abundance were used to update the harvest level every five years. Under a middle-of-the-road environmental scenario that assumed K and r would decline in proportion to projected sea-ice declines, annual removal of 0.02–0.03 of females resulted in a 0.8 probability of maintaining subpopulation abundance above maximum net productivity level for three polar bear generations (~34 years), our primary criterion for sustainability. Under more pessimistic and optimistic environmental scenarios, comparable female harvest rates were 0.01 and 0.055, respectively. Our coupled modeling-management framework can be used to inform tradeoffs between protection and sustainable use for wildlife populations experiencing habitat loss.

Polar bears are of international conservation concern due to climate change but are difficult to study because of low densities and an expansive, circumpolar distribution. In a collaborative U.S.–Russian effort in spring of 2016, we used aerial surveys to detect and estimate the abundance of polar bears on sea ice in the Chukchi Sea. Our surveys used a combination of thermal imagery, digital photography, and human observations. Using spatio-temporal statistical models that related bear and track densities to physiographic and biological covariates (e.g., sea ice extent, resource selection functions derived from satellite tags), we predicted abundance and spatial distribution throughout our study area. Estimates of 2016 abundance ranged from 3,435 (95% CI: 2,300–5,131) to 5,444 (95% CI: 3,636–8,152) depending on the proportion of bears assumed to be missed on the transect line during Russian surveys. Our point estimates are larger than, but of similar magnitude to, a recent estimate for the period 2008–2016 (2,937; 95% CI 1,522–5,944) derived from an integrated population model applied to a slightly smaller area. Although a number of factors (e.g., equipment issues, differing platforms, low sample sizes, size of the study area relative to sampling effort) required us to make a number of assumptions to generate estimates, it establishes a useful lower bound for abundance, and suggests high spring polar bear densities on sea ice in Russian waters south of Wrangell Island. With future improvements, we suggest that springtime aerial surveys may represent a plausible avenue for studying abundance and distribution of polar bears and their prey over large, remote areas.

Kane Basin (KB) is one of the world's most northerly polar bear (Ursus maritimus) subpopulations, where bears have historically inhabited a mix of thick multiyear and annual sea ice year‐round. Currently, KB is transitioning to a seasonally ice‐free region because of climate change. This ecological shift has been hypothesized to benefit polar bears in the near‐term due to thinner ice with increased biological production, although this has not been demonstrated empirically. We assess sea‐ice changes in KB together with changes in polar bear movements, seasonal ranges, body condition, and reproductive metrics obtained from capture–recapture (physical and genetic) and satellite telemetry studies during two study periods (1993–1997 and 2012–2016). The annual cycle of sea‐ice habitat in KB shifted from a year‐round ice platform (~50% coverage in summer) in the 1990s to nearly complete melt‐out in summer (<5% coverage) in the 2010s. The mean duration between sea‐ice retreat and advance increased from 109 to 160 days (p = .004). Between the 1990s and 2010s, adult female (AF) seasonal ranges more than doubled in spring and summer and were significantly larger in all months. Body condition scores improved for all ages and both sexes. Mean litter sizes of cubs‐of‐the‐year (C0s) and yearlings (C1s), and the number of C1s per AF, did not change between decades. The date of spring sea‐ice retreat in the previous year was positively correlated with C1 litter size, suggesting smaller litters following years with earlier sea‐ice breakup. Our study provides evidence for range expansion, improved body condition, and stable reproductive performance in the KB polar bear subpopulation. These changes, together with a likely increasing subpopulation abundance, may reflect the shift from thick, multiyear ice to thinner, seasonal ice with higher biological productivity. The duration of these benefits is unknown because, under unmitigated climate change, continued sea‐ice loss is expected to eventually have negative demographic and ecological effects on all polar bears.

Climate change has broad ecological implications for species that rely on sensitive habitats. For some top predators, loss of habitat is expected to lead to cascading behavioral, nutritional, and reproductive changes that ultimately accelerate population declines. In the case of the polar bear (Ursus maritimus), declining Arctic sea ice reduces access to prey and lengthens seasonal fasting periods. We used a novel combination of physical capture, biopsy darting, and visual aerial observation data to project reproductive performance for polar bears by linking sea ice loss to changes in habitat use, body condition (i.e., fatness), and cub production. Satellite telemetry data from 43 (1991–1997) and 38 (2009–2015) adult female polar bears in the Baffin Bay subpopulation showed that bears now spend an additional 30 d on land (90 d in total) in the 2000s compared to the 1990s, a change closely correlated with changes in spring sea ice breakup and fall sea ice formation. Body condition declined for all sex, age, and reproductive classes and was positively correlated with sea ice availability in the current and previous year. Furthermore, cub litter size was positively correlated with maternal condition and spring breakup date (i.e., later breakup leading to larger litters), and negatively correlated with the duration of the ice‐free period (i.e., longer ice‐free periods leading to smaller litters). Based on these relationships, we projected reproductive performance three polar bear generations into the future (approximately 35 yr). Results indicate that two‐cub litters, previously the norm, could largely disappear from Baffin Bay as sea ice loss continues. Our findings demonstrate how concurrent analysis of multiple data types collected over long periods from polar bears can provide a mechanistic understanding of the ecological implications of climate change. This information is needed for long‐term conservation planning, which includes quantitative harvest risk assessments that incorporate estimated or assumed trends in future environmental carrying capacity.

Large carnivores are imperiled globally, and characteristics making them vulnerable to extinction (e.g., low densities and expansive ranges) also make it difficult to estimate demographic parameters needed for management. Here we develop an integrated population model to analyze capture-recapture, radiotelemetry, and count data for the Chukchi Sea subpopulation of polar bears (Ursus maritimus), 2008–2016. Our model addressed several challenges in capture-recapture studies for polar bears by including a multievent structure reflecting location and life history states, while accommodating state uncertainty. Female breeding probability was 0.83 (95% credible interval [CRI] = 0.71–0.90), with litter sizes of 2.18 (95% CRI = 1.71–2.82) for age-zero and 1.61 (95% CRI = 1.46–1.80) for age-one cubs. Total adult survival was 0.90 (95% CRI = 0.86–0.92) for females and 0.89 (95% CRI = 0.83–0.93) for males. Spring on-ice densities west of Alaska were 0.0030 bears/km² (95% CRI = 0.0016–0.0060), similar to 1980s-era density estimates although methodological differences complicate comparison. Abundance of the Chukchi Sea subpopulation, derived by extrapolating density from the study area using a spatially-explicit habitat metric, was 2,937 bears (95% CRI = 1,552–5,944). Our findings are consistent with other lines of evidence suggesting the Chukchi Sea subpopulation has been productive in recent years, although it is uncertain how long this will continue given sea-ice loss due to climate change.

Climate change is expected to result in range shifts and habitat fragmentation for many species. In the Arctic, loss of sea ice will reduce barriers to dispersal or eliminate movement corridors, resulting in increased connectivity or geographic isolation with sweeping implications for conservation. We used satellite telemetry, data from individually marked animals (research and harvest), and microsatellite genetic data to examine changes in geographic range, emigration, and interpopulation connectivity of the Baffin Bay (BB) polar bear (Ursus maritimus) subpopulation over a 25‐year period of sea‐ice loss. Satellite telemetry collected from n = 43 (1991–1995) and 38 (2009–2015) adult females revealed a significant contraction in subpopulation range size (95% bivariate normal kernel range) in most months and seasons, with the most marked reduction being a 70% decline in summer from 716,000 km² (SE 58,000) to 211,000 km² (SE 23,000) (p < .001). Between the 1990s and 2000s, there was a significant shift northward during the on‐ice seasons (2.6° shift in winter median latitude, 1.1° shift in spring median latitude) and a significant range contraction in the ice‐free summers. Bears in the 2000s were less likely to leave BB, with significant reductions in the numbers of bears moving into Davis Strait (DS) in winter and Lancaster Sound (LS) in summer. Harvest recoveries suggested both short and long‐term fidelity to BB remained high over both periods (83–99% of marked bears remained in BB). Genetic analyses using eight polymorphic microsatellites confirmed a previously documented differentiation between BB, DS, and LS; yet weakly differentiated BB from Kane Basin (KB) for the first time. Our results provide the first multiple lines of evidence for an increasingly geographically and functionally isolated subpopulation of polar bears in the context of long‐term sea‐ice loss. This may be indicative of future patterns for other polar bear subpopulations under climate change.

Climate change is altering habitat for many species, leading to shifts in distributions that can increase levels of human–wildlife conflict. To develop effective strategies for minimizing human–wildlife conflict, we must understand the relative influences that climate change and other factors have on wildlife distributions. Polar bears (Ursus maritimus) are increasingly using land during summer and autumn due to sea ice loss, leading to higher incidents of conflict and concerns for human safety. We sought to understand the relative influence of sea ice conditions, onshore habitat characteristics, and human-provisioned food attractants on the distribution and abundance of polar bears while on shore. We also wanted to determine how mitigation measures might reduce human–polar bear conflict associated with an anthropogenic food source. We built a Bayesian hierarchical model based on 14 years of aerial survey data to estimate the weekly number and distribution of polar bears on the coast of northern Alaska in autumn. We then used the model to predict how effective two management options for handling subsistence-harvested whale remains in the community of Kaktovik, Alaska might be. The distribution of bears on shore was most strongly influenced by the presence of whale carcasses and to a lesser extent sea ice and onshore habitat conditions. The numbers of bears on shore were related to sea ice conditions. The two management strategies for handling the whale carcasses reduced the estimated number of bears near Kaktovik by > 75%. By considering multiple factors associated with the onshore distribution and abundance of polar bears we discerned what role human activities played in where bears occur and how successful efforts to manage the whale carcasses might be for reducing human–polar bear conflict.

Satellite telemetry studies provide information that is critical to the conservation and management of species affected by ecological change. Here we report on the performance and retention of two types (SPOT-227 and SPOT-305A) of ear-mounted Argos-linked satellite transmitters (i.e., platform transmitter terminal, or PTT) deployed on free-ranging polar bears in Eastern Greenland, Baffin Bay, Kane Basin, the southern Beaufort Sea, and the Chukchi Sea during 2007–2013.

Transmissions from 142 out of 145 PTTs deployed on polar bears were received for an average of 69.3 days. The average functional longevity, defined as the number of days they transmitted while still attached to polar bears, for SPOT-227 was 56.8 days and for SPOT-305A was 48.6 days. Thirty-four of the 142 (24%) PTTs showed signs of being detached before they stopped transmitting, indicating that tag loss was an important aspect of tag failure. Furthermore, 10 of 26 (38%) bears that were re-observed following application of a PTT had a split ear pinna, suggesting that some transmitters were detached by force. All six PTTs that were still on bears upon recapture had lost the antenna, which indicates that antenna breakage was a significant contributor to PTT failure. Finally, only nine of the 142 (6%) PTTs—three of which were still attached to bears—had a final voltage reading close to the value indicating battery exhaustion. This suggests that battery exhaustion was not a major factor in tag performance.

The average functional longevity of approximately 2 months for ear-mounted PTTs (this study) is poor compared to PTT collars fitted to adult female polar bears, which can last for several years. Early failure of the ear-mounted PTTs appeared to be caused primarily by detachment from the ear or antenna breakage. We suggest that much smaller and lighter ear-mounted transmitters are necessary to reduce the risk of tissue irritation, tissue damage, and tag detachment, and with a more robust antenna design. Our results are applicable to other tag types (e.g., iridium and VHF systems) and to research on other large mammals that cannot wear radio collars.