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Abstract A fundamental assumption of many ecological studies is that researchers are studying a representative sample of the population of interest. However, fish and wildlife studies, including many telemetry studies, often rely upon data from marked individuals that are sampled opportunistically rather than randomly and may provide little information on unmarked individuals. Data that are unrepresentative at the time of marking may need to be censored until marked animal distributions are representative of the population. In the absence of additional data on unmarked individuals, evaluating the representativeness of newly marked individuals within a population of interest can be difficult. If previously marked individuals can be assumed to be spatially representative of the population, comparing the spatial distributions of different cohorts of marked individuals can provide an alternative means to assess the representativeness of newly marked animals. We used simulations to assess the effectiveness of a randomization test that compared the spatial distribution between newly marked and previously marked individuals as an alternative to recapture‐based methods for evaluating mixing of newly marked individuals with the population of interest and compared different metrics of spatial overlap. We then tested this randomization test on radio‐collared caribou (Rangifer tarandus) from 2 different herds with very different marking strategies. We found that, based on simulation results, kernel‐based metrics outperformed cluster analysis metrics and that Bhattacharyya's Affinity (BA) index of kernel overlap was a good metric to identify differences in spatial distribution between marked cohorts. The time it took for newly marked caribou to become mixed with previously marked animals was primarily related to the timing of marking in relation to seasonal movement patterns (periods of aggregation and dispersion, and seasonal differences in spatial fidelity) and the spatial distribution of marking relative to seasonal distribution. In some cases, mixing did not occur until calving aggregations formed in early June. We recommend that researchers evaluate the assumption that marked individuals are spatially representative of their study population and carefully assess sampling time and location of their marking program to increase the likelihood that marked individuals represent the population of interest.

Abstract Warming temperatures and advancing spring are affecting annual snow and ice cycles, as well as plant phenology, across the Arctic and boreal regions. These changes may be linked to observed population declines in wildlife, including barren‐ground caribou (Rangifer tarandus), a key species of Arctic environments. We quantified how barren‐ground caribou, characteristically both gregarious and migratory, synchronize births in time and aggregate births in space and investigated how these tactics are influenced by variable weather conditions. We analyzed movement patterns to infer calving dates for 747 collared female caribou from seven herds across northern North America, totaling 1255 calving events over a 15‐year period. By relating these events to local weather conditions during the 1‐year period preceding calving, we examined how weather influenced calving timing and the ability of caribou to reach their central calving area. We documented continental‐scale synchrony in calving, but synchrony was greatest within an individual herd for a given year. Weather conditions before and during gestation had contrasting effects on the timing and location of calving. Notably, a combination of unfavorable weather conditions during winter and spring, including the pre‐calving migration, resulted in a late arrival on the calving area or a failure to reach the greater calving area in time for calving. Though local weather conditions influenced calving timing differently among herds, warm temperatures and low wind speed, which are associated with soft, deep snow, during the spring and pre‐calving migration, generally affected the ability of female caribou to reach central calving areas in time to give birth. Delayed calving may have potential indirect consequences, including reduced calf survival. Overall, we detected considerable variability across years and across herds, but no significant trend for earlier calving by caribou, even as broad indicators of spring and snow phenology trend earlier. Our results emphasize the importance of monitoring the timing and location of calving, and to examine how weather during summer and winter are affecting calving and subsequent reproductive success.

Abstract Background Migrations in temperate systems typically have two migratory phases, spring and autumn, and many migratory ungulates track the pulse of spring vegetation growth during a synchronized spring migration. In contrast, autumn migrations are generally less synchronous and the cues driving them remain understudied. Our goal was to identify the cues that migrants use in deciding when to initiate migration and how this is updated while en route. Methods We analyzed autumn migrations of Arctic barren-ground caribou (Rangifer tarandus) as a series of persistent and directional movements and assessed the influence of a suite of environmental factors. We fitted a dynamic-parameter movement model at the individual-level and estimated annual population-level parameters for weather covariates on 389 individual-seasons across 9 years. Results Our results revealed strong, consistent effects of decreasing temperature and increasing snow depth on migratory movements, indicating that caribou continuously update their migratory decision based on dynamic environmental conditions. This suggests that individuals pace migration along gradients of these environmental variables. Whereas temperature and snow appeared to be the most consistent cues for migration, we also found interannual variability in the effect of wind, NDVI, and barometric pressure. The dispersed distribution of individuals in autumn resulted in diverse environmental conditions experienced by individual caribou and thus pronounced variability in migratory patterns. Conclusions By analyzing autumn migration as a continuous process across the entire migration period, we found that caribou migration was largely related to temperature and snow conditions experienced throughout the journey. This mechanism of pacing autumn migration based on indicators of the approaching winter is analogous to the more widely researched mechanism of spring migration, when many migrants pace migration with a resource wave. Such a similarity in mechanisms highlights the different environmental stimuli to which migrants have adapted their movements throughout their annual cycle. These insights have implications for how long-distance migratory patterns may change as the Arctic climate continues to warm.

Abstract How animals use their range can have physiological, ecological, and demographic repercussions, as well as impact management decisions, species conservation, and human society. Fidelity, the predictable return to certain places, can improve fitness if it is associated with high‐quality habitat or helps enable individuals to locate heterogenous patches of higher‐quality habitat within a lower‐quality habitat matrix. Our goal was to quantify patterns of fidelity at different spatial scales to better understand the relative plasticity of habitat use of a vital subsistence species that undergoes long‐distance migrations. We analyzed a decade (2010–2019) of GPS data from 240 adult, female Western Arctic Herd (WAH) caribou (Rangifer tarandus) from northwest Alaska, U.S.A. We assessed fidelity at 2 spatial scales: to site‐specific locations within seasonal ranges and to regions within the herd's entire range by using 2 different null datasets. We assessed both area and consistency of use during 6 different seasons of the year. We also assessed the temporal consistency of migration and calving events. At the scale of the overall range, we found that caribou fidelity was greatest during the calving and insect relief (early summer) seasons, where the herd tended to maximally aggregate in the smallest area, and lowest in winter when the seasonal range is largest. However, even in seasons with lower fidelity, we found that caribou still showed fidelity to certain regions within the herd's range. Within those seasonal ranges, however, there was little individual site‐specific fidelity from year to year, with the exception of summer periods. Temporally, we found that over 90% of caribou gave birth within 7 days of the day they gave birth the previous year. This revealed fairly high temporal consistency, especially given the spatial and temporal variability of spring migration. Fall migration exhibited greater temporal variability than spring migration. Our results support the hypothesis that higher fidelity to seasonal ranges is related to greater environmental and resource predictability. Interestingly, this fidelity was stronger at larger scales and at the population level. Almost the entire herd would seek out these areas with predictable resources, and then, individuals would vary their use, likely in response to annually varying conditions. During seasons with lower presumed spatial and/or temporal predictability of resources, population‐level fidelity was lower but individual fidelity was higher. The herd would be more spread out during the seasons of low‐resource predictability, leading to lower fidelity at the scale of their entire range, but individuals could be closer to locations they used the previous year, leading to greater individual fidelity, perhaps resulting from memory of a successful outcome the previous year. Our results also suggest that fidelity in 1 season is related to fidelity in the subsequent season. We hypothesize that some differences in patterns of range fidelity may be driven by seasonal differences in group size, degree of sociality, and/or density‐dependent factors. Climate change may affect resource predictability and, thus, the spatial fidelity and temporal consistency of use of animals to certain seasonal ranges.

Abstract Wildlife management often involves trade‐offs between protecting species and allowing human activities and development. Ideally, these decisions are guided by scientific studies that quantify the impacts of proposed actions on the environment. However, critical information to assess impacts of proposed activities may be lacking, such as certainty in where actions will take place, which may hinder a robust impact assessment. To address this issue, we present the Development Impacts Analysis (DIA), which employs Monte Carlo simulation modeling to quantify the environmental consequences of proposed development scenarios, while accounting for uncertainty in the exact location of future development. We applied the DIA to five proposed oil leasing management scenarios under a revised management plan for the National Petroleum Reserve—Alaska. For each management scenario with differing levels of proposed development (“alternatives”), oil production pads and roads were randomly simulated in proportion to estimated undiscovered oil and following alternative‐specific restrictions. We assessed habitat displacement for two caribou (Rangifer tarandus) herds, eight shorebird species, and black brant (Branta bernicla) based on reported responses to development, repeating the process 100 times for each alternative. Some habitat loss was reported for each proposed alternative, but the amount of impact varied by alternative and species. One caribou herd and most bird species indicated greatest effects in the alternative with the least restrictions on development and lesser impacts under more protective alternatives. Our results emphasized the importance of considering spatial variation in development effects and species‐specific differences when evaluating management proposals. The DIA quantified potential impacts on a suite of species under proposed management alternatives, while accounting for uncertainty in where development will occur and providing confidence intervals on estimated impacts. This illustrates that uncertainty need not preclude management decisions about establishment of broad land use restrictions prior to submission of project‐level proposals but can instead be explicitly incorporated into decision making. While no single management approach will likely benefit all species, use of tools such as the DIA allows managers to quantify trade‐offs among species and pursue approaches that balance the needs of various taxa and other management objectives.

A distinguishing characteristic of many migratory animals is their annual return to distinct calving (birthing) areas in the spring, yet the navigational mechanisms employed during migration that result in this pattern are poorly understood. Effective conservation of these species requires reliable delineation of such areas, quantifying the factors that influence their selection, and understanding the underlying mechanisms resulting in use of calving areas. We used barren-ground caribou (Rangifer tarandus granti) as a study species and identified calving sites of the Western Arctic Herd in Alaska using GPS collar data from 2010–2017. We assessed variability in calving areas by comparing spatial delineations across all combinations of years. To understand calving area selection at a landscape scale, we performed a resource selection analysis comparing calving sites to available locations across the herd’s range and incorporated time-varying, remotely sensed metrics of vegetation quality and quantity. We found that whereas calving areas varied from year to year, this annual variation was centered on an area of recurring attraction consistent with previous studies covering the last six decades. Calving sites were characterized by high-quality forage at the average time of calving, but not peak calving that year, and by a narrow range of distinct physiographic factors. Each year, calving sites were located on areas of above-average conditions based on our predictive model. Our findings indicate that the pattern of spring migration for pregnant females was to migrate to areas that consistently provide high-quality forage when averaged across years, and then upon arriving at this calving ground, refine selection using their perception of annually varying conditions that are driven by environmental stochasticity. We suggest that the well-documented and widespread pattern of fidelity to calving grounds by caribou is supportive of a navigational mechanism based on spatial memory at a broad scale to optimize foraging and energy acquisition at a critical life-history stage. The extent to which migrants depend on memory to reach their spring destinations has implications for the adaptability of populations to changing climate and human impacts.

Abstract Body size reflects realized physiological niche width at the population level and provides insights into the potential resiliency of a species to natural and anthropogenic perturbations to ecosystems. We analyzed patterns of American black and brown bear body size through a meta‐analysis of data from 18 studies conducted across North America, to evaluate the effects of species, sex, latitude, sympatry, and harvest. We used a bootstrap model selection procedure, which accounted for differences in sample size and population variation between studies, to investigate patterns in body mass. As expected, we found that brown bears were generally larger than black bears and both species were sexually dimorphic (i.e., males were generally larger than females). Black bear body size was not related to latitude, whereas brown bear body size was greatest at intermediate latitudes, possibly due to the presence of salmon. Neither sympatry nor harvest was associated with body size for either species at the geographic scale of our meta‐analyses, but both may warrant consideration at the local scale. Body size, as an index of population health, reflects phenotypic plasticity within species and populations and may serve as a useful indicator of niche utilization. Indeed, understanding large‐scale physiological patterns such as these can assist in understanding past, present, and future changes to realized niches and subsequent resiliency of species and populations.

Abstract Selection and use of dens is a critical behavioral adaption of bears (Ursus spp.) to survive extended periods of resource limitation. We analyzed selection of den sites by brown bears (Ursus arctos) in an arctic montane environment from 2014 to 2017. We used GPS data from 51 instrumented bears to physically locate, identify, and describe den sites (n = 33) and to quantitatively model the selection of denning habitat (n = 65 dens over 3 yr). We conducted an analysis of den‐site selection at the home range (third order) scale using variables describing topographic, vegetative, and spatially‐explicit fish and wildlife models. We hypothesized four main factors influence den‐site selection by brown bears: (1) physiography, (2) proximity to food resources, (3) thermal insulation, and (4) den‐area fidelity. We used kernel density estimates to define space use and conditional logistic regression to estimate den selection within individual home ranges. Brown bears selected den sites on steep slopes (>31°) with high snow load potential near their previously used den site. Our results supported our hypothesis that den selection by brown bears was related to thermal insulation, structural stability, drainage, and den‐area fidelity.

Animals that deliver a toxic secretion through a wound or to the body surface without a wound are considered venomous and toxungenous, respectively. Hematophagous insects, such as mosquitoes (Aedes spp.), meet the criteria for venomous, and some endoparasitic insects, such as warble flies (Hypoderma tarandi), satisfy the definition for toxungenous. The impacts of these insects on their hosts are wide ranging. In the Arctic, their primary host is the most abundant ungulate, the caribou (Rangifer tarandus). The most conspicuous impacts of these insects on caribou are behavioral. Caribou increase their movements during peak insect harassment, evading and running away from these parasites. These behavioral responses scale up to physiological effects as caribou move to less productive habitats to reduce harassment which increases energetic costs due to locomotion, reduces nutrient intake due to less time spent foraging, and can lead to poorer physiological condition. Reduced physiological condition can lead to lower reproductive output and even higher mortality rates, with the potential to ultimately affect caribou demographics. Caribou affect all trophic levels in the Arctic and the processes that connect them, thus altering caribou demographics could impact the ecology of the region. Broadening the definitions of venomous and toxungenous animals to include hematophagous and endoparasitic insects should not only generate productive collaborations among toxinologists and parasitologists, but will also lead to a deeper understanding of the ecology of toxic secretions and their widespread influence.

Lichens are the primary winter forage for large herds of migratory caribou (Rangifer tarandus). Caribou select for lichens more than they are available across the landscape and they generally avoid, during winter, habitat that has been burned by wildfires for decades while lichen abundance recovers. However, the relative importance of lichens in the diet is subject to debate. From 2010-2013, we conducted one of the largest microhistological studies of the early fall (58 samples from 1 site) and late winter (338 samples from 58 sites) diets of barren-ground caribou. Lichens con­stituted ~ 71% of the late winter diets of caribou in northwest Alaska, whereas moss (11%) and shrubs (9%) were the next most common forage items. Early fall diets were very similar to late winter, perhaps because deciduous vegetation is senescent during both periods. Diets of males, non-pregnant females and pregnant females were not significantly different. Pregnancy was not associated with the abundance of any forage type during winter but was associated with higher physiological stress. This result was expected as fall body condition dictates conception, caribou are ‘capital’ breeders, and gestation can be energetically demanding. Caribou that migrated south (i.e., wintered south of 67.1°N) had lower levels of nutritional stress, higher levels of lichen in the diet, and lower levels of moss and shrubs compared to caribou that did not migrate south. Future investigations into the potential connection between lichen abundance in the winter diet and survivorship, as well as linking the late summer diets of individuals to their reproductive success, should be undertaken.