Tall shrubs and trees are advancing into many tundra and wetland ecosystems but at a rate that often falls short of that predicted due to climate change. For forest, tall shrub, and tundra ecosystems in two pristine mountain ranges of Alaska, we apply a Bayesian, error-propagated calculation of expected elevational rise (climate velocity), observed rise (biotic velocity), and their difference (biotic inertia). We show a sensitive dependence of climate velocity on lapse rate and derive biotic velocity as a rigid elevational shift. Ecosystem presence identified from recent and historic orthophotos ~50 years apart was regressed on elevation. Biotic velocity was estimated as the difference between critical point elevations of recent and historic logistic fits divided by time between imagery. For both mountain ranges, the 95% highest posterior density of climate velocity enclosed the posterior distributions of all biotic velocities. In the Kenai Mountains, mean tall shrub and climate velocities were both 2.8 m y(-1). In the better sampled Chugach Mountains, mean tundra retreat was 1.2 m y(-1) and climate velocity 1.3 m y(-1). In each mountain range, the posterior mode of tall woody vegetation velocity (the complement of tundra) matched climate velocity better than either forest or tall shrub alone, suggesting competitive compensation can be important. Forest velocity was consistently low at 0.1-1.1 m y(-1), indicating treeline is advancing slowly. We hypothesize that the high biotic inertia of forest ecosystems in south-central Alaska may be due to competition with tall shrubs and/or more complex climate controls on the elevational limits of trees than tall shrubs. Among tall shrubs, those that disperse farthest had lowest inertia. Finally, the rapid upward advance of woody vegetation may be contributing to regional declines in Dall's sheep (Ovis dalli), a poorly dispersing alpine specialist herbivore with substantial biotic inertia due to dispersal reluctance.

Increasing demands for energy have generated interest in expanding oil and gas production on the North Slope of Alaska, USA, raising questions about the resilience of barren‐ground caribou (Rangifer tarandus) populations to new development. Although the amount of habitat lost directly to energy development in the Arctic will likely be relatively small, there are significant concerns about habitat that may be indirectly affected because of caribou avoidance behaviors. Behavioral responses to energy development for wildlife have been documented, but such responses are often assumed to dissipate over time, despite scant information on the ability of animals to habituate. To understand the long‐term effects of energy development on barren‐ground caribou, we investigated the behavior of the Central Arctic Herd in northern Alaska, which has been exposed to oil development on its summer range for approximately 40 years. Using recent (2015–2017) location data from global positioning system (GPS)‐collared females, we conducted a zone of influence analysis to assess whether caribou reduced their use of habitat near energy development, and if so, the distance the effects attenuated. We conducted this analysis for the calving, post‐calving, and mosquito harassment periods when caribou exhibit distinct resource selection patterns, and contrasted our results to past research that investigated the responses of the Central Arctic Herd immediately following the construction of the oil fields. Despite the long‐term presence of energy development within the Central Arctic Herd summer range, we found that female caribou exhibited avoidance responses to infrastructure during all time periods, although the effects waned across the summer. Caribou reduced their use of habitat within 5 km of development during the calving period, within 2 km during the post‐calving period, and within 1 km during the mosquito harassment period; these areas were predicted to overlap 12%, 15%, and 17% of important calving, post‐calving, and mosquito period habitat, respectively. During the calving period, the indirect effects we observed were similar to those observed in past research, whereas during the post‐calving and mosquito periods, we detected avoidance responses that had not been previously reported. These findings corroborate a growing body of evidence suggesting that habituation to industrial development in caribou in the Arctic is likely to be weak or absent, and emphasizes the value of minimizing the footprint of infrastructure within important seasonal habitat to reduce behavioral effects to barren‐ground caribou. © 2019 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.

The satellite-derived Normalized Difference Vegetation Index (NDVI) is commonly used by researchers and managers to represent ungulate forage conditions in landscapes across the globe, despite limited information about how it compares to empirical measurements of forage quality and quantity. The application of NDVI as a forage metric is particularly appealing for studying migratory caribou (Rangifer tarandus) in remote Arctic ecosystems, where field assessments are logistically and financially prohibitive, and climate-mediated changes in vegetation have been hypothesized to influence population declines. To determine the utility of NDVI for adequately representing caribou forage conditions, we compared NDVI derived from Moderate Resolution Imaging Spectroradiometer (MODIS) satellite imagery to empirical measures of caribou forage biomass, nitrogen, digestible nitrogen, and digestible energy within the summer range of the Central Arctic Caribou Herd on the North Slope of Alaska. Specifically, we determined the strength of forage-NDVI relationships at the start of the growing season and across the summer, assessed the efficacy of NDVI variables for modeling spatiotemporal variation in field measurements of different forage components, and used long-term MODIS data to estimate temporal changes in forage between 2000 and 2016. We found that NDVI values were weakly correlated with caribou forage quality at the start of the growing season and throughout the summer. Although linear models of forage-NDVI relationships performed poorly, NDVI variables (NDVI and the number of days from when NDVI reached its maximum value) were useful for modeling spatiotemporal variation in empirical measurements of forage components across the growing season, but only when we incorporated nonlinear forage-NDVI relationships and other habitat covariates. Phenological advances in the date of peak NDVI were associated with significant changes in forage conditions, particularly nitrogen, which exhibited earlier seasonal declines. Using long-term MODIS data, predicted values of forage nitrogen declined between 2000 and 2016, driven by exceedingly low values in 2014 and 2015. Given our results, we caution the application of NDVI as a general (linear) proxy of caribou forage conditions across the growing season, and encourage practitioners to use NDVI variables to model spatiotemporal variation in specific forage conditions from empirical field data, accounting for nonlinear forage-NDVI relationships.

Spatiotemporal variation in forage is a primary driver of ungulate behavior, yet little is known about the nutritional components they select, and how selection varies across the growing season with changes in forage quality and quantity. We addressed these uncertainties in barren‐ground caribou (Rangifer tarandus), which experience their most important foraging opportunities during the short Arctic summer. Recent declines in Arctic caribou populations have raised concerns about the influence of climate change on summer foraging opportunities, given shifting vegetation conditions and insect harassment, and their potential effects on caribou body condition and demography. We examined Arctic caribou selection of summer forage by pairing locations from females in the Central Arctic Herd of Alaska with spatiotemporal predictions of biomass, digestible nitrogen (DN), and digestible energy (DE). We then assessed selection for these nutritional components across the growing season at landscape and patch scales, and determined whether foraging opportunities were constrained by insect harassment. During early summer, at the landscape scale, caribou selected for intermediate biomass and high DN and DE, following expectations of the forage maturation hypothesis. At the patch scale, however, caribou selected for high values of all forage components, particularly DN, suggesting that protein may be limiting. During late summer, after DN declined below the threshold for protein gain, caribou exhibited a switch at both spatial scales, selecting for higher biomass, likely enabling mass and fat deposition. Mosquito activity strongly altered caribou selection of forage and increased their movement rates, while oestrid fly activity had little influence. Our results demonstrate that early and late summer periods afford Arctic caribou distinct foraging opportunities, as they prioritize quality earlier in the summer and quantity later. Climate change may further constrain caribou access to DN as earlier, warmer Arctic summers may be associated with reduced DN and increased mosquito harassment.

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