Quantifying effects of snow depth on caribou winter range selection and movement in Arctic Alaska.

BACKGROUND: Caribou and reindeer across the Arctic spend more than two thirds of their lives moving in snow. Yet snow-specific mechanisms driving their winter ecology and potentially influencing herd health and movement patterns are not well known. Integrative research coupling snow and wildlife sciences using observations, models, and wildlife tracking technologies can help fill this knowledge void. METHODS: Here, we quantified the effects of snow depth on caribou winter range selection and movement. We used location data of Central Arctic Herd (CAH) caribou in Arctic Alaska collected from 2014 to 2020 and spatially distributed and temporally evolving snow depth data produced by SnowModel. These landscape-scale (90 m), daily snow depth data reproduced the observed spatial snow-depth variability across typical areal extents occupied by a wintering caribou during a 24-h period. RESULTS: We found that fall snow depths encountered by the herd north of the Brooks Range exerted a strong influence on selection of two distinct winter range locations. In winters with relatively shallow fall snow depth (2016/17, 2018/19, and 2019/20), the majority of the CAH wintered on the tundra north of the Brooks Range mountains. In contrast, during the winters with relatively deep fall snow depth (2014/15, 2015/16, and 2017/18), the majority of the CAH caribou wintered in the mountainous boreal forest south of the Brooks Range. Long-term (19 winters; 2001-2020) monitoring of CAH caribou winter distributions confirmed this relationship. Additionally, snow depth affected movement and selection differently within these two habitats: in the mountainous boreal forest, caribou avoided areas with deeper snow, but when on the tundra, snow depth did not trigger significant deep-snow avoidance. In both wintering habitats, CAH caribou selected areas with higher lichen abundance, and they moved significantly slower when encountering deeper snow. CONCLUSIONS: In general, our findings indicate that regional-scale selection of winter range is influenced by snow depth at or prior to fall migration. During winter, daily decision-making within the winter range is driven largely by snow depth. This integrative approach of coupling snow and wildlife observations with snow-evolution and caribou-movement modeling to quantify the multi-facetted effects of snow on wildlife ecology is applicable to caribou and reindeer herds throughout the Arctic.

Environment and physiology shape Arctic ungulate population dynamics.

Species conservation in a rapidly changing world requires an improved understanding of how individuals and populations respond to changes in their environment across temporal scales. Increased warming in the Arctic puts this region at particular risk for rapid environmental change, with potentially devastating impacts on resident populations. Here, we make use of a parameterized full life cycle, individual-based energy budget model for wild muskoxen, coupling year-round environmental data with detailed ontogenic metabolic physiology. We show how winter food accessibility, summer food availability, and density dependence drive seasonal dynamics of energy storage and thus life history and population dynamics. Winter forage accessibility defined by snow depth, more than summer forage availability, was the primary determinant of muskox population dynamics through impacts on calf recruitment and longer term carryover effects of maternal investment. Simulations of various seasonal snow depth and plant biomass and quality profiles revealed that timing of and improved/limited winter forage accessibility had marked influence on calf recruitment (±10-80%). Impacts on recruitment were the cumulative result of condition-driven reproductive performance at multiple time points across the reproductive period (ovulation to calf weaning) as a trade-off between survival and reproduction. Seasonal and generational condition effects of snow-rich winters interacted with age structure and density to cause pronounced long-term consequences on population growth and structure, with predicted population recovery times from even moderate disturbances of 10 years or more. Our results show how alteration in winter forage accessibility, mediated by snow depth, impacts the dynamics of northern herbivore populations. Further, we present here a mechanistic and state-based model framework to assess future scenarios of environmental change, such as increased or decreased snowfall or plant biomass and quality to impact winter and summer forage availability across the Arctic.

Winter snow and spring temperature have differential effects on vegetation phenology and productivity across Arctic plant communities.

Tundra dominates two-thirds of the unglaciated, terrestrial Arctic. Although this region has experienced rapid and widespread changes in vegetation phenology and productivity over the last several decades, the specific climatic drivers responsible for this change remain poorly understood. Here we quantified the effect of winter snowpack and early spring temperature conditions on growing season vegetation phenology (timing of the start, peak, and end of the growing season) and productivity of the dominant tundra vegetation communities of Arctic Alaska. We used daily remotely sensed normalized difference vegetation index (NDVI), and daily snowpack and temperature variables produced by SnowModel and MicroMet, coupled physically based snow and meteorological modeling tools, to (1) determine the most important snowpack and thermal controls on tundra vegetation phenology and productivity and (2) describe the direction of these relationships within each vegetation community. Our results show that soil temperature under the snowpack, snowmelt timing, and air temperature following snowmelt are the most important drivers of growing season timing and productivity among Arctic vegetation communities. Air temperature after snowmelt was the most important control on timing of season start and end, with warmer conditions contributing to earlier phenology in all vegetation communities. In contrast, the controls on the timing of peak season and productivity also included snowmelt timing and soil temperature under the snowpack, dictated in part by the snow insulating capacity. The results of this novel analysis suggest that while future warming effects on phenology may be consistent across communities of the tundra biome, warming may result in divergent, community-specific productivity responses if coupled with reduced snow insulating capacity lowers winter soil temperature and potential nutrient cycling in the soil.

Environmental conditions alter behavioural organization and rhythmicity of a large Arctic ruminant across the annual cycle.

The existence and persistence of rhythmicity in animal activity during phases of environmental change is of interest in ecology, evolution and chronobiology. A wide diversity of biological rhythms in response to exogenous conditions and internal stimuli have been uncovered, especially for polar vertebrates. However, empirical data supporting circadian organization in behaviour of large ruminating herbivores remains inconclusive. Using year-round tracking data of the largest Arctic ruminant, the muskox (Ovibos moschatus), we modelled rhythmicity as a function of behaviour and environmental conditions. Behavioural states were classified based on patterns in hourly movements, and incorporated within a periodicity analyses framework. Although circadian rhythmicity in muskox behaviour was detected throughout the year, ultradian rhythmicity was most prevalent, especially when muskoxen were foraging and resting in mid-winter (continuous darkness). However, when combining circadian and ultradian rhythmicity together, the probability of behavioural rhythmicity declined with increasing photoperiod until largely disrupted in mid-summer (continuous light). Individuals that remained behaviourally rhythmic during mid-summer foraged in areas with lower plant productivity (NDVI) than individuals with arrhythmic behaviour. Based on our study, we conclude that muskoxen may use an interval timer to schedule their behavioural cycles when forage resources are low, but that the importance and duration of this timer are reduced once environmental conditions allow energetic reserves to be replenished ad libitum. We argue that alimentary function and metabolic requirements are critical determinants of biological rhythmicity in muskoxen, which probably applies to ruminating herbivores in general.

An application of upscaled optimal foraging theory using hidden Markov modelling: year-round behavioural variation in a large arctic herbivore.

BACKGROUND: In highly seasonal environments, animals face critical decisions regarding time allocation, diet optimisation, and habitat use. In the Arctic, the short summers are crucial for replenishing body reserves, while low food availability and increased energetic demands characterise the long winters (9-10 months). Under such extreme seasonal variability, even small deviations from optimal time allocation can markedly impact individuals' condition, reproductive success and survival. We investigated which environmental conditions influenced daily, seasonal, and interannual variation in time allocation in high-arctic muskoxen (Ovibos moschatus) and evaluated whether results support qualitative predictions derived from upscaled optimal foraging theory. METHODS: Using hidden Markov models (HMMs), we inferred behavioural states (foraging, resting, relocating) from hourly positions of GPS-collared females tracked in northeast Greenland (28 muskox-years). To relate behavioural variation to environmental conditions, we considered a wide range of spatially and/or temporally explicit covariates in the HMMs. RESULTS: While we found little interannual variation, daily and seasonal time allocation varied markedly. Scheduling of daily activities was distinct throughout the year except for the period of continuous daylight. During summer, muskoxen spent about 69% of time foraging and 19% resting, without environmental constraints on foraging activity. During winter, time spent foraging decreased to 45%, whereas about 43% of time was spent resting, mediated by longer resting bouts than during summer. CONCLUSIONS: Our results clearly indicate that female muskoxen follow an energy intake maximisation strategy during the arctic summer. During winter, our results were not easily reconcilable with just one dominant foraging strategy. The overall reduction in activity likely reflects higher time requirements for rumination in response to the reduction of forage quality (supporting an energy intake maximisation strategy). However, deep snow and low temperatures were apparent constraints to winter foraging, hence also suggesting attempts to conserve energy (net energy maximisation strategy). Our approach provides new insights into the year-round behavioural strategies of the largest Arctic herbivore and outlines a practical example of how to approximate qualitative predictions of upscaled optimal foraging theory using multi-year GPS tracking data.

Transitions in high-Arctic vegetation growth patterns and ecosystem productivity tracked with automated cameras from 2000 to 2013.

Climate-induced changes in vegetation phenology at northern latitudes are still poorly understood. Continued monitoring and research are therefore needed to improve the understanding of abiotic drivers. Here we used 14 years of time lapse imagery and climate data from high-Arctic Northeast Greenland to assess the seasonal response of a dwarf shrub heath, grassland, and fen, to inter-annual variation in snow-cover, soil moisture, and air and soil temperatures. A late snow melt and start of growing season is counterbalanced by a fast greenup and a tendency to higher peak greenness values. Snow water equivalents and soil moisture explained up to 77 % of growing season duration and senescence phase, highlighting that water availability is a prominent driver in the heath site, rather than temperatures. We found a significant advance in the start of spring by 10 days and in the end of fall by 11 days, resulting in an unchanged growing season length. Vegetation greenness, derived from the imagery, was correlated to primary productivity, showing that the imagery holds valuable information on vegetation productivity.

Changing Arctic snow cover: A review of recent developments and assessment of future needs for observations, modelling, and impacts.

Snow is a critically important and rapidly changing feature of the Arctic. However, snow-cover and snowpack conditions change through time pose challenges for measuring and prediction of snow. Plausible scenarios of how Arctic snow cover will respond to changing Arctic climate are important for impact assessments and adaptation strategies. Although much progress has been made in understanding and predicting snow-cover changes and their multiple consequences, many uncertainties remain. In this paper, we review advances in snow monitoring and modelling, and the impact of snow changes on ecosystems and society in Arctic regions. Interdisciplinary activities are required to resolve the current limitations on measuring and modelling snow characteristics through the cold season and at different spatial scales to assure human well-being, economic stability, and improve the ability to predict manage and adapt to natural hazards in the Arctic region.