Corrigendum to: State-dependent foraging by caribou with different nutritional requirements.

[This corrects the article DOI: 10.1093/jmammal/gyaa003.].

Wave-like Patterns of Plant Phenology Determine Ungulate Movement Tactics.

Animals exhibit a diversity of movement tactics [1]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [2]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called "green-wave surfing" [3-5]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [6]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [7]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [8, 9]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [10], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

State-dependent foraging by caribou with different nutritional requirements.

Foraging by animals is hypothesized to be state-dependent, that is, varying with physiological condition of individuals. State often is defined by energy reserves, but state also can reflect differences in nutritional requirements (e.g., for reproduction, lactation, growth, etc.). Testing hypotheses about state-dependent foraging in ungulates is difficult because fine-scale data needed to evaluate these hypotheses generally are lacking. To evaluate whether foraging by caribou (Rangifer tarandus) was state-dependent, we compared bite and intake rates, travel rates, dietary quality, forage selection, daily foraging time, and foraging strategies of caribou with three levels of nutritional requirements (lactating adults, nonlactating adults, subadults 1-2 years old). Only daily foraging times and daily nutrient intakes differed among nutritional classes of caribou. Lactating caribou foraged longer per day than nonlactating caribou-a difference that was greatest at the highest rates of intake, but which persisted even when intake was below requirements. Further, at sites where caribou achieved high rates of intake, caribou in each nutritional class continued foraging even after satisfying daily nutritional requirements, which was consistent with a foraging strategy to maximize energy intake. Foraging time by caribou was partially state-dependent, highlighting the importance of accounting for physiological state in studies of animal behavior. Fine-scale foraging behaviors may influence larger-scale behavioral strategies, with potential implications for conservation and management.

Understanding predation risk and individual variation in risk avoidance for threatened boreal caribou.

Predation risk is a driver of species' distributions. Animals can increase risk avoidance in response to fluctuations in predation risk, but questions remain regarding individual variability and the capacity to respond to changes in spatial risk across human-altered landscapes. In northeast British Columbia, Canada, boreal caribou populations declined as roads and seismic lines have increased, which are theorized to increase gray wolf predation. Our goal was to model risk and to evaluate individual variability and the development of risk perception by examining individual risk avoidance in response to reproductive status and age. We used locations from collared caribou and wolves to identify landscape features associated with the risk of a potential wolf-caribou encounter and risk of being killed given an encounter. We built resource selection functions to estimate individual responses to risk. We used general linear regressions to evaluate individual risk and linear feature avoidance as a function of age and reproductive status (calf or no calf). Linear features increased the risk of encounter. Older caribou and caribou with calves demonstrated stronger avoidance of the risk of encounter and roads, but weaker avoidance in late summer to the risk of being killed relative to younger and calf-less individuals. Mechanisms explaining the inverse relationships between the risk of encounter and risk of being killed are uncertain, but it is conceivable that caribou learn to avoid the risk of encounter and roads. Responses by females with vulnerable calves to the risk of encounter and risk of being killed might be explained by a trade-off between these two risk types and a prioritization on the risk of encounter. Despite the capacity to alter their responses to risk, the global decline in Rangifer populations (caribou and wild reindeer) suggests these behaviors are insufficient to mitigate the impacts of anthropogenic disturbances.

Hand-rearing wild caribou calves for studies of nutritional ecology.

Caribou (Rangifer tarandus) are considered difficult to hand-rear in captivity because they are sensitive to the composition and volume of formulated milks. We used a strict feeding schedule and a commercial milk formula designed specifically for caribou to bottle-feed neonates captured from five wild caribou herds in Alaska. Under a feeding protocol adjusted for age and mass, the growth rates and body mass of 26 hand-reared caribou calves to weaning were similar to those of three maternally nursed caribou. This protocol allows caretakers to hand-rear caribou that are as representative as possible of maternally raised neonates.

Allocating protein to reproduction in arctic reindeer and caribou.

Reindeer (Rangifer tarandus tarandus) and caribou (Rangifer tarandus granti) use body stores (capital) and food intake (income) for survival and reproduction. Intakes of low-nitrogen (N) food declined in winter and increased in spring (51-83 g dry matter kg(-0.75) d(-1)). Reindeer calved before regaining food intake, whereas caribou calved 28 d later. Body N was conserved by minimizing oxidation of amino acid N to urea. Maternal protein stored from early winter was used for 96% of fetal growth in reindeer but only 84% of fetal growth in later-birthing caribou. Both subspecies rely on maternal body protein for 91% of the protein deposited in the neonate via milk over the first 4 wk. All females lost body protein over winter, but lactating females continued to lose protein while nonreproductive females regained protein. Net costs of lactation above maintenance were greater for N (110%-130%) than for energy (40%-59%). Large fat stores in reindeer spare body protein from oxidation in winter, whereas in caribou, less fat with the same body protein favors migration when food is inadequate. The resilience of Rangifer populations to variable patterns of food supply and metabolic demand may be related to their ability to alter the timing and allocation of body protein to reproduction.

Body protein stores and isotopic indicators of N balance in female reindeer (Rangifer tarandus) during winter.

We studied bred and unbred female reindeer (Rangifer tarandus tarandus) during 12 wk of winter when ambient temperatures were low and nitrogen (N) demand for fetal growth is highest in pregnant females. Animals were fed a complete pelleted diet ad lib. that contained 2.54% N in dry matter that was 80% +/- 2% (X +/- SD) digestible. Female reindeer lost 64% +/- 14% of body fat but gained 34% +/- 11% of lean mass from 10 wk prepartum to parturition. These changes were equivalent to average balances of -14.14 +/- 2.35 MJ d(-1) and 10 +/- 3 g N d(-1). Blood cells, serum, and urine declined in (15)N/(14)N in late winter as body protein was gained from the diet. Blood cells of newborn calves were more enriched in (15)N and (13)C than that of their mothers, indicating the deposition of fetal protein from maternal stores. To quantify pathways of N flow in reindeer, N balance was measured by confining animals to cages for 10 d at 4 wk from parturition. N balance was inversely related to (15)N/(14)N in urea-N but not related to (15)N/(14)N of blood cells, creatinine, and feces. The proportion of urea-N derived from body protein increased above 0.46 as N balance fell below -200 mg N kg(-0.75) d(-1). Proportions of urea-N from body protein were -0.01 +/- 0.21 in pregnant females before and after caging and were consistent with average body protein gain in winter. Storage of protein allows reindeer and caribou to tolerate diets that are low in N without impairing fetal development.

Foraging across a variable landscape: behavioral decisions made by woodland caribou at multiple spatial scales.

We examined the foraging behavior of woodland caribou (Rangifer tarandus caribou) relative to the spatial and temporal heterogeneity of their environment. We assessed (1) whether caribou altered their behavior over time while making trade-offs between forage abundance and accessibility; and (2) whether foraging decisions were consistent across spatial scales (i.e., as scale increased, similar decision criteria were used at each scale). We discuss whether caribou adjusted their behavior to take advantage of changing forage availability through time and space. At the scale of the feeding site (as revealed by discriminant function analyses), caribou in both forested and alpine (above tree-line) environments selected sites where the biomass of particular lichen species was greatest and snow the least deep. Caribou did not select those species with the highest nutritional value (i.e., digestible protein and energy) in either area. Where snow depth, density, and hardness limited access to terrestrial lichens in the forest, caribou foraged instead at those trees with the greatest amount of arboreal lichen. Selection of lichen species and the influence of snow differed across time, indicating that in this system the abundance or accessibility of forage temporally influenced foraging behavior. A path analysis of forest data and multiple regression analysis of alpine data were used to test the hypothesis that variables important at the scale of the feeding site explained foraging effort at the scale of the patch. For forest patches, our hypothesized model reliably explained foraging effort, but not all variables that were statistically important at the scale of the feeding site were significant predictors at the scale of the patch. For alpine patches, our hypothesized model did not explain a statistically significant portion of the variation in the number of feeding sites within the patch, and none of the individual variables from the feeding site remained statistically significant at the patch scale. The incongruity between those variables important at the scale of the feeding site and those important at the patch showed that spatial scale affects the foraging decisions of woodland caribou. At the scale of the landscape, there was a trade-off between forage abundance and accessibility. Relative to the alpine environment, caribou in the forest foraged at feeding sites and patches with greater amounts of less variably distributed lichen, but deeper less variable snow depths. Considering the behavioral plasticity of woodland caribou, there may be no distinct advantage to foraging in one landscape over the other.