Katherine L. Parker

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.

A MULTISCALE BEHAVIORAL APPROACH TO UNDERSTANDING THE MOVEMENTS OF WOODLAND CARIBOU

We assessed the response of woodland caribou (Rangifer tarandus caribou) to land-cover type, predation risk, energetic costs of movement, and patch configuration at multiple spatial scales. We applied a nonlinear model to frequent locations collected with Global Positioning System (GPS) collars to identify discontinuities in the scales of movement by caribou found in forested and alpine (above tree line) habitats. We differ- entiated intra- from interpatch movements and identified collections of patches (multiple- patch scale) where caribou concentrated intrapatch movements. On average, intra- and interpatch movements were 450.7 and 1268.8 m, respectively, and multiple-patch move- ments occurred over an area of 182 ha. Intrapatch movements were highly correlated, indicative of a strong relationship between behavior and place. Caribou in the forest selected patches of Pine terrace, whereas caribou in the alpine selected patches of Alpine-little vegetative cover. Predation risk was not a factor influencing movements of caribou at the intrapatch scale. Selection of cover types was more variable during interpatch movements. At that scale, caribou selected patches of Pine terrace, Lakes/rivers, Alpine-little vegetative cover, and Alpine-grass. The routes selected by caribou had lower energetic costs relative to surrounding terrain, and during some winters, caribou were subjected to higher levels of predation risk during those movements. At the multiple-patch scale, selection was more specific and encompassed patches of Alpine-little vegetative cover, Alpine-grass, and Pine terrace. Predation risk was relatively unimportant at the multi-patch scale, but animals that moved from forested to alpine habitats reduced their relative risk of predation. Patch con- figuration was a poor predictor of those areas where caribou concentrated intrapatch move- ments. There was some evidence of caribou selecting patches of Pine terrace within a matrix of Wetlands and Pine-black spruce/black spruce patches. Caribou in the alpine avoided patches of Alpine-little vegetative cover adjacent to forest types. Our results indicate that forest managers should maintain widely distributed patches of Pine terrace and implement silvicultural regimes that do not stimulate predator populations across areas used for in- terpatch movements.

Calf Survival of Woodland Caribou in a Multi-Predator Ecosystem

Abstract The proximate role of predation in limiting caribou (Rangifer tarandus) populations is well documented, but the long-term effects of predation pressure on selection of calving areas and the subsequent impacts to calving success remain unclear. We examined the relationships among calf survival, predation risk, and vegetation characteristics among 3 calving areas and across spatial scales in the Besa-Prophet River drainage of northern British Columbia. Fifty woodland caribou (R. t. caribou) neonates were collared and monitored twice daily for the first month and once weekly during the next month of life in 2 summer field seasons (2002 and 2003). Predation risk was estimated using resource selection functions (RSFs) from Global Positioning System (GPS) locations of 15 grizzly bears (Ursus arctos) and 5 gray wolf (Canis lupus) packs. The Normalized Difference Vegetation Index (NDVI) derived from Landsat Thematic Mapper (TM) and Enhanced Thematic Mapper (ETM) data were used to quantify large-scale characteristics of vegetation (indices of biomass and quality). We incorporated small- and large-scale characteristics (i.e., predation risk, vegetation, and movement of woodland caribou calves) of neonatal calving sites into logistic regression models to predict survival for the calving (25 May–14 Jun) and summer (15 Jun–31 Jul) seasons. Predation risk and vegetation characteristics were highly variable among calving areas and calving sites, and parturient woodland caribou responded to these characteristics at different scales. Minimizing gray wolf risk and selecting against areas of high vegetation biomass were important at large scales; areas with high biomass were likely associated with increased predation risk. Calving in areas high in vegetation quality was important across scales, as parturient woodland caribou took higher levels of predation risk to access areas of high vegetative change. Models using small-scale characteristics of calving sites to predict survival performed better in the calving season than in summer. Large-scale characteristics predicted survival of woodland caribou neonates better in summer than in the calving season, probably in part because of the unexpected role of wolverines (Gulo gulo) as the main predator of woodland caribou calves during calving. Gray wolves were the main cause of mortality during the summer. Movement away from calving sites corresponded to higher calf survival and appeared to be in response to increased access to forage during the peak demands of lactation and/or minimizing gray wolf risk in the summer. High variation in predation risk and vegetation attributes among calving areas and at calving sites within calving areas, with no differences in calf mortality related to that variation, illustrates the importance of behavioral plasticity as a life-history strategy for woodland caribou.

PROTEIN CONSERVATION IN FEMALE CARIBOU (RANGIFER TARANDUS): EFFECTS OF DECREASING DIET QUALITY DURING WINTER

Abstract Female caribou subsist primarily on lichens and some senescent browse during winter when demands for fetal growth add to costs of thermoregulation and mobility. Lichens, although potentially high in digestible energy, contain less protein than required for maintenance by most north-temperate ungulates. To understand the adaptations of caribou to the nutritional constraints of their primary food resource, we fed captive female caribou a sequence of 3 diets designed to resemble decreasing quality of forages during early, mid-, and late winter, respectively: high energy–high protein (HIGH), medium energy–low protein (MEDIUM), and low energy–low protein (LOW). In vitro digestibility of dry matter declined from 94% (HIGH) in November, to 66% (MEDIUM) in December and January, and to 53% (LOW) from February to April. Dietary protein averaged 19.8% in November and 4.3% from December to April. We used measures of body condition, stable isotopic signatures, and concentrations of nitrogen (N) metabolites to define protein dynamics in the animals. Subcutaneous rump fat declined between October and April from 2.3 cm ± 0.3 SE to <0.5 cm as intake of digestible energy declined from 44.0 ± 2.0 MJ/day to 16.3 ± 3.2 MJ/day. In erythrocytes, increasing enrichment of carbon (13C) throughout winter suggested that caribou reused body lipids, and increases in 15N during January and February indicated that they also recycled amino-N. Urinary N was primarily urea with an isotopic signature that tracked dietary 15N through late winter. Plasma urea-N declined from 44.0 ± 2.6 mg/dl to 8.5 ± 1.2 mg/dl as nitrogen intake declined from 91.5 ± 5.3 g N/day to 14.1 ± 0.9 g N/day. Examination of these data suggests that caribou catabolized dietary C and N in preference to endogenous fat reserves and body protein. Female caribou appear to tolerate low intakes of protein and energy in winter by minimizing net loss of body protein and reapportioning body reserves to support fetal growth.

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 \documentclass{aastex} \usepackage{amsbsy} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{bm} \usepackage{mathrsfs} \usepackage{pifont} \usepackage{stmaryrd} \usepackage{textcomp} \usepackage{portland,xspace} \usepackage{amsmath,amsxtra} \usepackage[OT2,OT1]{fontenc} \newcommand\cyr{ \renewcommand\rmdefault{wncyr} \renewcommand\sfdefault{wncyss} \renewcommand\encodingdefault{OT2} \normalfont \selectfont} \DeclareTextFontCommand{\textcyr}{\cyr} \pagestyle{empty} \DeclareMathSizes{10}{9}{7}{6} \begin{document} \landscape

Expectations and Realities of GPS Animal Location Collars: Results of Three Years in the Field

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Estimates of critical thermal environments for mule deer.

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