David D. Gustine

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.

Interpreting Resource Selection at Different Scales for Woodland Caribou in Winter

Abstract Scientists commonly use resource selection functions (RSFs) to identify areas important to large herbivores. Defining availability of resources is scale dependent and may limit inference on biological mechanisms of selection, particularly if variation in selection of resources is high among individuals within a population. We used logistic regression, the information-theoretic approach, and Global Positioning System (GPS) radiotelemetry data from 10 female woodland caribou (Rangifer tarandus caribou) and 5 wolf (Canis lupus) packs to model resource selection by individual caribou in the winter and late-winter seasons. We evaluated the influence of spatial scale on the relative importance of cost of movement and components of predation risk. We examined attributes of the risk of wolf predation within availability data at 2 spatial scales, and quantified variation in resource selection among individual caribou. Energetic cost of movement was the most important covariate for all caribou at a spatial scale defined by seasonal movement. Increasing distance to areas of high wolf risk was more important at the larger spatial scale of home range. Variation was high in the selection of resources among caribou, although commonalities among individuals enabled pooling data on use and availability into 2 selection strategies. Researchers and managers should conduct multiscale analyses with varied definitions of availability, quantify variation among individuals, and pool data into common selection strategies to identify mechanisms of selection and to map a populations selection for resources on the landscape.

Habitat selection by female Stone's sheep in relation to vegetation, topography, and risk of predation

ABSTRACT Stones sheep (Ovis dalli stonei) are susceptible to anthropogenic disturbances, but few data have been available to help minimize impacts. We used the movements of individuals to define use and availability, and resource-selection functions (RSF) and the information-theoretic approach to examine seasonal habitat selection and interannual variation in selection of attributes by groups of Stones sheep. Movement rates of Stones sheep followed consistent yearly patterns, with highest rates occurring in summer and fall. Models that contained vegetation, topography, and risk of predation best explained resource selection. Topographic features ranked better, however, than components of vegetation or risk of predation from grizzly bears (Ursus arctos) and wolves (Canis lupus) at explaining habitat selection. Selection strategies were variable among groups of Stones sheep and between years within groups even though consistencies in selection for steep slopes, ridge-like topography, south aspects, and upper elevations were common. This research provides the first comprehensive analysis of habitat selection by Stones sheep, which show strong fidelity to seasonal ranges but also exhibit behavioural plasticity in selection of attributes within those ranges. Nomenclature: Hultén, 1974; Nowak & Paradiso, 1983.

Range Expansion of Moose in Arctic Alaska Linked to Warming and Increased Shrub Habitat

Twentieth century warming has increased vegetation productivity and shrub cover across northern tundra and treeline regions, but effects on terrestrial wildlife have not been demonstrated on a comparable scale. During this period, Alaskan moose (Alces alces gigas) extended their range from the boreal forest into tundra riparian shrub habitat; similar extensions have been observed in Canada (A. a. andersoni) and Eurasia (A. a. alces). Northern moose distribution is thought to be limited by forage availability above the snow in late winter, so the observed increase in shrub habitat could be causing the northward moose establishment, but a previous hypothesis suggested that hunting cessation triggered moose establishment. Here, we use recent changes in shrub cover and empirical relationships between shrub height and growing season temperature to estimate available moose habitat in Arctic Alaska c. 1860. We estimate that riparian shrubs were approximately 1.1 m tall c. 1860, greatly reducing the available forage above the snowpack, compared to 2 m tall in 2009. We believe that increases in riparian shrub habitat after 1860 allowed moose to colonize tundra regions of Alaska hundreds of kilometers north and west of previous distribution limits. The northern shift in the distribution of moose, like that of snowshoe hares, has been in response to the spread of their shrub habitat in the Arctic, but at the same time, herbivores have likely had pronounced impacts on the structure and function of these shrub communities. These northward range shifts are a bellwether for other boreal species and their associated predators.

An Isotopic Approach to Measuring Nitrogen Balance in Caribou

ABSTRACT Nutritional restrictions in winter may reduce the availability of protein for reproduction and survival in northern ungulates. We refined a technique that uses recently voided excreta on snow to assess protein status in wild caribou (Rangifer tarandus) in late winter. Our study was the first application of this non-invasive, isotopic approach to assess protein status of wild caribou by determining dietary and endogenous contributions of nitrogen (N) to urinary urea. We used isotopic ratios of N (&dgr;15N) in urine and fecal samples to estimate the proportion of urea N derived from body N (p-UN) in pregnant, adult females of the Chisana Herd, a small population that ranged across the Alaska-Yukon border. We took advantage of a predator-exclosure project to examine N status of penned caribou in April 2006. Lichens were the primary forage (>40%) consumed by caribou in the pen and &dgr;15N of fiber tracked the major forages in their diets. The &dgr;15N of urinary urea for females in the pen was depleted relative (-1.3 ± 1.0 parts per thousand [‰], ±SD) to the &dgr;15N of body N (2.7 ±0.7‰). A similar proportion of animals in the exclosure lost core body mass (excluding estimates of fetal and uterine tissues; 55%) and body protein (estimated by isotope ratios; 54%). This noninvasive technique could be applied at various spatial and temporal scales to assess trends in protein status of free-ranging populations of northern ungulates. Intra- and inter-annual estimates of protein status could help managers monitor effects of foraging conditions on nutritional constraints in ungulates, increase the efficiency and efficacy of management actions, and help prepare stakeholders for potential changes in population trends.

Climate-driven effects of fire on winter habitat for caribou in the Alaskan-Yukon Arctic.

Climatic warming has direct implications for fire-dominated disturbance patterns in northern ecosystems. A transforming wildfire regime is altering plant composition and successional patterns, thus affecting the distribution and potentially the abundance of large herbivores. Caribou (Rangifer tarandus) are an important subsistence resource for communities throughout the north and a species that depends on terrestrial lichen in late-successional forests and tundra systems. Projected increases in area burned and reductions in stand ages may reduce lichen availability within caribou winter ranges. Sufficient reductions in lichen abundance could alter the capacity of these areas to support caribou populations. To assess the potential role of a changing fire regime on winter habitat for caribou, we used a simulation modeling platform, two global circulation models (GCMs), and a moderate emissions scenario to project annual fire characteristics and the resulting abundance of lichen-producing vegetation types (i.e., spruce forests and tundra >60 years old) across a modeling domain that encompassed the winter ranges of the Central Arctic and Porcupine caribou herds in the Alaskan-Yukon Arctic. Fires were less numerous and smaller in tundra compared to spruce habitats throughout the 90-year projection for both GCMs. Given the more likely climate trajectory, we projected that the Porcupine caribou herd, which winters primarily in the boreal forest, could be expected to experience a greater reduction in lichen-producing winter habitats (−21%) than the Central Arctic herd that wintered primarily in the arctic tundra (−11%). Our results suggest that caribou herds wintering in boreal forest will undergo fire-driven reductions in lichen-producing habitats that will, at a minimum, alter their distribution. Range shifts of caribou resulting from fire-driven changes to winter habitat may diminish access to caribou for rural communities that reside in fire-prone areas.

Habitat selection by a focal predator (Canis lupus) in a multiprey ecosystem of the northern Rockies

Abstract Large predators respond to land cover and physiography that maximize the likelihood of encountering prey. Using locations from global positioning system–collared wolves (Canis lupus), we examined whether land cover, vegetation productivity or change, or habitat-selection value for ungulate prey species themselves most influenced patterns of selection by wolves in a large, intact multiprey system of northern British Columbia. Selection models based on land cover, in combination with topographical features, consistently outperformed models based on indexes of vegetation quantity and quality (using normalized difference vegetation index) or on selection value to prey species (moose [Alces americanus], elk [Cervus elaphus], woodland caribou [Rangifer tarandus], and Stones sheep [Ovis dalli stonei]). Wolves generally selected for shrub communities and high diversity of cover across seasons and avoided conifer stands and non-vegetated areas and west aspects year-round. Seasonal selection strategies were not always reflected in use patterns, which showed highest frequency of use in riparian, shrub, and conifer classes. Patterns of use and selection for individual wolf packs did not always conform to global models, and appeared related to the distribution of land cover and terrain within respective home ranges. Our findings corroborate the biological linkages between wolves and their habitat related to ease of movement and potential prey associations.

Using ultrasound measurements of rump fat to assess nutritional condition of woodland caribou in northern British Columbia, Canada

Body reserves (fat and protein) of cervids are important to the reproductive success of individuals, and therefore may limit productivity of populations. We used a portable ultrasound machine to measure thickness of rump fat for 39 woodland caribou (Rangifer tarandus caribou) captured in the winters (January–February) of 2003 and 2004. We compared thickness of rump fat between pregnant and non-pregnant individuals in the Besa-Prophet drainage of northern British Columbia, Canada. Thirty-eight of the 39 females captured in British Columbia were adults and 34 of the adult caribou were pregnant (89.5 ± 5.1%, x– ± binomial SE). Pregnant individuals had more rump fat (0.60 ± 0.067 cm) than nonpregnant animals (0.20 ± 0.029 cm). Recognizing that deposition and mobilization of fat vary with age and possibly across the winter season, ultrasonography can be used as a non-invasive technique in the field to assist in estimating body fat of caribou.

Advancing the match-mismatch framework for large herbivores in the Arctic: Evaluating the evidence for a trophic mismatch in caribou

Climate-induced shifts in plant phenology may adversely affect animals that cannot or do not shift the timing of their reproductive cycle. The realized effect of potential trophic “mismatches” between a consumer and its food varies with the degree to which species rely on dietary income and stored capital. Large Arctic herbivores rely heavily on maternal capital to reproduce and give birth near the onset of the growing season but are they vulnerable to trophic mismatch? We evaluated the long-term changes in the temperatures and characteristics of the growing seasons (1970–2013), and compared growing conditions and dynamics of forage quality for caribou at peak parturition, peak lactation, and peak forage biomass, and plant senescence between two distinct time periods over 36 years (1977 and 2011–13). Despite advanced thaw dates (7−12 days earlier), increased growing season lengths (15−21 days longer), and consistent parturition dates, we found no decline in forage quality and therefore no evidence within this dataset for a trophic mismatch at peak parturition or peak lactation from 1977 to 2011–13. In Arctic ungulates that use stored capital for reproduction, reproductive demands are largely met by body stores deposited in the previous summer and autumn, which reduces potential adverse effects of any mismatch between food availability and timing of parturition. Climate-induced effects on forages growing in the summer and autumn ranges, however, do correspond with the demands of female caribou and their offspring to gain mass for the next reproductive cycle and winter. Therefore, we suggest the window of time to examine the match-mismatch framework in Arctic ungulates is not at parturition but in late summer-autumn, where the multiplier effects of small changes in forage quality are amplified by forage abundance, peak forage intake, and resultant mass gains in mother-offspring pairs.

Diversity of nitrogen isotopes and protein status in caribou: implications for monitoring northern ungulates

Abstract Nutritional condition is an important determinant of productivity and survival in caribou (Rangifer tarandus). We used samples of excreta (n  =  1,150) to estimate diet composition from microhistology and 2 isotopic proxies of protein status for 2 ecotypes of caribou in 4 herds in late winter (2006–2008). Isotopes of nitrogen (&dgr;15N in parts per thousand [‰]) from excreta samples (urea, diet, and body N) were used to estimate indexes of protein status: the proportion of urea N derived from body N (p-UN) and the difference between the &dgr;15N of the body and urinary urea (&Dgr;body-urea). We examined dietary and terrain characteristics, &dgr;15N, p-UN, and &Dgr;body-urea by ecotype, herd, year, and foraging site. Multiple regression and an information-theoretic approach were used to evaluate correlates of protein status at each foraging site. The dietary and terrain characteristics of foraging sites did not vary by ecotype or herd (P > 0.108); diets were dominated by lichens (68% ± 14.1% SD). The &dgr;15N of urea, diet, body N, p-UN, and &Dgr;body-urea varied among foraging sites within each herd (P < 0.001). Although highly variable, the &dgr;15N of urinary urea was typically low (−4.68‰ ± 2.67‰ SD). Dietary N also had low &dgr;15N (−4.18‰ ± 0.92‰ SD), whereas body N was generally heavier in 15N (2.20‰ ± 1.56‰ SD) than urinary urea or the diet. Both measures of protein status were similarly diverse between ecotypes and among herds, which limited their applicability to monitor protein status at the population level. Although we observed limitations to interpreting estimates of p-UN from highly vagile ungulates, the &Dgr;body-urea may prove to be a useful index of protein status at smaller spatial and temporal scales. Indeed, a portion of the observed variance (r2  =  0.26) in &Dgr;body-urea at each foraging site was explained by the proportion of shrubs in the winter diet. There remains potential in using &dgr;15N in excreta as a noninvasive tool for evaluating protein status in northern ungulates; however, considerable analytical and sampling challenges remain for applying these isotopic approaches at large scales.