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. Zoo Biol. 32:163–171, 2013. © 2012 Wiley Periodicals, Inc.     

Dynamic selection for forage quality and quantity in response to phenology and insects in an Arctic ungulate

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.     

Displacement of Mountain Caribou From Winter Habitat by Snowmobiles

Abstract: Mountain caribou are an ecotype of woodland caribou (Rangifer tarandus caribou) that live in subalpine forests in southeastern British Columbia, Canada, extending into northern Idaho and Washington, USA. These caribou are listed as Threatened in Canada, Endangered in the United States, and are the subject of recovery planning efforts in both countries. Many areas of mountain caribou winter habitat experience intensive use by recreational snowmobilers. During 4 surveys, we recorded caribou on all 4 census blocks with little or no snowmobile activity (x̄ density = 0.41 caribou/km²), but during 3 of 4 years, we observed no caribou on the census block with intensive snowmobile activity. The year we observed caribou on the snowmobile block, most were using areas inaccessible to snowmobiles. We used a Resource Selection Function (RSF) based on radiotelemetry data for the area to compare habitat quality among the different census blocks. The absence of caribou from the intensive snowmobile area during most years could not be explained by differences in habitat quality. The RSF predicted that the intensive snowmobile area could support 53-96 caribou (95% CI). We conclude that intensive snowmobiling has displaced caribou from an area of suitable habitat. We recommend that snowmobile activity be restricted from all or most high-quality mountain caribou habitat as part of the recovery planning process.     

Evaluation of Maternal Penning to Improve Calf Survival in the Chisana Caribou Herd

Predation is a major limiting factor for most small sedentary caribou (Rangifer tarandus) populations, particularly those that are threatened or endangered across the southern extent of the species’ range. Thus, reducing predation impacts is often a management goal for improving the status of small caribou populations, and lethal predator removal is the primary approach that has been applied. Given that predator control programs are often contentious, other management options that can garner broader public acceptance need to be considered. Substantial calf losses to predation in the few weeks following birth are common for these small caribou populations. Therefore, we employed a novel experimental approach of maternal penning with the goal of reducing early calf mortality in the Chisana Caribou Herd, a declining population in southwest Yukon and adjacent Alaska thought to number around 300 individuals. Maternal penning entailed temporarily holding pregnant females on their native range in a large pen secure from predators from late March through the initial weeks of calf rearing to mid-June. During 2003–2006, we conducted 4 annual penning trials with 17–50 pregnant females each year (n = 146 total), assessed survival of calves born in the pens, and evaluated survival and nutritional effects of penning for females that were held. We also investigated the herd's population dynamics during 2003–2008 to determine effects of maternal penning on calf recruitment and population growth. In addition to information gained during maternal penning, we determined natality and survival patterns via radiotelemetry, conducted autumn age-sex composition surveys each year, and censused the population in mid-October 2003, 2005, and 2007. Based on our penning trials and demographic investigations, we used simulation models to evaluate the effects of maternal penning relative to a population's inherent growth rate (finite rate of increase [λ] without maternal penning) and penning effort (proportion of calves born in penning) to provide perspective on utility of this approach for improving the status of small imperiled caribou populations. Pregnant females held in maternal penning tolerated captivity well in that they exhibited positive nutritional responses to ad libitum feed we provided and higher survival than free-ranging females (0.993 and 0.951 for penned and free-ranging females, respectively). Survival of pen calves from birth to mid-June was substantially higher than that of free-ranging calves ( = 0.950 and 0.376, respectively). This initial period accounted for 76% of the annual calf mortality in the free-ranging population. Pen-born calves maintained their survival advantage over wild-born calves to the end of their first year ( = 0.575 and 0.192, respectively) during years penning occurred. Females in the Chisana Herd were highly productive with 57% producing their first offspring at 2 years of age, and annual natality rates averaging 0.842 calves/female ≥2 years old. Age-specific natality rates exceeded 0.900 for 4–9-year-olds, then exhibited senescent decline to 0.467 by 19 years old. Annual survival of free-ranging adult females and calves averaged 0.892 and 0.184, respectively, over all study years; both were reduced during 2004 because of poor winter survival. We noted reduced nutritional condition of caribou late that winter in that females we captured were lighter than in other years and produced lighter calves. We suspect that the reduced survival during winter 2004 and the observed nutritional characteristics resulted from adverse snow conditions in combination with effects of the extreme drought experienced the previous summer. Age-specific survival of adult females was ≥0.900 through 10 years of age, then declined with age. The Chisana Herd numbered 720 caribou in mid-October 2003, or more than twice that estimated prior to initiating maternal penning, and increased to 766 caribou by mid-October 2007. We calculated that penning added 54.2 yearling recruits, or 40% of calves released from penning. Based on the maternal penning results and the population's vital rates, we determined that the herd would have been stable during 2003–2007 at about 713 caribou without maternal penning; thus, the increase in herd size we observed resulted from maternal penning and was equivalent to the estimate of additional yearling recruits. The improvement in the population trend invoked by maternal penning was limited by the larger than expected population size and resulting low penning effort ( = 11% of calves born in pen). Our simulations corroborated that maternal penning increased population size by the number of additional recruits provided, even at low penning effort, for inherently stable populations. As the inherent rate of increase dropped below λ = 1.000, more of the additional recruits from penning were needed to offset the downward population inertia, thus requiring increased penning effort to reach stability. For populations declining at λ < 0.890, stability could not be achieved with 100% penning effort given the vital rates in our models. Maternal penning in its limited application to date has proven to be broadly popular as a nonlethal management action aimed at reducing initial calf mortality from predation in small caribou populations. However, based on the Chisana program and 3 subsequent efforts elsewhere, improvement in population trends have been modest at best and come at a high financial cost. Given the necessity of maximizing penning effort, maternal penning may have a role in addressing conservation challenges for some small caribou populations that are stable or slowly declining, but its application should be primarily driven by objective assessment of the likelihood of improving population trends rather than popularity relative to other management options.     

Critical summer foraging tradeoffs in a subarctic ungulate

Summer diets are crucial for large herbivores in the subarctic and are affected by weather, harassment from insects and a variety of environmental changes linked to climate. Yet, understanding foraging behavior and diet of large herbivores is challenging in the subarctic because of their remote ranges. We used GPS video‐camera collars to observe behaviors and summer diets of the migratory Fortymile Caribou Herd (Rangifer tarandus granti) across Alaska, USA and the Yukon, Canada. First, we characterized caribou behavior. Second, we tested if videos could be used to quantify changes in the probability of eating events. Third, we estimated summer diets at the finest taxonomic resolution possible through videos. Finally, we compared summer diet estimates from video collars to microhistological analysis of fecal pellets. We classified 18,134 videos from 30 female caribou over two summers (2018 and 2019). Caribou behaviors included eating (mean = 43.5%), ruminating (25.6%), travelling (14.0%), stationary awake (11.3%) and napping (5.1%). Eating was restricted by insect harassment. We classified forage(s) consumed in 5,549 videos where diet composition (monthly) highlighted a strong tradeoff between lichens and shrubs; shrubs dominated diets in June and July when lichen use declined. We identified 63 species, 70 genus and 33 family groups of summer forages from videos. After adjusting for digestibility, monthly estimates of diet composition were strongly correlated at the scale of the forage functional type (i.e., forage groups composed of forbs, graminoids, mosses, shrubs and lichens; r = 0.79, p < .01). Using video collars, we identified (1) a pronounced tradeoff in summer foraging between lichens and shrubs and (2) the costs of insect harassment on eating. Understanding caribou foraging ecology is needed to plan for their long‐term conservation across the circumpolar north, and video collars can provide a powerful approach across remote regions.     

Impacts of spruce budworm defoliation on the habitat of woodland caribou,moose, and their main predators

Forest logging has contributed to the decline of several woodland caribou populations by causing the fragmentation of mature coniferous stands. Such habitat alterations could be worsened by spruce budworm (SBW) outbreaks. Using 6201 vegetation plots from provincial inventories conducted after the last SBW outbreak (1968–1992) in boreal forests of Québec (Canada), we investigated the influence of SBW‐caused tree defoliation and mortality on understory vegetation layers relevant to woodland caribou and its main predators. We found a positive association between severe outbreaks and the cover of most groups of understory plant species, especially in stands that were dominated by balsam fir before the outbreak, where a high canopy openness particularly benefited relatively fast‐growing deciduous plants. Such increases in early successional vegetation could provide high‐quality forage for moose, which is likely to promote higher wolf densities and increase predation pressure on caribou. SBW outbreaks may thus negatively affect woodland caribou by increasing predation risk, the main factor limiting caribou populations in managed forests. For the near future, we recommend updating the criteria used to define critical caribou habitat to consider the potential impacts of spruce budworm defoliation.     

Animal-defined resources reveal nutritional inadequacies for woodland caribou during summer–autumn

Populations of woodland caribou (Rangifer tarandus caribou) are declining throughout their range and many are at risk of extirpation, yet the role of nutrition in these declines remains poorly understood, in part owing to a lack of information about available nutritional resources during summer. We quantified rates of intake of digestible protein and digestible energy by tame caribou foraging in temporary enclosures in the predominant plant communities of northeastern British Columbia, Canada, during summer–autumn and compared intake rates to daily requirements for protein and energy during lactation. We tested hypotheses related to the nutritional adequacy of the environment to support nutritional requirements during lactation (with and without replenishment of body reserves) and simulated scenarios of foraging by caribou in these plant communities to better understand how wild caribou could meet nutritional demands on these landscapes. Nutritional resources varied among plant communities across seasonal, ecological, and successional gradients; digestible energy intake per minute and per day were significantly greater in younger than older forests; dietary digestible energy and per-minute and daily intake of digestible protein were greater, though not significantly so, in younger than older forests; and dietary digestible protein was greater in older than younger forests, though differences were not significant. Tame caribou were unable to satisfy protein and energy requirements during lactation, even without replenishment of body reserves, at most sites sampled. Further, foraging simulations suggested widespread nutritional inadequacies on ranges of wild caribou. Selection for habitats offering the best nutrition may mitigate some nutritional inadequacies, but given low availability of vegetation communities with high nutritional value, performance (e.g., calf production, growth, replenishment of body fat and protein) of caribou may be depressed at levels of nutrition documented herein. Our results, coupled with recent measurements of body fat of wild caribou in northeastern British Columbia, refute the hypothesis that the nutritional environment available to caribou during summer in northeastern British Columbia is adequate to fully support nutritional demands of lactating caribou, which has implications to productivity of caribou populations, recovery, and conservation.     

Behavioral “bycatch” from camera trap surveys yields insights on prey responses to human‐mediated predation risk

Human disturbance directly affects animal populations and communities, but indirect effects of disturbance on species behaviors are less well understood. For instance, disturbance may alter predator activity and cause knock‐on effects to predator‐sensitive foraging in prey. Camera traps provide an emerging opportunity to investigate such disturbance‐mediated impacts to animal behaviors across multiple scales. We used camera trap data to test predictions about predator‐sensitive behavior in three ungulate species (caribou Rangifer tarandus; white‐tailed deer, Odocoileus virginianus; moose, Alces alces) across two western boreal forest landscapes varying in disturbance. We quantified behavior as the number of camera trap photos per detection event and tested its relationship to inferred human‐mediated predation risk between a landscape with greater industrial disturbance and predator activity and a “control” landscape with lower human and predator activity. We also assessed the finer‐scale influence on behavior of variation in predation risk (relative to habitat variation) across camera sites within the more disturbed landscape. We predicted that animals in areas with greater predation risk (e.g., more wolf activity, less cover) would travel faster past cameras and generate fewer photos per detection event, while animals in areas with less predation risk would linger (rest, forage, investigate), generating more photos per event. Our predictions were supported at the landscape‐level, as caribou and moose had more photos per event in the control landscape where disturbance‐mediated predation risk was lower. At a finer‐scale within the disturbed landscape, no prey species showed a significant behavioral response to wolf activity, but the number of photos per event decreased for white‐tailed deer with increasing line of sight (m) along seismic lines (i.e., decreasing visual cover), consistent with a predator‐sensitive response. The presence of juveniles was associated with shorter behavioral events for caribou and moose, suggesting greater predator sensitivity for females with calves. Only moose demonstrated a positive behavioral association (i.e., longer events) with vegetation productivity (16‐day NDVI), suggesting that for other species bottom‐up influences of forage availability were generally weaker than top‐down influences from predation risk. Behavioral insights can be gleaned from camera trap surveys and provide complementary information about animal responses to predation risk, and thus about the indirect impacts of human disturbances on predator–prey interactions.     

Inferring parturition and neonate survival from movement patterns of female ungulates: a case study using woodland caribou

Analyses of animal movement data have primarily focused on understanding patterns of space use and the behavioural processes driving them. Here, we analyzed animal movement data to infer components of individual fitness, specifically parturition and neonate survival. We predicted that parturition and neonate loss events could be identified by sudden and marked changes in female movement patterns. Using GPS radio‐telemetry data from female woodland caribou (Rangifer tarandus caribou), we developed and tested two novel movement‐based methods for inferring parturition and neonate survival. The first method estimated movement thresholds indicative of parturition and neonate loss from population‐level data then applied these thresholds in a moving‐window analysis on individual time‐series data. The second method used an individual‐based approach that discriminated among three a priori models representing the movement patterns of non‐parturient females, females with surviving offspring, and females losing offspring. The models assumed that step lengths (the distance between successive GPS locations) were exponentially distributed and that abrupt changes in the scale parameter of the exponential distribution were indicative of parturition and offspring loss. Both methods predicted parturition with near certainty (>97% accuracy) and produced appropriate predictions of parturition dates. Prediction of neonate survival was affected by data quality for both methods; however, when using high quality data (i.e., with few missing GPS locations), the individual‐based method performed better, predicting neonate survival status with an accuracy rate of 87%. Understanding ungulate population dynamics often requires estimates of parturition and neonate survival rates. With GPS radio‐collars increasingly being used in research and management of ungulates, our movement‐based methods represent a viable approach for estimating rates of both parameters.