Woodland Caribou Extirpation and Anthropogenic Landscape Disturbance in Ontario

Abstract: The decline of woodland caribou (Rangifer tarandus caribou) has been attributed to anthropogenic landscape disturbances, but critical distance thresholds and time lags between disturbance and extirpation are unknown. Using a database of caribou presence and extirpation for northern Ontario, Canada, geo-coded to 10 times 10-km cells, we constructed logistic regression models to predict caribou extirpation based on distance to the nearest of each of 9 disturbance types: forest cutovers, fires, roads, utility corridors, mines, pits and quarries, lakes, trails, and rail lines. We used Akaike's Information Criterion to select parsimonious models and Receiver-Operating Characteristic curves to derive optimal thresholds. To deal with the effects of spatial autocorrelation on estimates of model significance, we used subsampling and restricted randomizations. Forest cutovers were the best predictor of caribou occupancy, with a tolerance threshold of 13 km to nearest cutover and a time lag of 2 decades between disturbance by cutting and caribou extirpation. Management of woodland caribou should incorporate buffers around habitat and requires long-term monitoring of range occupancy.     

Boreal Caribou Can Coexist with Natural but Not Industrial Disturbances

For species at risk, it is important that demographic models be consistent with our most recent knowledge because alternate model versions can have differing predictions for wildlife and natural resource management. To establish and maintain this consistency, we can compare predicted model values to current or past observations and demographic knowledge. When novel predictor information becomes available, testing for consistency between modeled and observed values ensures the best models are used for robust, evidence-based, wildlife management. We combine novel information on the extent of historical disturbance regimes (industrial and fire) to an existing demographic model and predict historical and projected demographics of woodland caribou (Rangifer tarandus caribou). Exploring 6 simulation experiments across 5 populations in Alberta, Canada, we identify the relative importance of industrial disturbance, fire, and population density to observed population size and growth rate. We confirm the onset of significant declines across all 5 populations began approximately 30 years ago, demonstrate these declines have been consistent, and conclude they are more likely due to industrial disturbance from the oil and gas sector within contemporary population ranges than historical fire regimes. These findings reinforce recent research on the cause of woodland caribou declines. Testing for consistency between observations and models prescribed for species recovery is paramount for assessing the cause of declines, projecting population trends, and refining recovery strategies for effective wildlife management. We provide a novel simulation method for conducting these tests. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Anthropogenic Disturbance and Population Viability of Woodland Caribou in Ontario

One of the most challenging tasks in wildlife conservation and management is to clarify how spatial variation in land cover due to anthropogenic disturbance influences wildlife demography and long-term viability. To evaluate this, we compared rates of survival and population growth by woodland caribou (Rangifer tarandus caribou) from 2 study sites in northern Ontario, Canada that differed in the degree of anthropogenic disturbance because of commercial logging and road development, resulting in differences in predation risk due to gray wolves (Canis lupus). We used an individual-based model for population viability analysis (PVA) that incorporated adaptive patterns of caribou movement in relation to predation risk and food availability to predict stochastic variation in rates of caribou survival. Field estimates of annual survival rates for adult female caribou in the unlogged ( 0.90) and logged ( 0.76) study sites recorded during 2010–2014 did not differ significantly (P > 0.05) from values predicted by the individual-based PVA model (unlogged: = 0.87; logged: 0.79). Outcomes from the individual-based PVA model and a simpler stage-structured matrix model suggest that substantial differences in adult survival largely due to wolf predation are likely to lead to long-term decline of woodland caribou in the commercially logged landscape, whereas the unlogged landscape should be considerably more capable of sustaining caribou. Estimates of population growth rates (λ) for the 2010–2014 period differed little between the matrix model and the individual-based PVA model for the unlogged (matrix model = 1.01; individual-based model = 0.98) and logged landscape (matrix model = 0.88; individual-based model = 0.89). We applied the spatially explicit PVA model to assess the viability of woodland caribou across 14 woodland caribou ranges in Ontario. Outcomes of these simulations suggest that woodland caribou ranges that have experienced significant levels of commercial forestry activities in the past had annual growth rates <0.89, whereas caribou ranges that had not experienced commercial forestry operations had population growth rates >0.96. These differences were strongly related to regional variation in wolf densities. Our results suggest that increased wolf predation risk due to anthropogenic disturbance is of sufficient magnitude to cause appreciable risk of population decline in woodland caribou in Ontario. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Estimating Size and Trend of the North Interlake Woodland Caribou Population Using Fecal-DNA and Capture-Recapture Models

A critical step in recovery efforts for endangered and threatened species is the monitoring of population demographic parameters. As part of these efforts, we evaluated the use of fecal-DNA based capture–recapture methods to estimate population sizes and population rate of change for the North Interlake woodland caribou herd (Rangifer tarandus caribou), Manitoba, Canada. This herd is part of the boreal population of woodland caribou, listed as threatened under the federal Species at Risk Act (2003) and the provincial Manitoba Endangered Species Act (2006). Between 2004 and 2009 (9 surveys), we collected 1,080 fecal samples and identified 180 unique genotypes (102 females and 78 males). We used a robust design survey plan with 2 surveys in most years and analysed the data with Program MARK to estimate encounter rates (p), apparent survival rates (φ), rates of population change (λ), and population sizes (N). We estimated these demographic parameters for males and females and for 2 genetic clusters within the North Interlake. The population size estimates were larger for the Lower than the Upper North Interlake area and the proportion of males was lower in the Lower (33%) than the Upper North Interlake (49%). Population rate of change for the entire North Interlake area (2005–2009) using the robust design Pradel model was significantly <1.0 (λ = 0.90, 95% CI: 0.82–0.99) and varied between sex and area with the highest being for males in Lower North Interlake (λ = 0.98, 95% CI: 0.83–1.13) and the lowest being for females in Upper North Interlake (λ = 0.83, 95% CI: 0.69–0.97). The additivity of λ between sex and area is supported on the log scale and translates into males having a λ that is 0.09 greater than females and independent of sex, Lower North Interlake having a λ that is 0.06 greater than Upper North Interlake. Population estimates paralleled these declining trends, which correspond to trends observed in other fragmented populations of woodland caribou along the southern part of their range. The results of this study clearly demonstrate the applicability and success of non-invasive genetic sampling in monitoring populations of woodland caribou. © 2012 The Wildlife Society.     

Determining Sustainable Levels of Cumulative Effects for Boreal Caribou

Abstract: Direct and indirect effects of industrial development have contributed, in part, to the threatened status of boreal ecotype caribou (Rangifer tarandus caribou) in Alberta and Canada. Our goal was to develop a model that would allow managers to identify landscape-scale targets for industrial development, while ensuring functional habitat for sustainable caribou populations. We examined the relationship between functional habitat loss resulting from cumulative effects of natural and anthropogenic disturbance, and the rate of population change (Λ) for 6 populations of boreal caribou in Alberta, Canada. We defined functional habitat loss according to 2 variables for which we had a priori reasons to suspect causative associations with Λ: 1) percentage area of caribou range within 250 m of anthropogenic footprint, and 2) percentage of caribou range disturbed by wildfire within the last 50 years. Multiple regression coefficients for both independent variables indicated significant effects on Λ. The 2-predictor model explained 96% (R²) of observed variation in Λ among population units (F_2,3 = 35.2, P = 0.008). The model may be used to evaluate plans for industrial development in relation to predicted wildfire rates and goals for caribou population growth rates.     

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.     

Coyote Habitat Selection and Management Implications for the Gaspésie Caribou

ABSTRACT Anthropogenic disturbances can promote establishment and growth of predator populations in areas where secondary prey can then become threatened. In this study, we investigated habitat selection of eastern coyotes (Canis latrans), a relatively new predator in the vicinity of an endangered population of caribou (Rangifer tarandus caribou). We hypothesized that coyotes in the boreal forest depend mainly on disturbed habitat, particularly that of anthropogenic origin, because these habitats provide increased food accessibility. Coyotes would likely take advantage of moose (Alces alces) carcasses, berries, and snowshoe hares (Lepus americanus) found in open habitats created by logging. To test these predictions, we described coyote diet and habitat selection at different spatial and temporal levels and then compared resource availability between habitats. To do so, we installed Global Positioning System radiocollars on 23 individual coyotes in the Gaspésie Peninsula, eastern Québec, Canada. Coyotes selected clear-cuts of 5–20 years and avoided mature coniferous forests both at the landscape and home-range levels. Clear-cuts of 5–20 years were found to contain a high availability of moose carcasses and berries, and vulnerability of snowshoe hares is known to increase in clear-cuts. The importance of these 3 food resources was confirmed by the characteristics of core areas used by coyotes and diet analysis. Moose remains were found at 45% of core areas and coyote diet comprised 51% moose on an annual basis. Anthropogenic disturbances in the boreal forest thus seem to benefit coyotes. Our results indicated that the relationship between coyotes and caribou likely involves spillover predation. This knowledge allows managers to consider spillover predation by coyotes as a possible threat for endangered caribou population when the predator depends mainly on habitat of anthropogenic origin and to suggest methods to alleviate it when developing management plans.     

Modeling the Western Arctic Caribou Herd During a Positive Growth Phase: Potential Effects of Wolves and Radiocollars

ABSTRACT Population modeling exercises can lead to both expected and unexpected results useful for wildlife research and management, even though inferences must often be qualitative, given underlying assumptions. Our main objective was to use empirical data on wolf (Canis lupus) kill rates and growth of the Western Arctic caribou (Rangifer tarandus) herd (WAH) of Alaska, USA, to assess the potential for predator regulation. We used available data and published literature to construct a deterministic density-dependent population model fitted to trends of the WAH from 1976 through 2003. By increasing wolf densities in the baseline model, we failed to reject the hypothesis that wolves at a density of 6.5 wolves per 1,000 km² could regulate a caribou herd at a density of 0.4 caribou per km². In addition, our model may be conservative by underestimating the regulatory potential of wolves. We suggest that this relatively simple predator-preysystem shows signs of a predation—food 2-state model. Elasticities from matrix models may be deceiving. Although herd growth is most sensitive to changes in adult female survival, survival of younger cohorts may be more easily influenced by natural conditions or management action. Management of the WAH near maximum sustained yield may not be attainable if desired, but modeling exercises such as this elucidate options. In conducting this research, we also discovered by Monte Carlo simulation that survival and productivity data from radiocollared females and calves were negatively biased and failed to predict herd growth. Thus, researchers should consider potential effects of neck collars on vital rates of female tundra caribou and concomitant offspring when using sample data to model population dynamics or test hypotheses.     

Caribou Use of Habitat Near Energy Development in Arctic Alaska

Increasing demands for energy have generated interest in expanding oil and gas production on the North Slope of Alaska, USA, raising questions about the resilience of barren-ground caribou (Rangifer tarandus) populations to new development. Although the amount of habitat lost directly to energy development in the Arctic will likely be relatively small, there are significant concerns about habitat that may be indirectly affected because of caribou avoidance behaviors. Behavioral responses to energy development for wildlife have been documented, but such responses are often assumed to dissipate over time, despite scant information on the ability of animals to habituate. To understand the long-term effects of energy development on barren-ground caribou, we investigated the behavior of the Central Arctic Herd in northern Alaska, which has been exposed to oil development on its summer range for approximately 40 years. Using recent (2015–2017) location data from global positioning system (GPS)-collared females, we conducted a zone of influence analysis to assess whether caribou reduced their use of habitat near energy development, and if so, the distance the effects attenuated. We conducted this analysis for the calving, post-calving, and mosquito harassment periods when caribou exhibit distinct resource selection patterns, and contrasted our results to past research that investigated the responses of the Central Arctic Herd immediately following the construction of the oil fields. Despite the long-term presence of energy development within the Central Arctic Herd summer range, we found that female caribou exhibited avoidance responses to infrastructure during all time periods, although the effects waned across the summer. Caribou reduced their use of habitat within 5 km of development during the calving period, within 2 km during the post-calving period, and within 1 km during the mosquito harassment period; these areas were predicted to overlap 12%, 15%, and 17% of important calving, post-calving, and mosquito period habitat, respectively. During the calving period, the indirect effects we observed were similar to those observed in past research, whereas during the post-calving and mosquito periods, we detected avoidance responses that had not been previously reported. These findings corroborate a growing body of evidence suggesting that habituation to industrial development in caribou in the Arctic is likely to be weak or absent, and emphasizes the value of minimizing the footprint of infrastructure within important seasonal habitat to reduce behavioral effects to barren-ground caribou. © 2019 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

A Burning Question: What are the Implications of Forest Fires for Woodland Caribou?

Canada's federal recovery strategy for boreal woodland caribou (Rangifer tarandus caribou) classifies areas burned by forest fire as disturbed habitat. This assignment of fire as a disturbance has potential economic and social implications across Canada, and influences plans and actions to achieve caribou conservation and recovery. Previous researchers have reported caribou avoid burned habitat, but these studies did not typically consider unburned residual patches within fire perimeters. Additionally, the implications of burned habitat on individual caribou survival is unclear. We examined resource selection by boreal woodland caribou of burns, and unburned residual patches, using global positioning system (GPS) locations for 201 caribou across 6 caribou populations in Alberta, Canada. We also examined if burned habitat affected the survival of adult female caribou. Caribou avoided burns and unburned residual patches. Increased use of burned habitats, however, did not lower the survival of adult caribou. Collectively, these results provide evidence to support current assertions that burns, and the embedded unburned residual patches are not preferred caribou habitat and increase our understanding of the implications of forest fire for caribou vital rates. Our investigation offers important information about the role of forest fire in caribou ecology and enhances the identification of disturbed habitat under recovery strategy guidelines to effectively address caribou population declines. © 2021 The Wildlife Society.     

Estimating ungulate recruitment and growth rates using age ratios

Trends in population growth can be monitored with data for key vital rates without knowledge of abundance. Although adult female survival has the highest elasticity for ungulate population dynamics, the more variable recruitment rates are commonly monitored to track local variation in growth rates. Specifically, recruitment is often measured using late winter young:adult age ratios, though these age ratios are difficult to reliably interpret given the contribution of multiple vital rates to annual ratios. We show that the supplementation of age ratio data with concurrent radio-telemetry monitoring of adult female survival allows both retrospective estimation of empirical population growth rates and the decomposition of recruitment-specific vital rates. We demonstrate the estimation of recruitment and population growth rates for 1 woodland caribou population using these methods, including elasticity and life-stage simulation analysis of the relative contribution of adult female survival and recruitment rates to variation in population growth. We show, for this woodland caribou population, that adult female survival and recruitment rates were nearly equivalent drivers of population growth. We recommend the concurrent monitoring of adult female survival to reliably interpret age ratios when managing caribou and other ungulates. © 2011 The Wildlife Society.     

Body Size Variations in Caribou Ecotypes and Relationships With Demography

ABSTRACT In many vertebrates size is one of the most influential and variable individual characteristics and a strong determinant of reproductive success. Body size is generally density dependent and decreases when intraspecific competition increases. Frequent and long-distance movements increase energy expenditures and, therefore, may also influence body size, particularly in highly mobile species. Caribou (Rangifer tarandus, also known as reindeer) exhibit tremendous variation in size and movements and thus represent an excellent candidate species to test the relationships between body size, population size, and movements. We analyzed body measurements of adult female caribou from 7 herds of the Québec-Labrador Peninsula, Canada, and we related their morphology to population size, movements, and annual ranges. The herds represented 3 ecotypes (migratory, montane, and sedentary). Ecotypes and herds differed in size (length), shape (roundness), and movements. The sedentary ecotype was larger and moved 4 to 7 times less than the migratory ecotype in the 1990s. At the start of a demographic growth period in the early 1960s, migratory caribou from the Rivière-George (hereafter George) herd had longer mandibles than caribou of the sedentary ecotype. Mandible length in the George herd declined in the 1980s after rapid population growth, while individuals performed extensive movements and the herd's annual range increased. Migratory caribou then became shorter than sedentary caribou. After the George herd decline in the 1990s, mandible length increased again near levels of the 1980s. Caribou from the migratory Rivière-aux-Feuilles herd later showed a similar decline in mandible length during a period of population growth, associated with longer movements and increasing annual range. We hypothesize that the density-dependent effect observed on body size might have been exerted through summer habitat degradation and movement variations during herd growth. Our study has 2 important implications for caribou management: the distinctiveness of different populations and ecotypes, and the correlations between population trajectories and changes in body condition and habitat.     

Comparison of Woodland Caribou Calving Areas Determined by Movement Patterns Across Northern Ontario

Adult female survival and calf recruitment influence population dynamics, but there is limited information on calving and neonatal mortality of boreal woodland caribou (Rangifer tarandus caribou; caribou) in Ontario, Canada. We identified calf parturition sites and 5-week neonatal mortality using a movement-based approach across 3 northern Ontario study regions (Pickle Lake, Nakina, and Cochrane) that vary in their capacity to support caribou populations. In comparing 22 caribou-years of video-collar footage during 2010–2013 to predictions of the movement-based approach, we found live parturition events were 100% correctly classified, date of parturition was within 1.08 ± 0.28 ( ± SE) days, and mortality events up to 5 weeks postpartum were 88% correctly classified. Across study regions, 87% of 186 caribou were pregnant and 76% of 107 caribou-years indicated birth events with median parturition dates a week later in Cochrane (23 May) than in Pickle Lake (17 May) and Nakina (16 May). Based on selection ratios of caribou-years with calves-at-heel (n = 80), caribou consistently selected for lowlands and closed-canopied forests and mostly against early-seral stands (<20 yrs old) and areas near linear features during the neonatal and the post-neonatal period (up to 35 days postpartum). Based on the video footage and movement models, 30% of 81 caribou-years that indicated live births also showed females lost their calf within the first 5 weeks postpartum, with higher risk of neonatal mortality associated with increased use of lowlands and greater postpartum movement rates. This study provides informative metrics of caribou reproduction across northern Ontario that will contribute to future population modeling and identifies important landscape features to be considered in future industrial development and land use planning for caribou conservation. © 2020 The Wildlife Society.     

Divergent estimates of herd‐wide caribou calf survival: Ecological factors and methodological biases

Population monitoring is a critical part of effective wildlife management, but methods are prone to biases that can hinder our ability to accurately track changes in populations through time. Calf survival plays an important role in ungulate population dynamics and can be monitored using telemetry and herd composition surveys. These methods, however, are susceptible to unrepresentative sampling and violations of the assumption of equal detectability, respectively. Here, we capitalized on 55 herd‐wide estimates of woodland caribou (Rangifer tarandus caribou) calf survival in Newfoundland, Canada, using telemetry (n = 1,175 calves) and 249 herd‐wide estimates of calf:cow ratios (C:C) using herd composition surveys to investigate these potential biases. These data included 17 herd‐wide estimates replicated from both methods concurrently (n = 448 calves and n = 17 surveys) which we used to understand which processes and sampling biases contributed to disagreement between estimates of herd‐wide calf survival. We used Cox proportional hazards models to determine whether estimates of calf mortality risk were biased by the date a calf was collared. We also used linear mixed‐effects models to determine whether estimates of C:C ratios were biased by survey date and herd size. We found that calves collared later in the calving season had a higher mortality risk and that C:C tended to be higher for surveys conducted later in the autumn. When we used these relationships to modify estimates of herd‐wide calf survival derived from telemetry and herd composition surveys concurrently, we found that formerly disparate estimates of woodland caribou calf survival now overlapped (within a 95% confidence interval) in a majority of cases. Our case study highlights the potential of under‐appreciated biases to impact our understanding of population dynamics and suggests ways that managers can limit the influence of these biases in the two widely applied methods for estimating herd‐wide survival.     

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.     

Caribou movement as a correlated random walk

Movement is a primary mechanism coupling animals to their environment, yet there exists little empirical analysis to test our theoretical knowledge of this basic process. We used correlated random walk (CRW) models and satellite telemetry to investigate long-distance movements of caribou, the most vagile, non-volant terrestrial vertebrate in the world. Individual paths of migratory and sedentary female caribou were quantified using measures of mean move length and angle, and net squared displacements at each successive move were compared to predictions from the models. Movements were modelled at two temporal scales. For paths recorded through one annual cycle, the CRW model overpredicted net displacement of caribou through time. For paths recorded over shorter intervals delineated by seasonal behavioural changes of caribou, there was excellent correspondence between model predictions and observations for most periods for both migratory and sedentary caribou. On the smallest temporal scale, a CRW model significantly overpredicted displacements of migratory caribou during 3 months following calving; this was also the case for sedentary caribou in late summer, and in late winter. In all cases of overprediction there was significant positive autocorrelation in turn direction, indicating that movements were more tortuous than expected. In one case of underprediction, significant negative autocorrelation of sequential turn direction was evident, indicating that migratory caribou moved in straightened paths during spring migration to calving grounds. Results are discussed in light of known migration patterns and possible limiting factors for caribou, and indicate the applicability of CRW models to animal movement at vast spatial and temporal scales, thus assisting in future development of more sophisticated models of population spread and redistribution for vertebrates. Received: 14 July 1999 / Accepted: 15 November 1999     

Effects of weather and hunting on wild reindeer population dynamics in Hardangervidda National Park

Wild reindeer have a range that extends across the circumpolar region. In the last few decades, however, populations of wild reindeer have been on the decline. The reasons for these declines are poorly understood, but are suggested to be linked to both local and global climatic factors, disease, and human interference. Hardangervidda plateau in Norway is home to the largest wild reindeer population in Europe, and is at the southern end of its European range. This population is therefore of particular importance, particularly in the light of climate change. We investigated how weather and hunting have affected the wild reindeer population in Hardangervidda over the last two decades. Our findings suggest that the wild reindeer population in Hardangervidda is most affected by winter temperature and hunting, where colder temperatures and lower harvest rates typically result in higher growth rates. We did not find significant evidence for linear density dependence. Our results show trends across Hardangervidda, and give an indication of how region-wide weather and hunting pressure can affect the wild reindeer population. As new data emerge, future investigations should look into the existence and nature of density dependence and the influence of other weather and human disturbance related factors.