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.     

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.     

Changing northern vegetation conditions are influencing barren ground caribou (Rangifer tarandus groenlandicus) post‐calving movement rates

Aim

To quantify changes in vegetation productivity over the past three decades across five barren ground caribou (Rangifer tarandus groenlandicus) herd ranges and assess how these changes are influencing caribou movement rates.

Location

Northwest Territories and Nunavut, Canada.

Methods

As an indicator of vegetation productivity, the enhanced vegetation index (EVI) was calculated on newly developed cloud‐free, gap‐free, Landsat surface reflectance image composites representing 1984–2012. Changes in EVI were assessed on a pixel basis using Theil‐Sen's nonparametric regression and compared across herd ranges and land cover types using generalized least squares regression. Animal movement velocity was calculated from caribou telemetry data and generalized additive mixed models were used to link movement rates with vegetation productivity during the post‐calving phase of the year (July and August).

Results

Vegetation productivity increased across the five caribou herd ranges examined. The largest productivity increase occurred over the ranges of the most western herds, with the largest observed changes in grassland or shrub habitats. Caribou tended to move more slowly through tundra habitats with elevated levels of productivity to a point, while grasslands movement rates decreased linearly with increasing productivity. Movement velocities peaked at intermediate productivity levels in shrub habitats.

Main conclusions

Over the three decades of collected data, barren ground caribou habitats have become more productive, which is consistent with other studies that have documented increases in Arctic vegetation productivity. The more western herds, whose ranges are also closest to the Arctic Ocean, experienced the largest increases in productivity. Finally, we demonstrate that barren ground caribou movement patterns will likely change as a result of changing vegetation productivity in complex manners depending on herd, habitat type and the magnitude of change in vegetation productivity.     

Modeling Caribou Movements: Seasonal Ranges and Migration Routes of the Central Arctic Herd

Migration is an important component of the life history of many animals, but persistence of large-scale terrestrial migrations is being challenged by environmental changes that fragment habitats and create obstacles to animal movements. In northern Alaska, the Central Arctic herd (CAH) of barren-ground caribou (Rangifer tarandus granti) is known to migrate over large distances, but the herd’s seasonal distributions and migratory movements are not well documented. From 2003–2007, we used GPS radio-collars to determine seasonal ranges and migration routes of 54 female caribou from the CAH. We calculated Brownian bridges to model fall and spring migrations for each year and used the mean of these over all 4 years to identify areas that were used repeatedly. Annual estimates of sizes of seasonal ranges determined by 90% fixed kernel utilization distributions were similar between summer and winter ($\bar{X}$ = 27,929 SE = 1,064 and $\bar{X}$ = 26,585 SE = 4912 km², respectively). Overlap between consecutive summer and winter ranges varied from 3.3–18.3%. Percent overlap between summer ranges used during consecutive years ($\bar{X}$ = 62.4% SE = 3.7%) was higher than for winter ranges ($\bar{X}$ = 42.8% SE = 5.9%). Caribou used multiple migration routes each year, but some areas were used by caribou during all years, suggesting that these areas should be managed to allow for continued utilization by caribou. Restoring migration routes after they have been disturbed or fragmented is challenging. However, prior knowledge of movements and threats may facilitate maintenance of migratory paths and seasonal ranges necessary for long-term persistence of migratory species.     

Modeling Sustainability of Arctic Communities: An Interdisciplinary Collaboration of Researchers and Local Knowledge Holders

How will climate change affect the sustainability of Arctic villages over the next 40 years? This question motivated a collaboration of 23 researchers and four Arctic communities (Old Crow, Yukon Territory, Canada; Aklavik, Northwest Territories, Canada; Fort McPherson, Northwest Territories, Canada; and Arctic Village, Alaska, USA) in or near the range of the Porcupine Caribou Herd. We drew on existing research and local knowledge to examine potential effects of climate change, petroleum development, tourism, and government spending cutbacks on the sustainability of four Arctic villages. We used data across eight disciplines to develop an Arctic Community Synthesis Model and a Web-based, interactive Possible Futures Model. Results suggested that climate warming will increase vegetation biomass within the herd’s summer range. However, despite forage increasing, the herd was projected as likely to decline with a warming climate because of increased insect harassment in the summer and potentially greater winter snow depths. There was a strong negative correlation between hypothetical, development-induced displacement of cows and calves from utilized calving grounds and calf survival during June. The results suggested that climate warming coupled with petroleum development would cause a decline in caribou harvest by local communities. Because the Synthesis Model inherits uncertainties associated with each component model, sensitivity analysis is required. Scientists and stakeholders agreed that (1) although simulation models are incomplete abstractions of the real world, they helped bring scientific and community knowledge together, and (2) relationships established across disciplines and between scientists and communities were a valuable outcome of the study. Additional project materials, including the Web-based Possible Futures Model, are available at http://www.taiga.net/sustain.     

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.     

Behaviour of West Greenland caribou during a population decline

The major decline of the West Greenland caribou herd during the 1970s prompted a study in 1977–78 of caribou behaviour in relation to environmental factors associated with the decline. Quantification of caribou activity revealed behavioural responses to critically low standing crops of preferred winter forage. Caribou on poor winter range were mostly inactive with low feeding intensities and abnormal diurnal activity patterns. In addition, these animals exhibited very restricted movements, frequently occurred alone or in small groups, and were unable to maintain normal social bonds. In contrast, where fruticose lichen winter forage was available, caribou did not display these behavioural traits. After the initiation of new vegetative growth, feeding intensity increased and social tendency returned. Comparison with behavioural data from other Rangifer populations suggests that the observed responses to low forage quality are not restricted to Greenland but are a normal response of barren-ground caribou to winter ranges poor in lichen forage.     

Interchange and Overlap Among Four Adjacent Arctic Caribou Herds

Barren ground caribou (Rangifer tarandus granti) are distributed in herds that seasonally use specific geographic regions within an annual range, with varying levels of fidelity during different periods (e.g., calving, insect relief, wintering). As a result, caribou management is generally tailored to individual herds that often range across administrative boundaries. Herd ranges can shift over time, seasonal ranges of adjacent herds often overlap, herds merge, and there is often little genetic differentiation among adjacent herds. If substantial herd interchange occurs, it would have important management implications by influencing estimates of herd size, herd composition, and harvest rates. We compiled satellite telemetry data from 2003–2015 for 4 large arctic caribou herds to quantify herd interchange rates. We calculated a metric of herd interchange based on the relationship of caribou locations to typical weekly herd ranges (all yrs combined) and the distance to other radio-collared caribou from each of the 4 herds (yr specific). Although herd membership cannot always be clearly defined based on location, this metric provides an objective measure of the strength of evidence of herd membership that can be used to make comparisons among herds and time periods. We also calculated herd overlap and quantified how it varied throughout the year. Herd interchange was rare in the 2 larger herds, generally occurring when caribou overwintered with an adjacent herd, whereas herd interchange from the 2 smaller herds was more frequent and could last longer than a year. Although sample sizes were limited, there were no clear patterns in herd interchange with year or annual herd size. The 2 smaller herds had large seasonal overlap with adjacent herds, suggesting that herd interchange may be related to spatiotemporal herd overlap and relative herd size. Our results can help managers understand herd interchange and overlap to make management decisions, interpret research results, and develop more accurate population models. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Behavioural trade-offs in response to external stimuli: time allocation of an Arctic ungulate during varying intensities of harassment by parasitic flies

1. Macroparasites may be a major factor shaping animal behaviour. Tundra ecosystems inhabited by caribou and reindeer (Rangifer tarandus) are known for large concentrations of ectoparasites including mosquitoes (Culicidae) and black flies (Simuliidae), as well as endoparasitic oestrid flies (Oestridae). 2. Increased intensity and duration of insect harassment because of climatic warming is hypothesized as a potential factor in recent declines of Rangifer across the circumpolar north. Although there is a well-observed relationship between insect harassment and caribou/reindeer behaviour, the influence of ecto- relative to endoparasitic species is unclear. Climatic changes may favour the activity patterns, distribution or abundance of certain insect species; thus, understanding differential effects on the behaviour of Rangifer is important. 3. We recorded caribou behaviour using group scan and focal sampling methods, while simultaneously trapping insects and recording weather conditions on the postcalving/summer range of the Bathurst barren-ground caribou herd in Northwest Territories and Nunavut, Canada, during 2007-2009. 4. We developed statistical model sets representing hypotheses about the effects of insects, weather, habitat/location, and date/time on caribou behaviour. We used multinomial logistic regression models to explore factors affecting the relative dominance of behaviour types within groups of caribou and fractional multinomial logistic regression models to determine factors influencing time allocation by individual caribou. We examined changes in feeding intensity using fractional logistic regression. 5. Relative dominance of insect avoidance behaviour within caribou groups and time allocation to insect avoidance by individual caribou increased when oestrid flies were present or black flies were active at moderate-high levels. Mosquito activity had relatively little effect on caribou behaviour. Time spent feeding was reduced by the greatest degree when all three insect types were present in combination. Feeding intensity was influenced to a greater extent by the accumulation of growing degree days over the course of the postcalving/summer season than by insect activity. Changes in Arctic systems that increase the activity/abundance of ecto- and endoparasites could have implications for the productivity of Rangifer populations.     

Linkages between large‐scale climate patterns and the dynamics of Arctic caribou populations

Recent research has linked climate warming to global declines in caribou and reindeer (both Rangifer tarandus) populations. We hypothesize large‐scale climate patterns are a contributing factor explaining why these declines are not universal. To test our hypothesis for such relationships among Alaska caribou herds, we calculated the population growth rate and percent change of four arctic herds using existing population estimates, and explored associations with indices of the Arctic Oscillation (AO) and the Pacific Decadal Oscillation (PDO). The AO, which more strongly affects eastern Alaska, was negatively associated with the population trends of the Porcupine Caribou Herd and Central Arctic Herd, the easternmost of the herds. We hypothesize that either increased snowfall or suboptimal growing conditions for summer forage plants could explain this negative relationship. Intensity of the PDO, which has greatest effects in western Alaska, was negatively associated with the growth rate of the Teshekpuk Caribou Herd in northwestern Alaska, but the Western Arctic Herd in western Alaska displayed the opposite trend. We suggest that the contrasting patterns of association relate to the spatial variability of the effects of the PDO on western and northwestern Alaska. Although predation and winter range quality have often been considered the primary causes of population variation, our results show that large‐scale climate patterns may play an important role in caribou population dynamics in arctic Alaska. Our findings reveal that climate warming has not acted uniformly to reduce caribou populations globally. Further research should focus on the relative importance of mechanisms by which climate indices influence caribou population dynamics.     

Summer Resource Selection and Identification of Important Habitat Prior to Industrial Development for the Teshekpuk Caribou Herd in Northern Alaska

Many caribou (Rangifer tarandus) populations are declining worldwide in part due to disturbance from human development. Prior to human development, important areas of habitat should be identified to help managers minimize adverse effects. Resource selection functions can help identify these areas by providing a link between space use and landscape attributes. We estimated resource selection during five summer periods at two spatial scales for the Teshekpuk Caribou Herd in northern Alaska prior to industrial development to identify areas of high predicted use for the herd. Additionally, given the strong influence parturition and insect harassment have on space use, we determined how selection differed between parturient and non-parturient females, and between periods with and without insect harassment. We used location data acquired between 2004–2010 for 41 female caribou to estimate resource selection functions. Patterns of selection varied through summer but caribou consistently avoided patches of flooded vegetation and selected areas with a high density of sedge-grass meadow. Predicted use by parturient females during calving was almost entirely restricted to the area surrounding Teshekpuk Lake presumably due to high concentration of sedge-grass meadows, whereas selection for this area by non-parturient females was less strong. When insect harassment was low, caribou primarily selected the areas around Teshekpuk Lake but when it was high, caribou used areas having climates where insect abundance would be lower (i.e., coastal margins, gravel bars). Areas with a high probability of use were predominately restricted to the area surrounding Teshekpuk Lake except during late summer when high use areas were less aggregated because of more general patterns of resource selection. Planning is currently underway for establishing where oil and gas development can occur in the herd’s range, so our results provide land managers with information that can help predict and minimize impacts of development on the herd.     

End of the road: Short-term responses of a large mammal community to forest road decommissioning

Anthropogenic disturbances are increasing worldwide, causing wildlife habitat loss, alteration, and fragmentation. In Canada, the decommissioning of linear anthropogenic structures is identified as a promising tool to restore the habitat of threatened populations of boreal caribou (Rangifer tarandus caribou) by reducing food availability for alternate prey and decreasing encounter probabilities with predators. In this study, we monitored the use of 40 km of decommissioned forest roads by caribou, gray wolves (Canis lupus), black bears (Ursus americanus), and moose (Alces americanus) 1–3 years after reclamation, using 232 motion-activated camera traps. We compared four additive treatments (meaning that each successive treatment included the treatment prior): closing the road to human access, decompacting its soil, planting black spruce (Picea mariana) trees, and adding enriched soil. We assessed the influence of treatments, use by other large mammals, and characteristics of the surrounding environment on road use by the four species. Caribou used the planted treatment (which also included closing and decompacting) more than the closed-only (reference) treatment, but treatments did not influence the use of decommissioned roads by bears and moose. We could not assess the use of treated roads by wolves because of low sample size. Road use by caribou declined with local moose density, but increased with local bear density. Caribou were observed more frequently on roads surrounded by regenerating and mature coniferous stands; caribou also preferentially used roads surrounded by wetlands. Our results suggest that the treatment combining road closure, soil decompaction, and tree planting could be beneficial to caribou, highlighting the relevance of including active restoration efforts in caribou conservation programs. We recommend that such a treatment be added to road decommissioning protocols for the conservation of caribou, alongside broad-scale habitat protection.     

Reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>) avoidance of a highway as revealed by lichen measurements

Reindeer and caribou Rangifer tarandus are reported to avoid human infrastructure such as roads, high-voltage power lines, pipelines, and tourist resorts. Lichens are important forage for reindeer during winter, and their relatively slow growth rates make them vulnerable to overgrazing. Height and volume of lichens are often used as an indicator of grazing pressure by reindeer and, thus, as an indirect measure of Rangifer avoidance of human infrastructure. We sampled lichen height in Cetraria nivalis-dominated communities along 4 and 3 parallel transects located on two parallel mountain ridges in Hardangervidda, south central Norway. The lichen measurements were analyzed in relation to altitude and the distance from four tourist cabins in the area and a highway (Rv7) running perpendicular to the 7 transects. The mountain ridge with 4 transects is part of a much used migratory corridor for wild reindeer R. tarandus tarandus. Along the nonmigratory ridge, lichen height decreased 35% over an 8-km distance from Rv7 and a tourist cabin, indicating reindeer aversion toward Rv7 and/or a tourist cabin. No similar relationship was found for the migration ridge in relation to distance from Rv7 or the tourist cabins. Our results suggest that avoidance of human infrastructure by wild reindeer might be limited where reindeer use of winter pastures is influenced by herd traditions and/or motivation to follow established migration corridors. This has important implications for addressing the use of similar pasture measurements when testing for Rangifer aversion toward human disturbances.     

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.     

Cumulative geoecological effects of 62 years of infrastructure and climate change in ice‐rich permafrost landscapes,Prudhoe Bay Oilfield,Alaska

Many areas of the Arctic are simultaneously affected by rapid climate change and rapid industrial development. These areas are likely to increase in number and size as sea ice melts and abundant Arctic natural resources become more accessible. Documenting the changes that have already occurred is essential to inform management approaches to minimize the impacts of future activities. Here, we determine the cumulative geoecological effects of 62 years (1949–2011) of infrastructure‐ and climate‐related changes in the Prudhoe Bay Oilfield, the oldest and most extensive industrial complex in the Arctic, and an area with extensive ice‐rich permafrost that is extraordinarily sensitive to climate change. We demonstrate that thermokarst has recently affected broad areas of the entire region, and that a sudden increase in the area affected began shortly after 1990 corresponding to a rapid rise in regional summer air temperatures and related permafrost temperatures. We also present a conceptual model that describes how infrastructure‐related factors, including road dust and roadside flooding are contributing to more extensive thermokarst in areas adjacent to roads and gravel pads. We mapped the historical infrastructure changes for the Alaska North Slope oilfields for 10 dates from the initial oil discovery in 1968–2011. By 2010, over 34% of the intensively mapped area was affected by oil development. In addition, between 1990 and 2001, coincident with strong atmospheric warming during the 1990s, 19% of the remaining natural landscapes (excluding areas covered by infrastructure, lakes and river floodplains) exhibited expansion of thermokarst features resulting in more abundant small ponds, greater microrelief, more active lakeshore erosion and increased landscape and habitat heterogeneity. This transition to a new geoecological regime will have impacts to wildlife habitat, local residents and industry.     

How landscape dynamics link individual‐ to population‐level movement patterns: a multispecies comparison of ungulate relocation data

Aim To demonstrate how the interrelations of individual movements form large‐scale population‐level movement patterns and how these patterns are associated with the underlying landscape dynamics by comparing ungulate movements across species. Locations Arctic tundra in Alaska and Canada, temperate forests in Massachusetts, Patagonian Steppes in Argentina, Eastern Steppes in Mongolia. Methods We used relocation data from four ungulate species (barren‐ground caribou, Mongolian gazelle, guanaco and moose) to examine individual movements and the interrelation of movements among individuals. We applied and developed a suite of spatial metrics that measure variation in movement among individuals as population dispersion, movement coordination and realized mobility. Taken together, these metrics allowed us to quantify and distinguish among different large‐scale population‐level movement patterns such as migration, range residency and nomadism. We then related the population‐level movement patterns to the underlying landscape vegetation dynamics via long‐term remote sensing measurements of the temporal variability, spatial variability and unpredictability of vegetation productivity. Results Moose, which remained in sedentary home ranges, and guanacos, which were partially migratory, exhibited relatively short annual movements associated with landscapes having very little broad‐scale variability in vegetation. Caribou and gazelle performed extreme long‐distance movements that were associated with broad‐scale variability in vegetation productivity during the peak of the growing season. Caribou exhibited regular seasonal migration in which individuals were clustered for most of the year and exhibited coordinated movements. In contrast, gazelle were nomadic, as individuals were independently distributed and moved in an uncoordinated manner that relates to the comparatively unpredictable (yet broad‐scale) vegetation dynamics of their landscape. Main conclusions We show how broad‐scale landscape unpredictability may lead to nomadism, an understudied type of long‐distance movement. In contrast to classical migration where landscapes may vary at broad scales but in a predictable manner, long‐distance movements of nomadic individuals are uncoordinated and independent from other such individuals. Landscapes with little broad‐scale variability in vegetation productivity feature smaller‐scale movements and allow for range residency. Nomadism requires distinct integrative conservation strategies that facilitate long‐distance movements across the entire landscape and are not limited to certain migration corridors.     

Impacts of climate change on the seasonal distribution of migratory caribou

Arctic ecosystems are especially vulnerable to global climate change as temperature and precipitation regimes are altered. An ecologically and socially highly important northern terrestrial species that may be impacted by climate change is the caribou, Rangifer tarandus . We predicted the current and potential future occurrence of two migratory herds of caribou [Rivière George herd (RG) and Rivière-aux-Feuilles (RAF) herd] under a Canadian General Circulation Model climate change scenario, across all seasons in the Québec–Labrador peninsula, using climatic and habitat predictor variables. Argos satellite-tracking collars have been deployed on 213 caribou between 1988 and 2003 with locations recorded every 4–5 days. In addition, we assembled a database of climate (temperature, precipitation, snowfall, timing and length of growing season) and habitat data obtained from the SPOT VEGETATION satellite sensor. Logistic regression models indicated that both climatic and physical habitat variables were significant predictors of current migratory caribou occurrence. Migratory caribou appeared to prefer regions with higher snowfall and lichen availability in the fall and winter. In the summer, caribou preferred cooler areas likely corresponding to a lower prevalence of insects, and they avoided disturbed and recently burnt areas. Climate change projections using climate data predicted an increased range for the RAF herd and decreased range for the RG herd during 2040–2069, limiting the herds to northeastern regions of the Québec–Labrador peninsula. Direct and indirect consequences of climate change on these migratory caribou herds possibly include alteration in habitat use, migration patterns, foraging behaviour, and demography, in addition to social and economic stress to arctic and subarctic native human populations.     

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     

Disentangling Woodland Caribou Movements in Response to Clearcuts and Roads across Temporal Scales

Although prey species typically respond to the most limiting factors at coarse spatiotemporal scales while addressing biological requirements at finer scales, such behaviour may become challenging for species inhabiting human altered landscapes. We investigated how woodland caribou, a threatened species inhabiting North-American boreal forests, modified their fine-scale movements when confronted with forest management features (i.e. clearcuts and roads). We used GPS telemetry data collected between 2004 and 2010 on 49 female caribou in a managed area in Québec, Canada. Movements were studied using a use – availability design contrasting observed steps (i.e. line connecting two consecutive locations) with random steps (i.e. proxy of immediate habitat availability). Although caribou mostly avoided disturbances, individuals nonetheless modulated their fine-scale response to disturbances on a daily and annual basis, potentially compromising between risk avoidance in periods of higher vulnerability (i.e. calving, early and late winter) during the day and foraging activities in periods of higher energy requirements (i.e. spring, summer and rut) during dusk/dawn and at night. The local context in which females moved was shown to influence their decision to cross clearcut edges and roads. Indeed, although females typically avoided crossing clearcut edges and roads at low densities, crossing rates were found to rapidly increase in greater disturbance densities. In some instance, however, females were less likely to cross edges and roads as densities increased. Females may then be trapped and forced to use disturbed habitats, known to be associated with higher predation risk. We believe that further increases in anthropogenic disturbances could exacerbate such behavioural responses and ultimately lead to population level consequences.     

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.