Caribou Rangifer Tarandus Research Articles

Spring diet of American black bears (Ursus americanus) in a moose (Alces alces) – woodland caribou (Rangifer tarandus caribou) system in northern Ontario, Canada

American black bears (Ursus americanus Pallas, 1780) are opportunistic foragers, and across most of their range in North America, their diet is predominantly vegetation with limited consumption of vertebrates and invertebrates. However, they are also predators of ungulates, especially neonates, with regional variation in the amount of predation pressure they exert. We used scat analysis to examine the diet of black bears during the calving season in a moose (Alces alces (Linnaeus, 1758)) – woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)) system in the Boreal forest of northern Ontario, Canada. Bears consumed herbaceous plants (46.5%), fruits and (or) seeds (20.0%), moose (3.3% adults; 4.3% calves), American beaver (Castor canadensis Kuhl, 1820; 8.5%), and insects (mostly ants; 4.2%). Bears had the highest consumption of moose and beaver in early spring, before switching to a more vegetation-dominated diet. We did not detect evidence of caribou consumption. Based on our results, black bear consumption of moose, particularly neonates, may warrant further investigation.

Red-Listed Ecosystem Status of Interior Wetbelt and Inland Temperate Rainforest of British Columbia, Canada

The Interior Wetbelt (IWB) of British Columbia, which includes the globally rare Inland Temperate Rainforest (ITR), contains primary forests poorly attributed and neglected in conservation planning. We evaluated the IWB and ITR using four IUCN Red List of Ecosystems Criteria: geographic distribution, environmental degradation (abiotic and biotic factors), and likelihood of ecosystem collapse. Clearcut logging (3.2M ha) represented 57% of all anthropogenic disturbances, reducing potential primary forest by 2.7 million ha (28%) for the IWB and 524,003 ha (39%) for the ITR. Decadal logging rates nearly doubled from 5.3% to 10.2% from 1970s–2000s. Core areas (buffered by 100-m from roads and developments) declined by 70% to 95% for the IWB and ITR, respectively. Vulnerable was assigned to karst, the only abiotic factor assessed, because it was associated with rare plants. For biotic factors, Old-Growth Birds were Vulnerable, Southern Woodland Caribou (Rangifer tarandus caribou) habitat and Sensitive Fish were Endangered, and Old-Growth Lichens habitat was Critical. Overall, the IWB was ranked as Endangered and the ITR as Critical with core area collapse possible within 9 to 18 years for the ITR, considered one of the world’s most imperiled temperate rainforests.

Seismic line edge effects on plants, lichens and their environmental conditions in boreal peatlands of Northwest Alberta (Canada)

Boreal peatlands of Alberta (Canada) are heavily dissected by a large network of linear disturbances, including seismic lines used for Oil and Gas exploration. Edge effects resulting from these linear disturbances can increase their footprint and affect wildlife habitat, including the quality and quantity of plant and lichen forage. In 2012, seismic line restoration became a priority in Alberta, under the realm of restoration of habitat of threatened boreal woodland caribou. More accurate understanding of biotic and abiotic interactions leading to edge effects can be important in choosing appropriate restoration practices and prompting natural regeneration. We assessed the extent of edge effects of seismic lines on the adjacent boreal peatland near Peace River, northwestern Alberta. We compared plant and lichen community composition and environmental conditions on seismic lines and along 22 transects, which extended perpendicularly 75 m into the peatland from both lines edges. Soil moisture and light were higher on the lines, but frozen ground was less frequent than in the adjacent peatland. Seismic lines were also subsided by approximately 1.2 cm. Lichens were less abundant in the 2–25 m zone from the edge—indicating edge effects. On the lines,Sphagnumconstituted a nearly 100% cover while lichens, trees, herbs, and other bryophytes were less abundant than in the peatland. We demonstrate how knowledge of edge effects on plants and lichens could be applied in assessing caribou habitat quality based on forage availability and predation risk, and how such knowledge could inform restoration practices of seismic lines in peatlands.

Context-dependent group size: effects of population density, habitat, and season

AbstractGroup size can vary in relation to population density, habitat, and season. Habitat and season may also interact with population density and affect group size through varying foraging benefits of social aggregation in different ecological contexts. We tested the hypothesis that group size varies across ecological contexts, including population density, habitat type, and season, for woodland caribou (Rangifer tarandus) in ten herds over 25 years in Newfoundland, Canada. We predicted that group size would increase as a function of population density. Based on the foraging benefits of social aggregation, we predicted larger groups as habitat openness increased because open areas tend to have higher quality foraging resources. We predicted larger groups during winter when foraging resources are covered in snow because caribou and other social animals exploit social information about the location of foraging resources. In contrast to our prediction, group size decreased as a function of population density. In support of our prediction, group size was larger in winter than calving and summer, and we found that group size increased with habitat openness in some, but not all, cases. Patterns of animal grouping are context-dependent and the additive effect of different ecological contexts on variation in group size informs our understanding of the implicit trade-offs between competition, predation risk, and profitability of forage.

Balancing Large-Scale Wildlife Protection and Forest Management Goals with a Game-Theoretic Approach

When adopted, wildlife protection policies in Canadian forests typically cover large areas and affect multiple economic agents working in these landscapes. Such measures are likely to increase the costs of timber for forestry companies operating in the area, which may hinder their acceptance of the policies unless harvesting remains profitable. We propose a bi-level wildlife protection problem that accounts for the profit-maximizing behavior of forestry companies operating in an area subject to protection. We consider the regulator with a wildlife protection mandate and forestry companies licensed to harvest public forest lands. We depict the relationship between the regulator and forestry companies as a leader-follower Stackelberg game. The leader sets the protected area target for each license area and the followers adjust their strategies to maximize payoffs while meeting the protection target set by the leader. The leader’s objective is to maximize the area-wide protection of spatially contiguous habitat while accounting for the followers’ profit-maximizing behavior. We apply the approach to investigate habitat protection policies for woodland caribou in the Churchill range, Ontario, Canada. We compare the game-theoretic solutions with solutions that do not consider the forest companies’ objectives and also with solutions equalizing the revenue losses among the companies.

Simultaneous monitoring of vegetation dynamics and wildlife activity with camera traps to assess habitat change

AbstractVegetation phenology and productivity drive resource use by wildlife. Vegetation dynamics also reveal patterns of habitat disturbance and recovery. Monitoring these fine‐scale vegetation patterns over large spatiotemporal extents can be difficult, but camera traps (CTs) commonly used to survey wildlife populations also collect data on local habitat conditions. We used CTs (n = 73) from 2016 to 2019 to assess impacts of habitat change in a boreal landscape of northern Canada, where seismic lines for petroleum exploration disturbed wildlife habitat and prompted vegetation restoration efforts. First, we quantified vegetation dynamics from CTs, comparing them to satellite‐based estimates that are typically used to monitor vegetation at broad spatial scales. We then used understory phenology and productivity estimated from CT time‐lapse images to assess vegetation recovery on seismic lines. Finally, we related vegetation dynamics with the habitat use of three wildlife species: sandhill cranes Grus canadensis, woodland caribou Rangifer tarandus, and white‐tailed deer Odocoileus virginianus. CTs provided unique insight into vegetation dynamics that were different from signals measured by satellites, with temporally inconsistent and even some negative correlations between CT and satellite metrics. We found some indication of vegetation recovery on seismic lines that had received restoration treatment, with understory patterns more similar to undisturbed habitat than to seismic lines that did not receive restoration treatment. CTs also provided inferences about wildlife activity related to vegetation resources, which approaches using satellite data failed to detect. Wildlife habitat use tracked vegetation phenology, but did not always increase with vegetation productivity at weekly, 16‐day, or annual intervals. Instead, associations with vegetation productivity depended on species, temporal scale, and productivity metrics. Given the widespread and growing use of CTs to monitor terrestrial wildlife, we recommend their use to simultaneously monitor habitat conditions to better understand the mechanisms that govern wildlife habitat use in changing environments.

Management‐mediated predation rate in the caribou–moose–wolf system: spatial configuration of logging activities matters

AbstractLandscape complexity can determine the population dynamics of interacting predators and prey. Yet, management plans are commonly developed from aspatial predictive models. This oversight may result in unexpected outcomes or the loss of opportunities to make spatial interventions that would increase a plan's effectiveness. The management of the threatened woodland caribou (Rangifer tarandus caribou), boreal population, provides an example of such shortcomings when using an aspatial approach. Currently, the most influential management recommendation is to maintain at least 65% of undisturbed forests in areas occupied by caribou populations, regardless of the spatial configuration of the forest cover. Using a spatially explicit individual‐based model (IBM), we evaluated the effects of the spatial configuration of cuts and roads on the mortality of boreal caribou living in sympatry with wolves (Canis lupus) and moose (Alces alces), an apparent competitor. Starting with a real forest landscape, we created forest management scenarios of the specific spatial distribution of cuts (mosaic, small, or large agglomeration) with increasing disturbance levels. We then ran the IBM with simulated agents, representing individuals of the three species, moving according to movement rules determined from radio‐collared individuals. We found that movement responses to land cover types and roads differed among species. For example, caribou and moose generally avoided areas close to roads, contrary to wolves. Those differences influenced the mortality of caribou agents, which not only depended on the levels of disturbance but also depended on the spatial distribution of cuts and roads. After controlling for disturbance level, wolves were more successful when forest management required an extensive road network resulting in relatively high habitat fragmentation. Caribou agents experienced lower mortality in landscapes with low densities of road and disturbance‐related edges. The effect remained much stronger, however, for the level than the spatial configuration of human disturbances. Still, our IBM demonstrated how landscape management could be used to manipulate species interactions, with the intent of either increasing or decreasing predation rates on specific populations, depending on management goals.

Determinants of vegetation regeneration on forest roads following restoration treatments: implications for boreal caribou conservation

Linear features are increasing worldwide and, in many jurisdictions, their decommissioning has been identified as a way to restore wildlife habitat. Few studies have assessed restoration practices on forest roads, yet they are the main linear disturbance throughout most circumboreal forests. In boreal forests of eastern Canada, such knowledge would be especially valuable for the conservation of boreal populations of woodland caribou (Rangifer tarandus caribou). We assessed the short‐term establishment of vegetation following four treatments applied across 40 km of forest roads, along a restoration gradient involving additive treatments (i.e. each successive treatment included the treatments prior): closing the road to traffic, decompacting the soil, planting black spruce trees, and adding enriched soil. We linked lateral cover (a proxy of movement obstruction for wildlife) and the occurrence and abundance of plant species to road treatments and environmental covariates. Vegetation establishment 3–4 years after decommissioning was mostly influenced by treatments but also by road width and stand composition in the vicinity of roads. The combination of closing, decompacting, and planting was the most effective treatment to establish regeneration that would lead to suitable caribou habitat as it reduced food availability for moose and bears (i.e. lower presence of herbaceous species, fruit‐bearing shrubs, and deciduous trees). It also reduced the presence of plants competing with spruce, such as ericaceous shrubs. Our results suggest that the decommissioning of forest roads could benefit caribou, provided it is performed at a sufficiently broad scale, and accompanied by other habitat restoration and protection practices.

Integrating counts, telemetry, and non‐invasive DNA data to improve demographic monitoring of an endangered species

AbstractPopulation monitoring can take many different forms, and monitoring elusive and endangered species frequently involves a variety of sparse data from different sources. Small populations are often hard to sample precisely and without bias, so when estimates of vital rates like survival or recruitment point to conflicting population trends, it can be hard to determine which is more correct. Furthermore, data can be extremely hard to collect on small populations and it can be helpful to find a way to use all available hard‐won data. To address these issues, we developed an integrated population model (IPM) using all available data to estimate vital rates and abundance for a case study of an endangered woodland caribou (Rangifer tarandus caribou) population. This IPM allowed us to incorporate data from juvenile recruitment surveys, telemetry‐based survival, aerial population counts and mark–resight data, and non‐invasive capture–recapture DNA data to better understand population status and trend. We estimated survival, abundance, and recruitment of four age classes of male and female caribou: young, juveniles, subadults, and adults. The four‐age class structure of the IPM allowed us to estimate recruitment from reproductive‐aged female caribou alone, even though it can be difficult to distinguish age classes—and even sexes—in the field. As part of our IPM, we developed a novel mixture model to break apart data from different age classes when age is unobservable, as it typically is from non‐invasive DNA samples. This helped us decrease bias in juvenile and adult survival estimates from scat data, which was important to our understanding of the population dynamics. Overall, our integrated model provided more precise estimates of population trends than any one method (e.g., telemetry or non‐invasive DNA) alone. This IPM provides a useful, flexible tool for biologists to monitor populations and provides a valuable example of the benefits of integrated population modeling approaches for endangered species management and recovery.

Habitat loss accelerates for the endangered woodland caribou in western Canada

AbstractHabitat loss is often the ultimate cause of species endangerment and is also a leading factor inhibiting species recovery. For this reason, species‐at‐risk legislation, policies and plans typically focus on habitat conservation and restoration as mechanisms for recovery. To assess the effectiveness of these instruments in decelerating habitat loss, we evaluated spatiotemporal habitat changes for an iconic endangered species, woodland caribou (Rangifer tarandus caribou). We quantified changes in forest cover, a key proxy of caribou habitat, for all caribou subpopulations in Alberta and British Columbia, Canada. Despite efforts under federal and provincial recovery plans, and requirements listed under Canada's Species at Risk Act, caribou subpopulations lost twice as much habitat as they gained during a 12‐year period (2000–2012). Drivers of habitat loss varied by ecotype, with Boreal and Northern Mountain caribou affected most by forest fire and Southern Mountain caribou affected more by forest harvest. Our case study emphasizes critical gaps between recovery planning and habitat management actions, which are a core expectation under most species‐at‐risk legislation. Loss of caribou habitat from 2000 to 2018 has accelerated. Linear features within caribou ranges have also increased over time, particularly seismic lines within Boreal caribou ranges, and we estimated that only 5% of seismic lines have functionally regenerated. Our findings support the idea that short‐term recovery actions such as predator reductions and translocations will likely just delay caribou extinction in the absence of well‐considered habitat management. Given the magnitude of ongoing habitat change, it is clear that unless the cumulative impacts of land‐uses are effectively addressed through planning and management actions that consider anthropogenic and natural disturbances, we will fail to achieve self‐sustaining woodland caribou populations across much of North America.

UAV and High Resolution Satellite Mapping of Forage Lichen (Cladonia spp.) in a Rocky Canadian Shield Landscape

Reindeer lichens (Cladonia spp.) are an important food source for woodland and barren ground caribou herds. In this study, we assessed Cladonia classification accuracy in a rocky, Canadian Shield landscape near Yellowknife, Northwest Territories using both Unmanned Aerial Vehicle (UAV) sensors and high-resolution satellite sensors. At the UAV scale, random forest classifications derived from a multispectral, visible-near infrared sensor (Micasense Altum) had an average 5% higher accuracy for mapping Cladonia (i.e., 95.5%) than when using a conventional color RGB camera (DJI Phantom 4 RTK). We aggregated Altum lichen classifications from three 5 ha study sites to train random forest regression models of fractional lichen cover using predictor features from WorldView-3 and Planet CubeSat satellite imagery. WorldView models at 6 m resolution had an average 6.8% RMSE (R 2 = 0.61) when tested at independent study sites and outperformed the 6 m Planet models, which had a 9.9% RMSE (R 2 = 0.34). These satellite results are comparable to previous lichen mapping studies focusing on woodlands, but the small cover of Cladonia in our study area (11.6% or 16.8% within the barren portions) results in a high relative RMSE (62.2%) expressed as a proportion of mean lichen cover.

Dynamic patterns in winter ungulate browse succession in the Boreal Plains of Alberta

Wildfires are a key driver of boreal forest structure and community composition that alter food resources affecting the behaviour and ecology of wildlife. In the first 50 years post-wildfire, woody browse availability in upland forests increase in quantity and quality for generalist ungulates, such as moose (Alces alces) and white-tailed deer (Odocoileus virginianus). Greater favorable habitat for these generalist ungulates results in increases to their respective populations, and through apparent competition, leads to increases in wolf populations; thus, causing unsustainable levels of predation on threatened woodland caribou (Rangifer tarandus caribou) populations. However, the duration of post-wildfire browse availability is not well understood in the Boreal Plains of Alberta as previous studies are primarily from the Taiga and Boreal Shield where vegetation communities are structurally different. This study examines the changes in winter browse richness, evenness, abundance, and community composition, as well as their use (browse levels) by moose and white-tailed deer, over a 150-year post-wildfire period. In the summer of 2019, we collected vegetation and ungulate browsing data from 164 upland and lowland forest sites in northeastern Alberta, Canada. We used analysis of covariance (ANCOVA) and ordinal logistic regression to examine changes in browse measures. Species richness and evenness showed a double peak at 10–25 years and 90 years post-wildfire in mixedwood forests, as a result of fluctuations in browse palatability, while browse abundance was constant. In contrast, black spruce and lowland forests had similar species richness, evenness and abundance over the 150-year chronosequence. However, browse abundance in lowland forests was higher than mixedwood forests, but this consisted of low palatable browse. Browsing was significant in jack pine forests, mixedwood forests and poor fens; coniferous saplings were generally avoided, whereas 35–65% of available deciduous saplings were browsed. Understanding post-wildfire succession and ungulate browsing in post-wildfire forests provides useful information for managing alternative prey populations necessary for long-term woodland caribou conservation.

Spatial familial networks to infer demographic structure of wild populations.

In social species, reproductive success and rates of dispersal vary among individuals resulting in spatially structured populations. Network analyses of familial relationships may provide insights on how these parameters influence population‐level demographic patterns. These methods, however, have rarely been applied to genetically derived pedigree data from wild populations.Here, we use parent–offspring relationships to construct familial networks from polygamous boreal woodland caribou (Rangifer tarandus caribou) in Saskatchewan, Canada, to inform recovery efforts. We collected samples from 933 individuals at 15 variable microsatellite loci along with caribou‐specific primers for sex identification. Using network measures, we assess the contribution of individual caribou to the population with several centrality measures and then determine which measures are best suited to inform on the population demographic structure. We investigate the centrality of individuals from eighteen different local areas, along with the entire population.We found substantial differences in centrality of individuals in different local areas, that in turn contributed differently to the full network, highlighting the importance of analyzing networks at different scales. The full network revealed that boreal caribou in Saskatchewan form a complex, interconnected familial network, as the removal of edges with high betweenness did not result in distinct subgroups. Alpha, betweenness, and eccentricity centrality were the most informative measures to characterize the population demographic structure and for spatially identifying areas of highest fitness levels and family cohesion across the range. We found varied levels of dispersal, fitness, and cohesion in family groups. Synthesis and applications: Our results demonstrate the value of different network measures in assessing genetically derived familial networks. The spatial application of the familial networks identified individuals presenting different fitness levels, short‐ and long‐distance dispersing ability across the range in support of population monitoring and recovery efforts.

Functional Changes to the Slate Islands Provincial Park Ecosystem with Successive Arrival of Wolves, Canis lupus, from the Lake Superior Coast

Observations from 1974-2016 of Caribou (Rangifer tarandus) on the archipelago that comprises Slate Islands Provincial Park allowed us to infer direct and indirect effects of the arrival of Wolf (Canis lupus) pairs in winters of 1993-94 and 2003-04. Wolves created conditions that led to the near demise of Caribou from the islands, including some, but not all, behavioural changes in Caribou consistent with avoiding predators. Caribou on SIPP did not appear to return to calving locations near shoreline areas, nor use them to escape from Wolves by entering water. Shorelines and locations of Patterson Island near a Wolf-occupied Red Fox (Vulpes vulpes) den were the most common Caribou kill locations. Wolves also functionally shifted the ecosystem in Slate Islands Provincial Park via direct and indirect effects on North American Beavers (Castor canadensis), Red Foxes and Snowshoe Hares (Lepus americanus).

Fall supplemental feeding increases population growth rate of an endangered caribou herd.

Most woodland caribou (Rangifer tarandus caribou) populations are declining primarily because of unsustainable predation resulting from habitat-mediated apparent competition. Wolf (Canis lupus) reduction is an effective recovery option because it addresses the direct effect of predation. We considered the possibility that the indirect effects of predation might also affect caribou population dynamics by adversely affecting summer foraging behaviour. If spring and/or summer nutrition was inadequate, then supplemental feeding in fall might compensate for that limitation and contribute to population growth. Improved nutrition and therefore body condition going into winter could increase adult survival and lead to improved reproductive success the next spring. To test that hypothesis, we fed high-quality food pellets to free-ranging caribou in the Kennedy Siding caribou herd each fall for six years, starting in 2014, to see if population growth rate increased. Beginning in winter 2015–16, the Province of British Columbia began a concurrent annual program to promote caribou population increase by attempting to remove most wolves within the Kennedy Siding and the adjacent caribou herds’ ranges. To evaluate the impact of feeding, we compared lambdas before and after feeding began, and to the population trend in the adjacent Quintette herd over the subsequent four years. Supplemental feeding appeared to have an incremental effect on population growth. Population growth of the Kennedy Siding herd was higher in the year after feeding began (λ = 1.06) compared to previous years (λ = 0.91) and to the untreated Quintette herd (λ = 0.95). Average annual growth rate of the Kennedy Siding herd over the subsequent four years, where both feeding and wolf reduction occurred concurrently, was higher than in the Quintette herd where the only management action in those years was wolf reduction (λ = 1.16 vs. λ = 1.08). The higher growth rate of the Kennedy Siding herd was due to higher female survival (96.2%/yr vs. 88.9%/yr). Many caribou were in relatively poor condition in the fall. Consumption of supplemental food probably improved their nutritional status which ultimately led to population growth. Further feeding experiments on other caribou herds using an adaptive management approach would verify the effect of feeding as a population recovery tool. Our results support the recommendation that multiple management actions should be implemented to improve recovery prospects for caribou.

Multiple lines of evidence for predator and prey responses to caribou habitat restoration

How to mitigate the impacts of anthropogenic habitat alteration and restore ecological processes has become an imperative question facing applied ecology. One high-profile example in Canada is boreal woodland caribou, which are declining across North America largely due to anthropogenic habitat alteration and associated changes to predator-prey dynamics. Habitat restoration is increasingly being implemented to recover habitat, as is mandated by federal law. But given the extent of the disturbance and the cost to conduct restoration, evaluating the effectiveness of restoration treatments is needed for effective recovery of caribou populations. We evaluated the effectiveness of silvicultural treatments to reduce predator (wolf and bear) and prey (moose and caribou) use of linear features using a multiple lines of evidence approach. All four species were less likely to be present at treated sites than untreated sites, and daily photo capture rates of moose and wolves when present at treated sites also declined; though effect sizes were typically small. Complimenting the camera-based results, individual moose, bears, and wolves monitored with GPS collars also showed a decline in the use of treated linear features, particularly those with higher intensity treatments, though the response was non-significant. The reduction in the use of treated lines did not scale-up into a significant decline in overall line-use within the treatment area. While we found more evidence than not supporting that animals reduced use of the restoration sites, our study highlights the complexity of monitoring and evaluating the success of habitat restoration. Understanding long-term responses is imperative to ensure habitat restoration is effective.

Coyote (Canis latrans) diet and spatial co-occurrence with woodland caribou (Rangifer tarandus caribou)

The interplay of predator encounters and antipredator responses is an integral part of understanding predator–prey interactions and spatial co-occurrence and avoidance can elucidate these interactions. We conducted hard-part dietary analysis of coyotes (Canis latrans Say, 1823) and space use of coyotes and woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)) to test two competing hypotheses about coyote and caribou predator–prey spatial dynamics using resource selection functions. The high encounter hypothesis predicts that coyotes would maximize encounters with caribou via high spatial co-occurrence, whereas the predator stealth hypothesis predicts that through low spatial co-occurrence with caribou, coyotes act as stealth predators by avoiding habitats that caribou typically select. Our dietary analysis revealed that ∼46% of sampled coyote diet is composed of caribou. We found that coyote share space with caribou in lichen-barren habitat in both summer and winter and that coyotes co-occur with caribou in forested habitat during summer, but not during winter. Our findings support predictions associated with the high encounter predator hypothesis whereby coyotes and caribou have high spatial co-occurrence promoting caribou in coyote diet.

Ecological Interactions Involving Feral Horses and Predators: Review with Implications for Biodiversity Conservation

ABSTRACTFor many ecosystems, feral horses are increasingly becoming an important if not dominant component of ungulate biomass and hence influence on community dynamics. Yet we still know little of how horses contribute to key ecological interactions including predator‐prey and indirect competitive relationships at a community level. Notably, feral species like horses can exhibit life‐history traits that differ from that of native (mainly artiodactyl) herbivore competitors. Artificial selection for traits like increased, early, or extended reproduction that have yet to be reversed by natural selection, coupled with naturally selected differences in anatomy and behavior, in addition to unique management objectives for horses compared to other species, means that the dynamics of feral horse populations are not likely to align with what might be expected of other large herbivores. Unexpected population dynamics and inherent biological asymmetries between native ungulates and feral horses may therefore influence the former via direct competition for shared resources and through enemy‐mediated interactions like apparent competition. In several localities feral horses now co‐exist with multiple native prey species, some of which are in decline or are species at risk. Compounding risks to native species from direct or indirect competitive exclusion by horses is the unique nature and socio‐political context of feral horse management, which tends towards allowing horse populations to be limited largely by natural, density‐dependent factors. We summarize the inherent asymmetries between feral horse biology and that of other ungulate prey species with consequences for conservation, focusing on predator‐prey and emerging indirect interactions in multi‐prey systems, and highlight future directions to address key knowledge gaps in our understanding of how feral horses may now be contributing to the (re)structuring of food webs. Observations of patterns of rapid growth and decline, and associated skews in sex ratios of feral horse populations, indicate a heightened potential for indirect interactions among large ungulate prey species, where there is a prevalence of feral horses as preferred prey, particularly where native prey are declining. In places like western North America, we expect predator‐prey interactions involving feral horses to become an increasingly important factor in the conservation of wildlife. This applies not only to economically or culturally important game species but also at‐risk species, both predators (e.g., wolves [Canis lupus], grizzly bears [Ursus arctos]) and prey (e.g., woodland caribou [Rangifer tarandus caribou]), necessitating an ecological understanding of the role of horses in natural environments that goes beyond that of population control. © 2021 The Wildlife Society.

Drivers of site fidelity in ungulates.

While the tendency to return to previously visited locations-termed 'site fidelity'-is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals' recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a 'win-stay, lose-switch' strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.

Close encounters of the fatal kind: Landscape features associated with central mountain caribou mortalities.

In western Canada, anthropogenic disturbances resulting from resource extraction activities are associated with habitat loss and altered predator–prey dynamics. These habitat changes are linked to increased predation risk and unsustainable mortality rates for caribou (Rangifer tarandus caribou). To inform effective habitat restoration, our goal was to examine whether specific linear disturbance features were associated with caribou predation in central mountain caribou ranges. We used predation‐caused caribou mortalities and caribou GPS‐collar data collected between 2008 and 2015 to assess caribou predation risk within and outside of protected areas at four spatio‐temporal scales: habitat use during the (a) 30 days, (b) 7 days, and (c) 24 hours prior to caribou being killed, and (d) characteristics at caribou kill site locations. Outside of protected areas, predation risk increased closer to pipelines, seismic lines, and streams. Within protected areas, predation risk increased closer to alpine habitat. Factors predicting predation risk differed among spatio‐temporal scales and linear feature types: predation risk increased closer to pipelines during the 30 and 7 days prior to caribou being killed and closer to seismic lines during the 30 days, 7 days, and 24 hours prior, but decreased closer to roads during the 30 days prior to being killed. By assessing habitat use prior to caribou being killed, we identified caribou predation risk factors that would not have been detected by analysis of kill site locations alone. These results provide further evidence that restoration of anthropogenic linear disturbance features should be an immediate priority for caribou recovery in central mountain caribou ranges.