Mobility of moose—comparing the effects of wolf predation risk,reproductive status,and seasonality

In a predator–prey system, prey species may adapt to the presence of predators with behavioral changes such as increased vigilance, shifting habitats, or changes in their mobility. In North America, moose (Alces alces) have shown behavioral adaptations to presence of predators, but such antipredator behavioral responses have not yet been found in Scandinavian moose in response to the recolonization of wolves (Canis lupus). We studied travel speed and direction of movement of GPS‐collared female moose (n = 26) in relation to spatiotemporal differences in wolf predation risk, reproductive status, and time of year. Travel speed was highest during the calving (May–July) and postcalving (August–October) seasons and was lower for females with calves than females without calves. Similarly, time of year and reproductive status affected the direction of movement, as more concentrated movement was observed for females with calves at heel, during the calving season. We did not find support for that wolf predation risk was an important factor affecting moose travel speed or direction of movement. Likely causal factors for the weak effect of wolf predation risk on mobility of moose include high moose‐to‐wolf ratio and intensive hunter harvest of the moose population during the past century.     

Response of Moose Hunters to Predation following Wolf Return in Sweden

Background

Predation and hunter harvest constitute the main mortality factors affecting the size and dynamics of many exploited populations. The re-colonization by wolves (Canis lupus) of the Scandinavian Peninsula may therefore substantially reduce hunter harvest of moose (Alces alces), the main prey of wolves.

Methodology/Principal findings

We examined possible effects of wolf presence on hunter harvest in areas where we had data before and after wolf establishment (n = 25), and in additional areas that had been continuously exposed to wolf predation during at least ten years (n = 43). There was a general reduction in the total number of moose harvested (n = 31,827) during the ten year study period in all areas irrespective of presence of wolves or not. However, the reduction in hunter harvest was stronger within wolf territories compared to control areas without wolves. The reduction in harvest was larger in small (500-800 km²) compared to large (1,200-1,800 km²) wolf territories. In areas with newly established wolf territories moose management appeared to be adaptive with regard to both managers (hunting quotas) and to hunters (actual harvest). In these areas an instant reduction in moose harvest over-compensated the estimated number of moose killed annually by wolves and the composition of the hunted animals changed towards a lower proportion of adult females.

Conclusions/Significance

We show that the re-colonization of wolves may result in an almost instant functional response by another large predator—humans—that reduced the potential for a direct numerical effect on the density of wolves’ main prey, the moose. Because most of the worlds’ habitat that will be available for future colonization by large predators are likely to be strongly influenced by humans, human behavioural responses may constitute a key trait that govern the impact of large predators on their prey.     

What Cues Do Ungulates Use to Assess Predation Risk in Dense Temperate Forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.     

Encounter frequencies between GPS-collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory

Over 6,000 GPS fixes from two wolves (Canis lupus) and 30,000 GPS fixes from five moose (Alces alces) in a wolf territory in southern Scandinavia were used to assess the static and dynamic interactions between predator and prey individuals. Our results showed that wolves were closer to some of the moose when inside their home ranges than expected if they had moved independently of each other, and we also found a higher number of close encounters (<500 m) than expected. This suggests that the wolves were actively seeking the individual moose within their territory. Furthermore, the wolves showed a preference for moving on gravel forest roads, which may be used as convenient travel routes when patrolling the territory and seeking areas where the moose are. However, due to the particularly large size of the wolf territory combined with relatively high moose densities, the wolves generally spent a very small proportion of their time inside the home range of each individual moose, and the frequency of encounters between the wolves and any particular moose was very low. We suggest that the high moose:wolf ratio in this large Scandinavian wolf territory compared to that typically occurring in North America, results in a relatively low encounter frequency and a low predation risk for individual moose, as the predation pressure is spread over a high number of prey individuals.     

The distribution of <Emphasis Type="Italic">Echinococcus granulosus</Emphasis> in moose: evidence for parasite-induced vulnerability to predation by wolves?

The role of parasites in influencing the trophic dynamics of hosts is becoming increasingly recognized in the ecological literature. Echinococcus granulosus is a tapeworm that relies on the predator-prey relationship between the definitive host (wolf, Canis lupus) and the intermediate host, (moose, Alces alces) to complete its life cycle. Heavy infection by E. granulosus may predispose moose to increased risk of predation by wolves. Theory predicts that parasite-induced vulnerability to predation will reduce the degree of aggregation of parasites in a host population. We tested for different levels of aggregation of E. granulosus in moose in areas of low, moderate, and high levels of wolf predation using Greens coefficient of dispersion. Parasite aggregation was lower in an area with high predation rate, thus we hypothesize that heavy infection by E. granulosus predisposes moose to predation by wolves. This increase in predation rate due to parasite infection may influence the role of wolves in regulating moose populations. We discuss alternative explanations for the negative correlation between predation rate and parasite aggregation.An erratum to this article can be found at     

Invading white-tailed deer change wolf–caribou dynamics in northeastern Alberta

Human-caused habitat change has been implicated in current woodland caribou (Rangifer tarandus caribou) population declines across North America. Increased early seral habitat associated with industrial footprint can result in an increase in ungulate densities and subsequently those of their predator, wolves (Canis lupus). Higher wolf densities can result in increased encounters between wolves and caribou and consequently higher caribou mortality. We contrasted changes in moose (Alces alces) and deer (Odocoileus spp.) densities and assessed their effects on wolf–caribou dynamics in northeastern Alberta, Canada, pre (1994–1997) versus post (2005–2009) major industrial expansion in the region. Observable white-tailed deer (O. virginianus) increased 17.5-fold but moose remained unchanged. Wolf numbers also increased from approximately 6–11.5/1,000 km². Coincident with these changes, spatial overlap between wolf pack territories and caribou range was high relative to the mid-1990s. The high number of wolf locations in caribou range suggests that forays were not merely exploratory, but rather represented hunting forays and denning locations. Scat analysis indicated that wolf consumption of moose declined substantively during this time period, whereas use of deer increased markedly and deer replaced moose as the primary prey of wolves. Caribou increased 10-fold in the diet of wolves and caribou population trends in the region changed from stable to declining. Wolf use of beaver (Castor canadensis) increased since the mid-1990s. We suggest that recent declines in woodland caribou populations in the southerly extent of their range have occurred because high deer densities resulted in a numeric response by wolves and consequently higher incidental predation on caribou. Our results indicate that management actions to conserve caribou must now include deer in primary prey and wolf reduction programs. © 2010 The Wildlife Society     

Moose-wolf dynamics and the natural regulation of moose populations

Summary In southwestern Québec, non-harvested moose populations stabilize at a density of 0.40 animal·km^-2. In an attempt to test whether or not moose were regulated by predators, we investigated wolf predation near this equilibrium density (0.37) and at 2 lower densities (0.23, 0.17). Scat analysis in summer and feeding observations in winter indicated a greater use of alternative food resources by wolves at lower moose densities. Each wolf pack killed on average 5.3, 1.8, 1.1 moose·100 days in the area of 0.37, 0.23, and 0.17 moose·km^-2, respectively. Consumption of moose per wolf was 2.8, 1.7, and 1.6 kg/day, respectively. January wolf densities were estimated at 1.38, 0.82, and 0.36 animals·100 km^-2, respectively. Year-long predation rates proved to be density-dependent, increasing with moose density from 6.1 to 19.3% of the postnatal populations. We conclude that moose populations in southwestern Québec are regulated largely by predators (wolves and maybe black bears) at a density where competition for forage produces no detrimental effect. We support the concept that wolf predation can have an important regulatory effect at low moose densities but also a depensatory (inversely density-dependent) effect at higher densities.     

Diet shift of a facultative scavenger, the wolverine, following recolonization of wolves

1. Wolves Canis lupus L. recolonized the boreal forests in the southern part of the Scandinavian peninsula during the late 1990s, but so far there has been little attention to its effect on ecosystem functioning. Wolf predation increases the availability of carcasses of large prey, especially moose Alces alces L., which may lead in turn to a diet switch in facultative scavengers such as the wolverine Gulo gulo L. 2. Using 459 wolverine scats collected during winter-spring 2001-04 for DNA identity and dietary contents, we compared diet inside and outside wolf territories while controlling for potential confounding factors, such as prey density. We tested the hypothesis that wolverine diet shifted towards moose in the presence of wolves, while taking into account possible sexual segregation between the sexes. Occurrence of reindeer, moose and small prey was modelled against explanatory covariates using logistic mixed-effects models. Furthermore, we compared diet composition and breadth among habitats and sexes. 3. Occurrence of reindeer, moose and small prey in the diet varied with prey availability and habitat. As expected, diet contained more moose and less reindeer and small prey in the presence of wolves. Their diet in tundra consisted of 40% reindeer Rangifer tarandus L., 39% moose and 9% rodents. In forest with wolf, their diet shifted to 76% moose, 18% reindeer and 5% rodents; compared to 42% moose, 32% reindeer and 15% rodents in forest without wolf. This diet switch could not be explained by higher moose density in wolf territories. Female diet consisted of more small prey than for males, but there was a tendency for females to use the highly available moose carrion opportunistically and to hunt less on small prey within wolf territories. 4. Our study highlights how wolves increase scavenging opportunities for wolverines, and how sexual differences in diet may also apply to large scavengers. Due to their more restricted home range, female wolverines are forced to rely more on hunting small prey. The relatively high occurrence of wolf kills, however, forms an important food source to wolverines in this area. The recolonization of wolves may therefore have contributed to the consequent recolonization of wolverines into the same area.     

Summer kill rates and predation pattern in a wolf–moose system: can we rely on winter estimates?

So far the vast majority of studies on large carnivore predation, including kill rates and consumption, have been based on winter studies. Because large carnivores relying on ungulates as prey often show a preference for juveniles, kill rates may be both higher and more variable during the summer season than during the rest of the year leading to serious underestimates of the total annual predation rate. This study is the first to present detailed empirical data on kill rates and prey selection in a wolf–moose system during summer (June–September) as obtained by applying modern Global Positioning System-collar techniques on individual wolves (Canis lupus) in Scandinavia. Moose (Alces alces) was the dominant prey species both by number (74.4%) and biomass (95.6%); 89.9% of all moose killed were juveniles, representing 76.0% of the biomass consumed by wolves. Kill rate in terms of the kilogram biomass/kilogram wolf per day averaged 0.20 (range: 0.07–0.32) among wolf territories and was above, or well above, the daily minimum food requirements in most territories. The average number of days between moose kills across wolf territories and study periods was 1.71 days, but increased with time and size of growing moose calves during summer. Over the entire summer (June–September, 122 days), a group (from two to nine) of wolves killed a total of 66 (confidence interval 95%; 56–81) moose. Incorporation of body growth functions of moose calves and yearlings and wolf pups over the summer period showed that wolves adjusted their kill rate on moose, so the amount of biomass/kilogram wolf was relatively constant or increased. The kill rate was much higher (94–116%) than estimated from the winter period. As a consequence, projecting winter kill rates to obtain annual estimates of predation in similar predator–prey systems may result in a significant underestimation of the total number of prey killed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Wolf-avoidance strategies of moose

Two possible anti-predator strategies of prey were suggested by the distribution of moose at Isle Royale, a 544 km² island with numerous offshore islets. Aerial surveys in winter indicated that moose density on small islets (<1.5 km²) averaged 415% greater than on the main island, with about 11% of the calf population existing on the small islets which comprise less than 2% of the total land area. On small islets and shorelines, protection from wolves (in the form of escape terrain) is provided by nearby water in summer and coniferous cover in winter. Calves on islands appear to have a higher survival rate. In summer, cow moose with calves were seen more commonly in camps with human activity (and no wolves) than away from camps, suggesting that people provided a refuge from wolf predation for this unhunted moose population. These subtle patterns of habitat selection by moose are attributed to the intensive natural selection pressure of wolf predation.     

Carnivore-livestock conflicts: determinants of wolf (Canis lupus) depredation on sheep farms in Finland

Wolves (Canis lupus) have recently expanded their distribution range into western and southern Finland, which has not hosted breeding wolves for over 100 years. This has raised concerns and public debate over wolf-livestock conflicts. Between 1998 and 2004 there were 45 wolf attacks on sheep on 34 farms. To assess the risk wolves may pose to sheep husbandry, we used data on depredation, sheep management, landscape structure and moose and wolf populations from continental Finland outside the area of reindeer husbandry to build models of the factors that may predispose sheep farms to wolf depredation. Our results provided evidence that sheep farms with the highest risk of wolf depredation were those located in regions where wolves were abundant. These farms were usually located close to the Russian border where the landscape is a mosaic of forest, wetlands and clear cut areas. These regions are sparsely populated by humans and farms are located far from each other. Finally, we generated probability maps based on generalised additive modelling to predict the risk of wolf predation on livestock in farms of southern Finland.     

Ecosystem Scale Declines in Elk Recruitment and Population Growth with Wolf Colonization: A Before-After-Control-Impact Approach

The reintroduction of wolves (Canis lupus) to Yellowstone provided the unusual opportunity for a quasi-experimental test of the effects of wolf predation on their primary prey (elk – Cervus elaphus) in a system where top-down, bottom-up, and abiotic forces on prey population dynamics were closely and consistently monitored before and after reintroduction. Here, we examined data from 33 years for 12 elk population segments spread across southwestern Montana and northwestern Wyoming in a large scale before-after-control-impact analysis of the effects of wolves on elk recruitment and population dynamics. Recruitment, as measured by the midwinter juvenile∶female ratio, was a strong determinant of elk dynamics, and declined by 35% in elk herds colonized by wolves as annual population growth shifted from increasing to decreasing. Negative effects of population density and winter severity on recruitment, long recognized as important for elk dynamics, were detected in uncolonized elk herds and in wolf-colonized elk herds prior to wolf colonization, but not after wolf colonization. Growing season precipitation and harvest had no detectable effect on recruitment in either wolf treatment or colonization period, although harvest rates of juveniles∶females declined by 37% in wolf-colonized herds. Even if it is assumed that mortality due to predation is completely additive, liberal estimates of wolf predation rates on juvenile elk could explain no more than 52% of the total decline in juvenile∶female ratios in wolf-colonized herds, after accounting for the effects of other limiting factors. Collectively, these long-term, large-scale patterns align well with prior studies that have reported substantial decrease in elk numbers immediately after wolf recolonization, relatively weak additive effects of direct wolf predation on elk survival, and decreased reproduction and recruitment with exposure to predation risk from wolves.     

Contrasting Effects of Wolves and Human Hunters on Elk Behavioral Responses to Predation Risk

ABSTRACT Prey behavioral responses to predation risk in wolf-ungulate-plant systems are of interest to wildlife managers. Using Global Positioning System data collected from telemetry-collared elk (Cervus elaphus) and wolves (Canis lupus), we evaluated elk behavioral responses to spatial and temporal variation in wolf- and human-predation risk on a winter range in the Greater Yellowstone Area, USA. We found elk changed grouping patterns and increased movement rates as predation risk increased and that these behavioral changes were habitat dependent. Elk behavioral responses to wolf- and human-predation risk were similar; however, responses to human-predation risk were stronger than responses to wolf-predation risk. These results suggest that predation risk from wolves or human hunters may result in elk spending more time on private rangelands away from public-land winter ranges, which may exacerbate problems of landowner tolerance of elk on livestock pastures. However, increased movement and changing grouping patterns on winter ranges may also disperse elk grazing impacts and lessen elk impacts on any one area.     

The scale-dependent impact of wolf predation risk on resource selection by three sympatric ungulates

Resource selection is a fundamental ecological process impacting population dynamics and ecosystem structure. Understanding which factors drive selection is vital for effective species- and landscape-level management. We used resource selection probability functions (RSPFs) to study the influence of two forms of wolf (Canis lupus) predation risk, snow conditions and habitat variables on white-tailed deer (Odocoileus virginianus), elk (Cervus elaphus) and moose (Alces alces) resource selection in central Ontario's mixed forest French River-Burwash ecosystem. Direct predation risk was defined as the frequency of a predator's occurrence across the landscape and indirect predation risk as landscape features associated with a higher risk of predation. Models were developed for two winters, each at two spatial scales, using a combination of GIS-derived and ground-measured data. Ungulate presence was determined from snow track transects in 64 16- and 128 1-km(2) resource units, and direct predation risk from GPS radio collar locations of four adjacent wolf packs. Ungulates did not select resources based on the avoidance of areas of direct predation risk at any scale, and instead exhibited selection patterns that tradeoff predation risk minimization with forage and/or mobility requirements. Elk did not avoid indirect predation risk, while both deer and moose exhibited inconsistent responses to this risk. Direct predation risk was more important to models than indirect predation risk but overall, abiotic topographical factors were most influential. These results indicate that wolf predation risk does not limit ungulate habitat use at the scales investigated and that responses to spatial sources of predation risk are complex, incorporating a variety of anti-predator behaviours. Moose resource selection was influenced less by snow conditions than cover type, particularly selection for dense forest, whereas deer showed the opposite pattern. Temporal and spatial scale influenced resource selection by all ungulate species, underlining the importance of incorporating scale into resource selection studies.     

Black bear (<Emphasis Type="Italic">Ursus americanus</Emphasis>) and wolf (<Emphasis Type="Italic">Canis</Emphasis> spp.) summer diet composition and ungulate prey selectivity in Ontario,Canada

Understanding predator-prey dynamics is an important component of management strategy development for wildlife populations that are directly affected by predation. Ungulates often serve as a significant source of prey for many large mammal predators, and patterns of predation are known to influence population dynamics. Although black bear and wolf diets have been investigated extensively, prey preference has been less commonly examined, especially in analyses that take into account age class (i.e., juvenile and adult) of the ungulate prey. We examined black bear (Ursus americanus), wolf (Canis lupus), and hybrids (Canis spp.) prey preference in Ontario based on the availability of three ungulate species—elk (Cervus elaphus), moose (Alces alces), and white-tailed deer (Odocoileus virginianus). We analyzed the presence of prey items in black bear and wolf scats collected over 3 years by examining prey hair cuticular scale patterns. We applied correction factors to frequencies of occurrence of prey items found in predator scat and related diet composition to the availability of ungulates, determined by fecal pellet transect surveys. In addition, non-ungulate diet items were identified to obtain full diet composition profiles. We found that black bear diet consisted of more than 87% vegetation, and they were opportunistic, not selecting for any particular ungulate species in either adult or juvenile age class. Wolf diet was comprised mainly of ungulates (~?73.2%), muskrat (Ondatra zibethica; ~?8.5%), and beaver (Castor canadensis; ~?14.6%), and although moose were at least 1.5 times more abundant then each of the other ungulate prey species in the study area, wolves preferred elk, using moose less than expected. Although we found black bear diet to be opportunistic during the summer, wolves in our study heavily utilized both juvenile and adult ungulates, however, among ungulate species, displayed preference for elk. The preference displayed by wolves provides insight that wildlife managers can use to guide further investigation and assist with the development of strategies to ensure continued elk reintroduction success, and moose and white-tailed deer population sustainability.     

Biomass Flow and Scavengers Use of Carcasses after Re-Colonization of an Apex Predator

Background

Reestablishment of apex predators influences the availability and distribution of biomass for scavengers and can therefore be an important agent for structuring species communities. We studied how the re-colonization of the Scandinavian Peninsula by wolves (Canis lupus) affected the amount and temporal variation in use of moose (Alces alces) carcasses.

Methodology/Principal Findings

We compared the availability of biomass from remains at wolf kills with those killed by hunters, vehicle collisions and natural death. Movement-triggered cameras monitored patterns of use on wolf kills and remains from hunter harvest by scavengers (n = 15 276) in relation to time of year, available carcass biomass, time since the death of the moose and presence of wolves. Remains from hunter harvest were the largest food source for scavengers both within wolf territories (57%) and in areas without wolves (81%). The total annual biomass available were similar in areas with (25 648 kg) and without (24 289 kg) wolves. Presence of wolves lowered the peak biomass available from hunter harvest in October (20%) and increased biomass available during December to August (38–324% per month). The probability of scavengers being present decreased faster with time at remains from hunter harvest compared to wolf kills and both the probability of being present and the number of visits by scavengers to wolf kills increased as the amount of biomass available on the carcass increased.

Conclusions/Significance

Wolves reduced the seasonal variation of biomass from moose carcasses and most important increased it during spring. Scavengers also visited wolf kills most frequently during spring when most scavenging species have young, which may lead to an increase in survival and/or reproductive success of scavengers within wolf territories. This applies both for abundant scavenging species that were the most frequent visitors at wolf kills and threatened scavengers with lower visit frequency.     

Conservation of wildlife populations: factoring in incremental disturbance

Progressive anthropogenic disturbance can alter ecosystem organization potentially causing shifts from one stable state to another. This potential for ecosystem shifts must be considered when establishing targets and objectives for conservation. We ask whether a predator–prey system response to incremental anthropogenic disturbance might shift along a disturbance gradient and, if it does, whether any disturbance thresholds are evident for this system. Development of linear corridors in forested areas increases wolf predation effectiveness, while high density of development provides a safe‐haven for their prey. If wolves limit moose population growth, then wolves and moose should respond inversely to land cover disturbance. Using general linear model analysis, we test how the rate of change in moose (Alces alces) density and wolf (Canis lupus) harvest density are influenced by the rate of change in land cover and proportion of land cover disturbed within a 300,000 km² area in the boreal forest of Alberta, Canada. Using logistic regression, we test how the direction of change in moose density is influenced by measures of land cover change. In response to incremental land cover disturbance, moose declines occurred where <43% of land cover was disturbed; in such landscapes, there were high rates of increase in linear disturbance and wolf density increased. By contrast, moose increases occurred where >43% of land cover was disturbed and wolf density declined. Wolves and moose appeared to respond inversely to incremental disturbance with the balance between moose decline and wolf increase shifting at about 43% of land cover disturbed. Conservation decisions require quantification of disturbance rates and their relationships to predator–prey systems because ecosystem responses to anthropogenic disturbance shift across disturbance gradients.     

Spatial Partitioning of Predation Risk in a Multiple Predator-Multiple Prey System

ABSTRACT Minimizing risk of predation from multiple predators can be difficult, particularly when the risk effects of one predator species may influence vulnerability to a second predator species. We decomposed spatial risk of predation in a 2-predator, 2-prey system into relative risk of encounter and, given an encounter, conditional relative risk of being killed. Then, we generated spatially explicit functions of total risk of predation for each prey species (elk [Cervus elaphus] and mule deer [Odocoileus hemionus]) by combining risks of encounter and kill. For both mule deer and elk, topographic and vegetation type effects, along with resource selection by their primary predator (cougars [Puma concolor] and wolves [Canis lupus], respectively), strongly influenced risk of encounter. Following an encounter, topographic and vegetation type effects altered the risk of predation for both ungulates. For mule deer, risk of direct predation was largely a function of cougar resource selection. However, for elk, risk of direct predation was not only a function of wolf occurrence, but also of habitat attributes that increased elk vulnerability to predation following an encounter. Our analysis of stage-based (i.e., encounter and kill) predation indicates that the risk effect of elk shifting to structurally complex habitat may ameliorate risk of direct predation by wolves but exacerbate risk of direct predation by cougars. Information on spatiotemporal patterns of predation will be become increasingly important as state agencies in the western United States face pressure to integrate predator and prey management.     

Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

Moose movement rates are altered by wolf presence in two ecosystems

Predators directly impact prey populations through lethal encounters, but understanding nonlethal, indirect effects is also critical because foraging animals often face trade‐offs between predator avoidance and energy intake. Quantifying these indirect effects can be difficult even when it is possible to monitor individuals that regularly interact. Our goal was to understand how movement and resource selection of a predator (wolves; Canis lupus) influence the movement behavior of a prey species (moose; Alces alces). We tested whether moose avoided areas with high predicted wolf resource use in two study areas with differing prey compositions, whether avoidance patterns varied seasonally, and whether daily activity budgets of moose and wolves aligned temporally. We deployed GPS collars on both species at two sites in northern Minnesota. We created seasonal resource selection functions (RSF) for wolves and modeled the relationship between moose first‐passage time (FPT), a method that discerns alterations in movement rates, and wolf RSF values. Larger FPT values suggest rest/foraging, whereas shorter FPT values indicate travel/fleeing. We found that the movements of moose and wolves peaked at similar times of day in both study areas. Moose FPTs were 45% lower in areas most selected for by wolves relative to those avoided. The relationship between wolf RSF and moose FPT was nonlinear and varied seasonally. Differences in FPT between low and high RSF values were greatest in winter (?82.1%) and spring (?57.6%) in northeastern Minnesota and similar for all seasons in the Voyageurs National Park ecosystem. In northeastern Minnesota, where moose comprise a larger percentage of wolf diet, the relationship between moose FPT and wolf RSF was more pronounced (ave. across seasons: ?60.1%) than the Voyageurs National Park ecosystem (?30.4%). These findings highlight the role wolves can play in determining moose behavior, whereby moose spend less time in areas with higher predicted likelihood of wolf resource selection.