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.     

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.     

Wolf predation on moose and roe deer: chase distances and outcome of encounters

We examined chase distances of gray wolves Canis lupus Linnaeus, 1758 hunting moose Alces alces and roe deer Capreolus capreolus, and recorded details of encounters between wolves and prey on the Scandinavian Peninsula, 1997–2003. In total, 252 wolf attacks on moose and 64 attacks on roe deer were registered during 4200 km of snow tracking in 28 wolf territories. Average chase distances were 76 m for moose and 237 m for roe deer, a difference likely due to variation in body size and vigilance between prey species. A model including prey species, outcome of the attack, and snow depth explained 15–19% of the variation found in chase distances, with shorter chase distances associated with greater snow depth and with successful attacks on moose but not on roe deer. Wolf hunting success did not differ between prey species (moose 43%, roe deer 47%) but in 11% of the wolf attacks on moose at least one moose was injured but not killed, whereas no injured roe deer survived. Compared with most North American wolf studies chase distances were shorter, hunting success was greater, and fewer moose made a stand when attacked by wolves in our study. Differences in wolf encounters with moose and roe deer likely result from different anti-predator behaviour and predator-prey history between prey species.     

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.     

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     

Comparative Patterns of Predation by Cougars and Recolonizing Wolves in Montana's Madison Range

Abstract: Numerous studies have documented how prey may use antipredator strategies to reduce the risk of predation from a single predator. However, when a recolonizing predator enters an already complex predator—prey system, specific antipredator behaviors may conflict and avoidance of one predator may enhance vulnerability to another. We studied the patterns of prey selection by recolonizing wolves (Canis lupus) and cougars (Puma concolor) in response to prey resource selection in the northern Madison Range, Montana, USA. Elk (Cervus elaphus) were the primary prey for wolves, and mule deer (Odocoileus hemionus) were the primary prey for cougars, but elk made up an increasingly greater proportion of cougar kills annually. Although both predators preyed disproportionately on male elk, wolves were most likely to prey on males in poor physical condition. Although we found that the predators partitioned hunting habitats, structural complexity at wolf kill sites increased over time, whereas complexity of cougar kill sites decreased. We concluded that shifts by prey to structurally complex refugia were attempts by formerly naïve prey to lessen predation risk from wolves; nevertheless, shifting to more structurally complex refugia might have made prey more vulnerable to cougars. After a change in predator exposure, use of refugia may represent a compromise to minimize overall risk. As agencies formulate management strategies relative to wolf recolonization, the potential for interactive predation effects (i.e., facilitation or antagonism) should be considered.     

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.     

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.     

The influence of prey consumption and demographic stochasticity on population growth rate of Isle Royale wolves Canis lupus

The relationship between the rates of prey capture and predator population growth is a fundamental aspect of predation, yet it is rarely measured for vertebrate predators. For the isolated wolf population on Isle Royale, annual variation in kill rate explains 22% of the variation in wolf population growth rate. From the slope of this relationship, we estimate that the production efficiency (ratio of production to respiration) of wolves is between 0.5% and 1.5%. More generally, we assess the relative extent to which wolf population growth rate is affected by density dependence, prey availability (moose, Alces alces ), winter weather, and demographic stochasticity. Prey availability explains the most variation in wolf growth rate (42%), but this is only recognized after accounting for the influence of a disease-induced population crash and age structure of the prey population (i.e. number of vulnerable moose, >9 years of age). Demographic stochasticity accounts for approximately 30% of the variation in wolf growth rate. This recognition is important, but not surprising, given that the average population size of Isle Royale wolves is 22. Previous work indicates that the effect of winter climate, as mediated through prey vulnerability and kill rates, is substantial. This work indicates that the direct effect of winter climate is weak, and explains only about 4% of the variation in wolf growth rate (P=0.10).     

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.     

Assessing the influence of prey–predator ratio,prey age structure and packs size on wolf kill rates

Traditional predation theory assumes that prey density is the primary determinant of kill rate. More recently, the ratio of prey‐to‐predator has been shown to be a better predictor of kill rate. However, the selective behavior of many predators also suggests that age structure of the prey population should be an important predictor of kill rate. We compared wolf–moose predation dynamics in two sites, south‐central Scandinavia (SCA) and Isle Royale, Lake Superior, North America (IR), where prey density was similar, but where prey age structure and prey‐to‐predator ratio differed. Per capita kill rates of wolves preying on moose in SCA are three times greater than on IR. Because SCA and IR have similar prey densities differences in kill rate cannot be explained by prey density. Instead, differences in kill rate are explained by differences in the ratio of prey‐to‐predator, pack size and age structure of the prey populations. Although ratio‐dependent functional responses was an important variable for explaining differences in kill rates between SCA and IR, kill rates tended to be higher when calves comprised a greater portion of wolves’ diet (p =0.05). Our study is the first to suggest how age structure of the prey population can affect kill rate for a mammalian predator. Differences in age structure of the SCA and IR prey populations are, in large part, the result of moose and forests being exploited in SCA, but not in IR. While predator conservation is largely motivated by restoring trophic cascades and other top–down influences, our results show how human enterprises can also alter predation through bottom–up processes.     

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.     

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.     

Differential risk effects of wolves on wild versus domestic prey have consequences for conservation

Predators play integral roles in shaping ecosystems through cascading effects to prey and vegetation. Such effects occur when prey species alter their behavior to avoid predators, a phenomenon called the risk effects of predators. Risk effects of wild predators such as wolves are well documented for wild prey, but not for free ranging domestic animals such as cattle despite their importance for ecosystem function and conservation. We compared risk effects of satellite‐collared wolves (n = 16) on habitat selection by global‐positioning‐system‐collared elk (n = 10) and cattle (n = 31). We calculated resource selection functions (RSFs) in periods before, during and after wolf visits in elk home ranges or cattle pastures. The habitat variables tested included: distance to roads and trails, terrain ruggedness, food‐quality and distance to forest. When wolves were present, elk stayed closer to forest cover and selected less for high‐quality‐food habitat. Thus, the risk effects of wolf presence on elk produced a change in the tradeoff between food and cover selection. Cattle responded by avoiding high‐quality‐food habitat and selecting areas closer to roads and trails (where people likely provided security), but these effects manifested only after wolves had left. Artificial selection in cattle may have attenuated natural anti‐predator behaviors. The effects of predators on ecosystems are likely different when mediated through risk effects on domestic compared to wild animals. Furthermore, predator control in response to livestock predation, an important conservation issue, may produce broad ecosystem effects triggered by decrease of an important predator species. Conservation planners should consider these effects where domestic herbivores are dominant species in the ecosystem.     

Disturbance-Mediated Apparent Competition Decouples in a Northern Boreal Caribou Range

The most widely reported threat to boreal and mountain populations of woodland caribou (Rangifer tarandus caribou; caribou) involves habitat- or disturbance-mediated apparent competition (DMAC). With DMAC, natural and anthropogenic disturbances that increase the abundance of deciduous-browsing cervids (e.g., moose [Alces alces], deer [Odocoileus spp.]) are thought to promote predator (especially wolf [Canis lupus]) numbers, which heightens predation risk to caribou. We know most about the effects of DMAC on caribou where the species is under threat by anthropogenic activities in relatively productive southern boreal and mountain systems. Yet, >60% of extant boreal caribou range in North America consists of northern shield and taiga ecoregions of low productivity where caribou may compete with only 1 ungulate species (moose) in the context of DMAC. In this environment, we know very little of how DMAC acts as a limiting factor to caribou. In Saskatchewan, Canada, from 2014–2018, using a combination of vegetation sampling, aerial surveys, and telemetry data (n = 38 wolves), we searched for evidence of DMAC (trends in data consistent with the hypothesis) in an 87,193-km² section of the Western Boreal Shield, a poorly productive but pristine region (0.18% of land cover classed as an anthropogenic feature) with a historically high fire-return interval (47% of stands aged <40 years). Despite the high levels of disturbance, moose density was relatively low (47 moose/1,000 km²), likely because of the scarcity of deciduous or mixed-wood stands and low abundance of deciduous browse in the young conifer stands that dominated the landscape. In contrast, boreal caribou density was relatively high for the species (37 caribou/1,000 km²). Wolf density (3.1 wolves/1,000 km²) and pack sizes ( = 4.0 wolves/pack) were low and resident (established) territories were large ( = 4,360 km²; 100% minimum convex polygon). The low density of wolves mirrored the low (standardized) ungulate biomass index (UBI; moose + boreal caribou) of the study area (0.36 UBI/km²). We conclude that wolf and hence caribou populations were not responding in accordance with the outcomes generally predicted by DMAC in our study area because the requisite strong, positive response to fire of deciduous-browse and alternate-prey abundance was lacking. As a limiting factor to caribou, DMAC is likely modulated at a macroecological scale by factors such as net primary productivity, a corollary to the general hypothesis that we advance here (i.e., primary productivity hypothesis of DMAC). We caution against managing for caribou based on the presumption of DMAC where the mechanism does not apply, which may include much of boreal caribou range in the north. © 2020 The Wildlife Society.     

Ecologically based definition of seasons clarifies predator–prey interactions

Species interactions within food webs are driven by multiple constraints, including those imposed by seasonal changes in the environment. Ecologically sound definitions of seasons may therefore be a prerequisite for clarifying predator prey interactions. Most studies define biological seasons based on fixed schedules or on temporal changes in a single movement measurement. We used a novel clustering approach based on homogeneous space‐use patterns of GPS‐collared animals to reveal 7 biological seasons for caribou Rangifer tarandus caribou, and 5 for both moose Alces alces and grey wolves Canis lupus interacting in a boreal ecosystem. Subsequent evaluation of niche overlap showed that, as predicted, wolves had a stronger spatio‐temporal connection with moose, its main prey, than with caribou. Movement constraints and limiting resource distributions similarly affected all species in some instances, but also caused temporal changes in the extent of niche overlap between wolves and its two prey. The risk that caribou faced was not only linked to the niche overlap with wolves, but also to the extent of wolf‐moose niche overlap during the same period. Food‐web properties emerged from the analysis, with temporal changes in relative niche overlap reflecting the strength of trophic interactions during the year. Our study demonstrates how the study of trophic interactions can benefit from comprehensive definitions of biological seasons.     

Prey selection and dietary response by wolves in a high-density multi-species ungulate community

Studies on predation by the wolf (Canis lupus) have often reported contradictory results about the role of prey density and vulnerability on wolf prey use. We investigated dietary response and prey selection by wolves in a high-density and multi-species ungulate community, analysing scats collected over a period of 11 years in the Casentinesi Forests, Italy. The second most abundant species, wild boar (Sus scrofa), was found to be the main wolf prey, and we did not observe any dietary response of wolves to variations in the density of either primary or secondary prey species. Selection patterns were uniform throughout the study period. Wolves strongly selected for wild boar piglets, while roe deer (Capreolus capreolus) fawns and adults, red deer (Cervus elaphus) adults and fallow deer (Dama dama) adults were avoided. Wolf preference for wild boar was inversely density dependent. Within each species, juveniles were preferred to adults. Medium-sized, young individuals of both wild boar and roe deer were optimal prey, although with different selection patterns related to the different anti-predator strategies adopted by each prey species. The results of this study suggest that in productive ecosystems with high density and high renewal rates of prey, selection patterns by wolves are determined by prey vulnerability, which is connected to prey age and body size. The different patterns of wild boar versus cervids use by wolf across Europe seems to be related to their relative abundances, while the strong selection of wild boar in Italian Apennines with respect to the more frequent avoidance in central-eastern Europe is better explained by higher piglet productivity and smaller body size of adults boar in Mediterranean temperate forests.     

Recolonizing gray wolves increase parasite infection risk in their prey

The recent recolonization of Central Europe by the European gray wolf (Canis lupus) provides an opportunity to study the dynamics of parasite transmission for cases when a definitive host returns after a phase of local extinction. We investigated whether a newly established wolf population increased the prevalence of those parasites in ungulate intermediate hosts representing wolf prey, whether some parasite species are particularly well adapted to wolves, and the potential basis for such adaptations. We recorded Sarcocystis species richness in wolves and Sarcocystis prevalence in ungulates harvested in study sites with and without permanent wolf presence in Germany using microscopy and DNA metabarcoding. Sarcocystis prevalence in red deer (Cervus elaphus) was significantly higher in wolf areas (79.7%) than in control areas (26.3%) but not in roe deer (Capreolus capreolus) (97.2% vs. 90.4%) or wild boar (Sus scrofa) (82.8% vs. 64.9%). Of 11 Sarcocystis species, Sarcocystis taeniata and Sarcocystis grueneri occurred more often in wolves than expected from the Sarcocystis infection patterns of ungulate prey. Both Sarcocystis species showed a higher increase in prevalence in ungulates in wolf areas than other Sarcocystis species, suggesting that they are particularly well adapted to wolves, and are examples of “wolf specialists”. Sarcocystis species richness in wolves was significantly higher in pups than in adults. “Wolf specialists” persisted during wolf maturation. The results of this study demonstrate that (1) predator–prey interactions influence parasite prevalence, if both predator and prey are part of the parasite life cycle, (2) mesopredators do not necessarily replace the apex predator in parasite transmission dynamics for particular parasites of which the apex predator is the definitive host, even if meso‐ and apex predators were from the same taxonomic family (here: Canidae, e.g., red foxes Vulpes vulpes), and (3) age‐dependent immune maturation contributes to the control of protozoan infection in wolves.     

Consequences of ratio-dependent predation by wolves for elk population dynamics

A growing number of studies suggest ratio-dependence may be common in many predator–prey systems, yet in large mammal systems, evidence is limited to wolves and their prey in Isle Royale and Yellowstone. More importantly, the consequences of ratio-dependent predation have not been empirically examined to understand the implications for prey. Wolves recolonized Banff National Park in the early 1980s, and recovery was correlated with significant elk declines. I used time-series data of wolf kill rates of elk, wolf and elk densities in winter from 1985–2007 to test for support for prey-, ratio-, or predator dependent functional and numeric responses of wolf killing rate to elk density. I then combined functional and numeric responses to estimate the total predation response to identify potential equilibrium states. Evidence suggests wolf predation on elk was best described by a type II ratio-dependent functional response and a type II numeric response that lead to inversely density-dependent predation rate on elk. Despite support for ratio-dependence, like other wolf-prey systems, there was considerable uncertainty amongst functional response models, especially at low prey densities. Consistent with predictions from ratio-dependent models, however, wolves contributed to elk population declines of over 80 % in our Banff system. Despite the statistical signature for ratio-dependence, the biological mechanism remains unknown and may be related to multi-prey dynamics in our system. Regardless, ratio-dependent models strike a parsimonious balance between theory and empiricism, and this study suggests that large mammal ecologists need to consider ratio-dependent models in predator–prey dynamics.     

Cross‐continental comparison of parasite communities in a wide‐ranging carnivore suggests associations with prey diversity and host density

1. Parasites are integral to ecosystem functioning yet often overlooked. Improved understanding of host–parasite associations is important, particularly for wide‐ranging species for which host range shifts and climate change could alter host–parasite interactions and their effects on ecosystem function.
2. Among the most widely distributed mammals with diverse diets, gray wolves (Canis lupus) host parasites that are transmitted among canids and via prey species. Wolf–parasite associations may therefore influence the population dynamics and ecological functions of both wolves and their prey. Our goal was to identify large‐scale processes that shape host–parasite interactions across populations, with the wolf as a model organism.
3. By compiling data from various studies, we examined the fecal prevalence of gastrointestinal parasites in six wolf populations from two continents in relation to wolf density, diet diversity, and other ecological conditions.
4. As expected, we found that the fecal prevalence of parasites transmitted directly to wolves via contact with other canids or their excreta was positively associated with wolf density. Contrary to our expectations, the fecal prevalence of parasites transmitted via prey was negatively associated with prey diversity. We also found that parasite communities reflected landscape characteristics and specific prey items available to wolves.
5. Several parasite taxa identified in this study, including hookworms and coccidian protozoans, can cause morbidity and mortality in canids, especially in pups, or in combination with other stressors. The density–prevalence relationship for parasites with simple life cycles may reflect a regulatory role of gastrointestinal parasites on wolf populations. Our result that fecal prevalence of parasites was lower in wolves with more diverse diets could provide insight into the mechanisms by which biodiversity may regulate disease. A diverse suite of predator–prey interactions could regulate the effects of parasitism on prey populations and mitigate the transmission of infectious agents, including zoonoses, spread via trophic interactions.