Is “Wolf‐Pack” Predation by Antimicrobial Bacteria Cooperative? Cell Behaviour and Predatory Mechanisms Indicate Profound Selfishness,Even when Working Alongside Kin

For decades, myxobacteria have been spotlighted as exemplars of social “wolf‐pack” predation, communally secreting antimicrobial substances into the shared public milieu. This behavior has been described as cooperative, becoming more efficient if performed by more cells. However, laboratory evidence for cooperativity is limited and of little relevance to predation in a natural setting. In contrast, there is accumulating evidence for predatory mechanisms promoting “selfish” behavior during predation, which together with conflicting definitions of cooperativity, casts doubt on whether microbial “wolf‐pack” predation really is cooperative. Here, it is hypothesized that public‐goods‐mediated predation is not cooperative, and it is argued that a holistic model of microbial predation is needed, accounting for predator and prey relatedness, social phenotypes, spatial organization, activity/specificity/transport of secreted toxins, and prey resistance mechanisms. Filling such gaps in our knowledge is vital if the evolutionary benefits of potentially costly microbial behaviors mediated by public goods are to be properly understood.     

Distribution, status and conservation of Indian gray wolf (Canis lupus pallipes) in Karnataka, India

The Indian gray wolf Canis lupus pallipes is the major large carnivore in the plains of Karnataka, India. We carried out a study on its distribution and status from November 2001 to July 2004. We estimated 555 wolves occupying about 123 330 km² of the state. In the past 40 years, wolves have disappeared from the southern plateau from an area of about 31 801 km². Their distribution is now largely restricted to the north-eastern dry plains. The wolf has also disappeared in recent years from some 'protected areas' such as Melkote Temple Wildlife Sanctuary, and their present population is largely found in 'non-protected' areas. Blackbucks are the only natural prey of wolves in Karnataka, but their density in most parts of the state is extremely low. The major prey species is domestic livestock, especially sheep. The available 'remote area' (forests or rocky terrains) in the wolf-occupied regions determined the status of the wolf. Killing of adult wolves and pups was common throughout the range of the wolf. However, such killings were made largely by local sheepherders with small sheep holdings and not by nomadic shepherds who maintained large sheep herds. The forests in the north-eastern parts of the state exist in small patches every few kilometers. Because each wolf pack ranges over large distances and is by and large a commensal species, we propose that the management of these small forest patches, considering them as components of a larger landscape, is the only effective conservation practice for the wolf. Although existing locally in low densities, because of a large ranging area of a single pack, the seemingly isolated wolf packs can become parts of a large metapopulation, providing a sustainable population.     

What best explains vigilance in elk: characteristics of prey, predators, or the environment?

We quantified the vigilance levels of elk (Cervus elaphus) preyedon by wolves (Canis lupus) in Yellowstone National Park, betweenJanuary and May in 2005 and 2006, and used Akaike's informationcriterion to compare a set of 38 regression models for vigilancelevels. These models combined up to 9 predictor variables of3 types: characteristics of the prey group (herd size and composition),characteristics of the predator (wolf pack size, distance away,and the presence/absence of a kill), and characteristics ofthe local environment (distance to woodland edges, snow depth,and snow cover). The set of models spanned a range of complexityfrom simple univariate models to complex combinations with upto 3 variables of each type. Complex models incorporating thecharacteristics of the wolf pack, the structure of the elk herd,and the environmental conditions had higher information contentthan simple models. Although univariate models of vigilancedetect significant relationships, they have low informationcontent relative to multivariate models. These results showthat elk assesses factors of several types when assessing riskand deciding how much time to allocate to vigilance. In particular,we found that all well-supported models of vigilance includedseveral "prey" variables and several "predator" variables. Thisresult highlights the need to consider information about predatorswhen trying to explain variation in vigilance levels in prey.     

Effect of sociality and season on gray wolf (Canis lupus) foraging behavior: implications for estimating summer kill rate

Background

Understanding how kill rates vary among seasons is required to understand predation by vertebrate species living in temperate climates. Unfortunately, kill rates are only rarely estimated during summer.

Methodology/Principal Findings

For several wolf packs in Yellowstone National Park, we used pairs of collared wolves living in the same pack and the double-count method to estimate the probability of attendance (PA) for an individual wolf at a carcass. PA quantifies an important aspect of social foraging behavior (i.e., the cohesiveness of foraging). We used PA to estimate summer kill rates for packs containing GPS-collared wolves between 2004 and 2009. Estimated rates of daily prey acquisition (edible biomass per wolf) decreased from 8.4±0.9 kg (mean ± SE) in May to 4.1±0.4 kg in July. Failure to account for PA would have resulted in underestimating kill rate by 32%. PA was 0.72±0.05 for large ungulate prey and 0.46±0.04 for small ungulate prey. To assess seasonal differences in social foraging behavior, we also evaluated PA during winter for VHF-collared wolves between 1997 and 2009. During winter, PA was 0.95±0.01. PA was not influenced by prey size but was influenced by wolf age and pack size.

Conclusions/Significance

Our results demonstrate that seasonal patterns in the foraging behavior of social carnivores have important implications for understanding their social behavior and estimating kill rates. Synthesizing our findings with previous insights suggests that there is important seasonal variation in how and why social carnivores live in groups. Our findings are also important for applications of GPS collars to estimate kill rates. Specifically, because the factors affecting the PA of social carnivores likely differ between seasons, kill rates estimated through GPS collars should account for seasonal differences in social foraging behavior.     

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.     

Spatial organization in the Ethiopian wolf Canis simensis: large packs and small stable home ranges

The spatial organization of the rare Ethiopian wolf ( Canis simensis ) was studied in the Afroalpine heathlands of Bale Mountains National Park, southern Ethiopia, between 1988 and 1992. Nineteen animals were radio-tracked, 48 ear-tagged and 64 others recognized by coat patterns and observed directly. Dry season (October—March) home ranges of resident wolves covered between 2 and 15 km². The ranges of adult males were slightly larger than those of females, and subadults' home ranges were slightly smaller than those of adults. The population density of the wolves was correlated with prey biomass. In optimal habitat, wolves lived in packs of 3—13 adults (mean 5.9 wolves > 1year old) containing several close-kin males; adult sex ratio favoured males 1.88: 1 and combined pack home ranges averaged 6.0km². In an area of lower prey productivity, wolves lived in pairs or small groups (mean 2.7), adult sex ratio was 1:1 and home ranges averaged 13.4 km².
Home ranges overlapped extensively (mean 85%) between members of the same pack. Four to seven percent of the population was additionally composed of non-resident females, inhabiting larger ranges (mean 11.1 km²). Home ranges of neighbouring packs were largely discrete, forming a tessellating mosaic of packs occupying all available habitat. Pack home ranges were stable in time, drifting only during major pack readjustment after the disappearance of a pack or significant demographic changes. Ethiopian wolf home ranges were smaller than would be expected for a carnivore of its size and sociality, presumably as a result of the high rodent productivity of the Afroalpine ecosystem.     

Experimental assessment of trophic ecology in a generalist spider predator: Implications for biocontrol in Uruguayan crops

Conservative biological control promotes the use of native natural enemies to limit the size and growth of pest populations. Although spiders constitute one of the most important groups of native predators in several crops, their trophic ecology remains largely unknown, especially for several generalist taxa. In laboratory, we assessed the predatory behaviour of a wandering spider (the wolf spider Lycosa thorelli (Keyserling, 1877) against several arthropods varying in size and trophic positions, all found in South American soybean and rice crops. As prey we used the bug Piezodorus guildinii (Westwood, 1837) as well as larvae and adults of the moth Spodoptera frugiperda (Smith, 1797), both being considered important pests in Uruguayan crops. We also used several non-pest arthropods as prey, sarcophagid flies, carabid beetles and wolf spiders. All prey were attacked in more or less high, although not statistically differing, proportions. However, carabids were not consumed, and bugs were consumed in significantly lower proportions than flies. A negative correlation was found between prey size and acceptance rate. Immobilization times were longer against larvae when compared to moths and flies, while predatory sequences were longer for bugs when compared to flies, moths and spiders. In addition, we found a positive effect of prey size on predatory sequence length and complexity. Our results confirm the ability of spiders to attack and feed upon prey with different morphologies, included well-defended arthropods, and their potential use as natural enemies of several pests in South American crops.     

Landscape of fear in Europe: wolves affect spatial patterns of ungulate browsing in Białowieża Primeval Forest,Poland

Large carnivores can either directly influence ungulate populations or indirectly affect their behaviour. Knowledge from European systems, in contrast to North American systems, on how this might lead to cascading effects on lower trophic levels is virtually absent. We studied whether wolves Canis lupus via density‐mediated and behaviorally‐mediated effects on their ungulate prey species influence patterns of browsing and tree regeneration inside the Bia?owie?a National Park, Poland. Browsing intensity of tree saplings (height class < 150 cm), irrespective of tree species or forest type, was lower inside a wolf core area (50.5%) where predator presence is highest, than in the remainder of the wolf pack’s home range (58.3%). Additionally, browsing intensity was reduced when the amount of coarse woody debris (CWD), which can act as a ?ungulate escape impediment?, increased (within 5‐m radius) inside the wolf core area. No relationship existed outside the core area. As a result, the proportion of trees growing out of herbivore control increased more strongly with increasing amount of CWD inside compared to outside the wolf core area. This suggests that next to direct effects of wolves on ungulate density caused by a higher predation pressure inside the core area, risk effects are important and are enhanced by habitat characteristics. These results indicate that behaviorally‐mediated effects of predators on prey can become more important than density‐mediated effects in affecting lower trophic levels. This is the first study we are aware of, that shows CWD can create fine‐scale risk effects on ungulates with the potential for cascading effects of large predators on patterns of tree regeneration for a European forest system. This knowledge broadens the discussion on how the impact of large predators on ecosystem functioning depends on the physical landscape, by illustrating these effects for a system which largely contrasts in this respect to the North American systems.     

Territory size of wolves Canis lupus: linking local (Białowieża Primeval Forest, Poland) and Holarctic-scale patterns

Factors affecting territory size in wolves Canis lupus were studied at 2 scales, the local population (Bia?owie?a Primeval Forest (BPF), eastern Poland) and the geographic range of species (literature review from 14 localities in the Holarctic). Four packs of wolves were studied by radio‐tracking in BPF from 1994 to 1999. The annual territories of packs (Minimum convex polygons with 95% of locations) averaged 201 km² (SD 63, range 116–310). Core areas of territories (50% MCP) covered from 14 to 78 km² (mean 35). Territory sizes and core areas both were negatively correlated to the encounter rates of ungulates (mean number of ungulates seen per unit time spent in the forest by human observers). Pack size (3–8 wolves) did not influence territory size. Home ranges of individual wolves from the same pack varied with season as well as the age, sex, and reproductive status of the wolf. Review of literature from North America and Europe (42–66^oN), showed that latitude and prey biomass were essential factors shaping the biogeographic variation in wolf territory size. Territories increased with latitude and declined with growing biomass of prey. The analysis showed that latitude acted partly independently of the south–north gradient in prey abundance. At similar standing crop of ungulate biomass (100 kg km^?2), wolf territories would average 140 km² at 40^oN, 370 km² at 50^oN, and 950 km² at 60^oN. Pack size was larger at northern latitudes, but the increase did not keep pace with enlargement of territories. Within‐territory density of wolves declined from 2.5–3 wolves 100 km^?2 at 40–45^oN to 0.7 wolves 100 km^?2 at 60^oN. Our analyses documented similarities regarding the role of prey resources in shaping wolf territoriality at the different scales. Furthermore, a macroecological approach revealed additional factors affecting wolf territory size that were not emergent from knowledge of local population.     

ESS analysis of mechanistic models for territoriality: the value of scent marks in spatial resource partitioning

In this paper elements of game theory are used to analyse a spatially explicit home range model for interacting wolf packs. The model consists of a system of nonlinear partial differential equations whose parameters reflect the movement behavior of individuals within each pack and whose solutions describe the patterns of space-use by each pack. By modifying the behavioral parameters, packs adjust their patterns of movement so as to maximize their reproductive output. This involves a tradeoff between maximizing prey intake and minimizing conflict with neighbors. Evolutionarily stable choices of the behavioral parameters yields territories that are immune to invasion by groups with alternate behaviors.     

A reappraisal of the evidence for regulation of wolf populations

The dogma that gray wolf (Canis lupus) population densities in naturally occurring systems are limited almost solely by available ungulate biomass is based upon studies that fit straight line linear regressions (Type 1 numerical response) to data collected at 32 sites across North America. We fit Type 1, 2, and 3 response functions to the data using linear and nonlinear regression as appropriate and found that the evidence supported wolf population regulation by density-dependence as much as limitation by prey availability. When we excluded 4 of 32 points from the original data set because those points represented exploited or expanding wolf populations the data suggested that wolf populations are self regulated rather than limited by prey biomass by at least a 3:1 margin. In establishing goals for sustainable wolf population levels, managers of wolf reintroductions and species recovery efforts should account for the possibility that some regulatory mechanism plays an important role in wolf population dynamics. © 2011 The Wildlife Society.     

Predatory senescence in ageing wolves

It is well established that ageing handicaps the ability of prey to escape predators, yet surprisingly little is known about how ageing affects the ability of predators to catch prey. Research into long-lived predators has assumed that adults have uniform impacts on prey regardless of age. Here we use longitudinal data from repeated observations of individually-known wolves ( Canis lupus ) hunting elk ( Cervus elaphus ) in Yellowstone National Park to demonstrate that adult predatory performance declines with age and that an increasing ratio of senescent individuals in the wolf population depresses the rate of prey offtake. Because this ratio fluctuates independently of population size, predatory senescence may cause wolf populations of equal size but different age structure to have different impacts on prey populations. These findings suggest that predatory senescence is an important, though overlooked, factor affecting predator-prey dynamics.     

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).     

Home range size variation in a recovering wolf population: evaluating the effect of environmental, demographic, and social factors

Home range size in mammals is a key ecological trait and an important parameter in conservation planning, and has been shown to be influenced by ecological, demographic and social factors in animal populations. Information on space requirements is especially important for carnivore species which range over very large areas and often come into direct conflict with human interest. We used long-term telemetry-location data from a recovering wolf population in Scandinavia to investigate variation in home range size in relation to environmental and social characteristics of the different packs. Wolves showed considerable variation in home range size, which ranged from 259 to 1,676 km². Although wolf density increased fourfold during the study period, we found no evidence that intraspecific competition influenced range size. Local variation in moose density, which was the main prey for most packs, did not influence wolf home range size. Home ranges increased with latitude and elevation and decreased with increased roe deer density. Although prey biomass alone did not influence range size, our data suggest that there is a correlation between habitat characteristics, choice of prey species and possible hunting success, which currently combine to shape home range size in Scandinavian wolves.     

Predators in natural fragments: foraging ecology of wolves in British Columbia's central and north coast archipelago

Aim Predator–prey dynamics in fragmented areas may be influenced by spatial features of the landscape. Although little is known about these processes, an increasingly fragmented planet underscores the urgency to predict its consequences. Accordingly, our aim was to examine foraging behaviour of an apex mammalian predator, the wolf (Canis lupus), in an archipelago environment. Location Mainland and adjacent archipelago of British Columbia, Canada; a largely pristine and naturally fragmented landscape with islands of variable size and isolation. Methods We sampled 30 mainland watersheds and 29 islands for wolf faeces in summers 2000 and 2001 and identified prey remains. We examined broad geographical patterns and detailed biogeographical variables (area and isolation metrics) as they relate to prey consumed. For island data, we used Akaike Information Criteria to guide generalized linear regression model selection to predict probability of black‐tailed deer (main prey; Odocoileus hemionus) in faeces. Results Black‐tailed deer was the most common item in occurrence per faeces (63%) and occurrence per item (53%) indices, representing about 63% of mammalian biomass. Wolves consumed more deer on islands near the mainland (65% occurrence per item) than on the mainland (39%) and outer islands (45%), where other ungulates (mainland only) and small mammals replaced deer. On islands, the probability of detecting deer was influenced primarily by island distance to mainland (not by area or inter‐landmass distance), suggesting limited recolonization by deer from source populations as a causal mechanism. Main conclusions Although sampling was limited in time, consistent patterns among islands suggest that population dynamics in isolated fragments are less stable and can result in depletion of prey. This may have important implications in understanding predator–prey communities in isolation, debate regarding wolf–deer systems and logging in temperate rain forests, and reserve design.     

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.     

Implications of Harvest on the Boundaries of Protected Areas for Large Carnivore Viewing Opportunities

The desire to see free ranging large carnivores in their natural habitat is a driver of tourism in protected areas around the globe. However, large carnivores are wide-ranging and subject to human-caused mortality outside protected area boundaries. The impact of harvest (trapping or hunting) on wildlife viewing opportunities has been the subject of intense debate and speculation, but quantitative analyses have been lacking. We examined the effect of legal harvest of wolves (Canis lupus) along the boundaries of two North American National Parks, Denali (DNPP) and Yellowstone (YNP), on wolf viewing opportunities within the parks during peak tourist season. We used data on wolf sightings, pack sizes, den site locations, and harvest adjacent to DNPP from 1997–2013 and YNP from 2008–2013 to evaluate the relationship between harvest and wolf viewing opportunities. Although sightings were largely driven by wolf population size and proximity of den sites to roads, sightings in both parks were significantly reduced by harvest. Sightings in YNP increased by 45% following years with no harvest of a wolf from a pack, and sightings in DNPP were more than twice as likely during a period with a harvest buffer zone than in years without the buffer. These findings show that harvest of wolves adjacent to protected areas can reduce sightings within those areas despite minimal impacts on the size of protected wolf populations. Consumptive use of carnivores adjacent to protected areas may therefore reduce their potential for non-consumptive use, and these tradeoffs should be considered when developing regional wildlife management policies.     

Population responses of roe deer to the recolonization of the French Vercors by wolves

In a context of changing carnivore populations worldwide, it is crucial to understand the consequences of these changes for prey populations. The recolonization by wolves of the French Vercors mountain range and the long-term monitoring (2001–2017) of roe deer in this area provided a unique opportunity to assess the effects of wolves on this prey. Roe deer was the main prey of wolves in the west Vercors mountain range during this recolonization. We compared roe deer abundance and fawn body mass in two contrasted areas of a wolf pack territory: a central area (core of the territory characterized by an intense use by wolves) and a peripheral area (used more occasionally). Roe deer population growth rates were lower in the central area between 2001 and 2006, resulting in a decline in roe deer abundance. Roe deer abundance substantially dropped in the two study areas after an extremely severe winter but the abundance of roe deer in the central area facing with wolves was slower to recover and remained at lower abundance levels for 6 years. Fawn body mass was consistently lower in the central area, varied similarly as roe deer abundance, and was not influenced by weather conditions or red deer population abundance. Altogether, the effects of wolves on roe deer in the central area occurred during a 10-year period following the establishment of wolves, through the interplay between wolf predation (before wolves started preying on red deer), harsh winter conditions and possibly naivety of prey to this recolonizing predator.     

Linking anti‐predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore

Ecological theory predicts that the diffuse risk cues generated by wide‐ranging, active predators should induce prey behavioural responses but not major, population‐ or community‐level consequences. We evaluated the non‐consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high‐risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20‐fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.     

Effects of wolf pack size and winter conditions on elk mortality

Elk (Cervus canadensis) are high-profile game animals for many states in the western United States, yet over the past several decades some populations have experienced a persistent and broad-scale decline in recruitment. Over this same period, gray wolves (Canis lupus) have become an integral component of many western landscapes and agencies are increasingly challenged to maximize hunting opportunities of ungulates via predator management while simultaneously ensuring wolf conservation. To better understand the implications of predator management on elk populations, we monitored survival of 1,244 adult female elk and 806 6-month-old calves from 29 populations distributed throughout Idaho, USA, from 2004 to 2016. We developed predictive models of mortality that related mortality risk to wolf pack size, winter conditions, and individual-level characteristics. Annual mortality rates (excluding harvest) for adult females and calves were 0.09 and 0.40, respectively. Calf mortality was predicted best with a model that included additive effects of chest girth at time of capture, mean size of surrounding wolf packs, and snow depth. Adult female mortality was predicted best with a model that included female age, mean size of surrounding wolf packs, and snow depth. Based on a sensitivity analysis, chest girth had the largest effect on risk of mortality for calves followed by pack size and snow depth. Other than the effect of senescence in the oldest (>15 yr) individuals, pack size and snow depth had the largest effect on risk of mortality for adult females. We estimated cause-specific mortality and predation was the dominant cause of known-fate mortalities for adult females (35% mountain lion [Puma concolor] and 32% wolf) and calves (45% mountain lion and 28% wolf), whereas malnutrition accounted for 9% and 10% of adult female and calf mortalities, respectively. Wolves preferentially selected smaller calves and older adult females, whereas mountain lions showed little preference for calf size or age class of adult females. Our study indicates managers can increase elk survival by reducing wolf pack sizes on surrounding winter ranges, especially in areas where, or during years when, snow is deep. Additionally, managers interested in improving over-winter calf survival can implement actions to increase the size of calves entering winter by increasing the nutritional quality of summer and early fall forage resources. Although our study was prompted by management questions related to wolves, mountain lions killed more elk than wolves and differences in selection of individual elk indicate mountain lions may have comparably more of an effect on elk population dynamics. Although we were unable to relate changes in mountain lion populations to elk survival in our study, future research should seek a better understanding of multi-predator systems, including how management of one predator affect others and ultimately how these interactions affect elk survival. © 2019 The Wildlife Society