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Predators may create healthier prey populations by selectively removing diseased individuals. Predators typically prefer some ages of prey over others, which may, or may not, align with those prey ages that are most likely to be diseased.The interaction of age‐specific infection and predation has not been previously explored and likely has sizable effects on disease dynamics. We hypothesize that predator cleansing effects will be greater when the disease and predation occur in the same prey age groups.We examine the predator cleansing effect using a model where both vulnerability to predators and pathogen prevalence vary with age. We tailor this model to chronic wasting disease (CWD) in mule deer and elk populations in the Greater Yellowstone Ecosystem, with empirical data from Yellowstone grey wolves and cougars.Model results suggest that under moderate, yet realistic, predation pressure from cougars and wolves independently, predators may decrease CWD outbreak size substantially and delay the accumulation of symptomatic deer and elk. The magnitude of this effect is driven by the ability of predators to selectively remove late‐stage CWD infections that are likely the most responsible for transmission, but this may not be the age class they typically select. Thus, predators that select for infected young adults over uninfected juveniles have a stronger cleansing effect, and these effects are strengthened when transmission rates increase with increasing prey morbidity. There are also trade‐offs from a management perspective—that is, increasing predator kill rates can result in opposing forces on prey abundance and CWD prevalence.Our modelling exploration shows that predators have the potential to reduce prevalence in prey populations when prey age and disease severity are considered, yet the strength of this effect is influenced by predators' selection for demography or body condition. Current CWD management focuses on increasing cervid hunting as the primary management tool, and our results suggest predators may also be a useful tool under certain conditions, but not necessarily without additional impacts on host abundance and demography. Protected areas with predator populations will play a large role in informing the debate over predator impacts on disease. [ABSTRACT FROM AUTHOR]

Understanding trophic cascades in terrestrial wildlife communities is a major challenge because these systems are difficult to sample properly. We show how a tradition of non‐random sampling has confounded this understanding in a textbook system (Yellowstone National Park) where carnivore [Canis lupus (wolf)] recovery is associated with a trophic cascade involving changes in herbivore [Cervus canadensis (elk)] behaviour and density that promote plant regeneration. Long‐term data indicate a practice of sampling only the tallest young plants overestimated regeneration of overstory aspen (Populus tremuloides) by a factor of 4–7 compared to random sampling because it favoured plants taller than the preferred browsing height of elk and overlooked non‐regenerating aspen stands. Random sampling described a trophic cascade, but it was weaker than the one that non‐random sampling described. Our findings highlight the critical importance of basic sampling principles (e.g. randomisation) for achieving an accurate understanding of trophic cascades in terrestrial wildlife systems. [ABSTRACT FROM AUTHOR]

Many species exhibit selective foraging behaviour, where consumers use a nonrandom subset of available food types. Yet little is known about how selective foraging behaviour varies with environmental conditions and the community level consequences of such selection dynamics. We examined selective foraging by wolves preying primarily on elk in Yellowstone National Park (YNP) over a 12-year period and on moose in Isle Royale National Park (IRNP) over a 47-year period. Specifically, we assessed how selection for calves and senescent adults varied with their frequency in the environment, wolf abundance and winter severity. Selection for senescent adults decreased as the relative abundance of senescent prey increased (i.e. negative frequency-dependent selection) in both study sites. In IRNP, selection for calves was also negatively frequency dependent and declined with increasing wolf abundance. These results are inconsistent with the pattern of positive frequency-dependent selection expected under the prey-switching hypothesis. These results suggest that selection is primarily driven by intraspecific differences in prey vulnerability and wolves' interest in minimizing their risk of injury, as opposed to maximizing intake rates. Lastly, we ran simulations to evaluate how predator–prey dynamics were influenced by dynamic patterns of selection, like those observed in YNP and IRNP. The simulations indicated that predators are more efficient (i.e. steeper slope of the numerical response) when selection for calves is negatively frequency dependent, which results in a lower mean abundance of prey. More importantly, predation is a stronger destabilizing force when selection for calves is negatively frequency dependent. That stronger destabilizing force is indicated by greater variability in the abundance of prey and predators, prey populations being less resilient and a steeper negative slope of the relationship between predation rate and prey population growth rate. As such, our simulation analyses suggest that some of the observed patterns of negative-frequency dependent selection may have important consequences for predator–prey dynamics. • Wolves showed negative frequency-dependent selection for calves and senescent prey. • Wolves did not exhibit prey-switching behaviour when preferred prey became rare. • Selection is likely driven by the predator's interest to reduce their risk of injury. • Selection for calves may result in more efficient predators and less resilient prey. [ABSTRACT FROM AUTHOR]

Environmentally mediated changes in body size often underlie population responses to environmental change, yet this is not a universal phenomenon. Understanding when phenotypic change underlies population responses to environmental change is important for obtaining insights and robust predictions of population dynamics in a changing world. We develop a dynamic integral projection model that mechanistically links environmental conditions to demographic rates and phenotypic traits (body size) via changes in resource availability and individual energetics. We apply the model to the northern Yellowstone elk population and explore population responses to changing patterns of seasonality, incorporating the interdependence of growth, demography, and density-dependent processes operating through population feedback on available resources. We found that small changes in body size distributions can have large impacts on population dynamics but need not cause population responses to environmental change. Environmental changes that altered demographic rates directly, via increasing or decreasing resource availability, led to large population impacts in the absence of substantial changes to body size distributions. In contrast, environmentally driven shifts in body size distributions could occur with little consequence for population dynamics when the effect of environmental change on resource availability was small and seasonally restricted and when strong density-dependent processes counteracted expected population responses. These findings highlight that a robust understanding of how associations between body size and demography influence population responses to environmental change will require knowledge of the shape of the relationship between phenotypic distributions and vital rates, the population status with regard to its carrying capacity, and importantly the nature of the environmentally driven change in body size and carrying capacity. [ABSTRACT FROM AUTHOR]

The extent to which prey space use actively minimizes predation risk continues to ignite controversy. Methodological reasons that have hindered consensus include inconsistent measurements of predation risk, biased spatiotemporal scales at which responses are measured and lack of robust null expectations.We addressed all three challenges in a comprehensive analysis of the spatiotemporal responses of adult female elk (Cervus elaphus) to the risk of predation by wolves (Canis lupus) during winter in northern Yellowstone, USA.We quantified spatial overlap between the winter home ranges of GPS‐collared elk and three measures of predation risk: the intensity of wolf space use, the distribution of wolf‐killed elk and vegetation openness. We also assessed whether elk varied their use of areas characterized by more or less predation risk across hours of the day, and estimated encounter rates between simultaneous elk and wolf pack trajectories. We determined whether observed values were significantly lower than expected if elk movements were random with reference to predation risk using a null model approach.Although a small proportion of elk did show a tendency to minimize use of open vegetation at specific times of the day, overall we highlight a notable absence of spatiotemporal response by female elk to the risk of predation posed by wolves in northern Yellowstone.Our results suggest that predator–prey interactions may not always result in strong spatiotemporal patterns of avoidance. [ABSTRACT FROM AUTHOR]

Prey switching occurs when a generalist predator kills disproportionately more of an abundant prey species and correspondingly spares a rarer species. Although this behaviour is a classic stabilizing mechanism in food web models, little is known about its operation in free-living systems which often include dangerous prey species that resist predation., We used long-term (1995-2015) data from a large mammal system in northern Yellowstone National Park, USA, to understand how prey preference of a wild, generalist predator ( Canis lupus) responds to a shift in prey species evenness involving rising numbers of dangerous prey ( Bison bison) and dropping numbers of relatively safer prey ( Cervus elaphus)., Contrary to the prey switching hypothesis, wolves attacked and killed disproportionately more of the rarer, but safer, species. Wolves maintained a strong preference against bison even when this species was nearly twice as abundant as elk. [Correction added after online publication on 26 April 2017: 'more than' changed to 'nearly']. There was also evidence that wolves were increasingly averse to hunting bison as relative bison abundance increased., Wolves seldom hunted bison because capture success was limited to a narrow set of conditions: larger packs (>11 wolves) chasing smaller herds (10-20 bison) with calves. Wolves scavenged bison carrion instead and did so more frequently as bison abundance increased., Our study demonstrates the overarching importance of prey vulnerability to understanding the prey preferences of generalist predators in ecological communities with dangerous prey. The formidable defences of such prey diminish the potential for switching and its stabilizing influence on population dynamics. In these communities, shifts from hunting to scavenging are perhaps more likely than shifts in prey preference. The assumption of switching may therefore overestimate the stability of multi-prey systems that include dangerous prey species., A is available for this article. [ABSTRACT FROM AUTHOR]

Trophic interactions are a fundamental topic in ecology, but we know little about how competition between apex predators affects predation, the mechanism driving top-down forcing in ecosystems. We used long-term datasets from Scandinavia (Europe) and Yellowstone National Park (North America) to evaluate how grey wolf (Canis lupus) kill rate was affected by a sympatric apex predator, the brown bear (Ursus arctos). We used kill interval (i.e. the number of days between consecutive ungulate kills) as a proxy of kill rate. Although brown bears can monopolize wolf kills, we found no support in either study system for the common assumption that they cause wolves to kill more often. On the contrary, our results showed the opposite effect. In Scandinavia, wolf packs sympatric with brown bears killed less often than allopatric packs during both spring (after bear den emergence) and summer. Similarly, the presence of bears at wolf-killed ungulates was associated with wolves killing less often during summer in Yellowstone. The consistency in results between the two systems suggests that brown bear presence actually reduces wolf kill rate. Our results suggest that the influence of predation on lower trophic levels may depend on the composition of predator communities. [ABSTRACT FROM AUTHOR]

Knowledge of characteristics that promote group success during intraspecific encounters is key to understanding the adaptive advantages of sociality for many group-living species. In addition, some individuals in a group may be more likely than others to influence intergroup conflicts, a relatively neglected idea in research on social animals. Here we use observations of aggressive interactions between wolf (Canis lupus) packs over an extended period and use pack characteristics to determine which groups had an advantage over their opponents. During 16 years of observation in Yellowstone National Park from 1995 to 2010, we documented 121 interpack aggressive interactions. We recorded pack sizes, compositions, and spatial orientation related to residency to determine their effects on the outcomes of interactions between packs. Relative pack size (RPS) improved the odds of a pack displacing its opponent. However, pack composition moderated the effect of RPS as packs with relatively more old members (>6.0 years old) or adult males had higher odds of winning despite a numerical disadvantage. The location of the interaction with respect to pack territories had no effect on the outcome of interpack interactions. Although the importance of RPS in successful territorial defense suggests the evolution and maintenance of group living may be at least partly due to larger packs' success during interpack interactions, group composition is also an important factor, highlighting that some individuals are more valuable than others during interpack conflicts. [ABSTRACT FROM AUTHOR]

Understanding the population dynamics of top-predators is essential to assess their impact on ecosystems and to guide their management. Key to this understanding is identifying the mechanisms regulating vital rates., Determining the influence of density on survival is necessary to understand the extent to which human-caused mortality is compensatory or additive. In wolves ( Canis lupus), empirical evidence for density-dependent survival is lacking. Dispersal is considered the principal way in which wolves adjust their numbers to prey supply or compensate for human exploitation. However, studies to date have primarily focused on exploited wolf populations, in which density-dependent mechanisms are likely weak due to artificially low wolf densities., Using 13 years of data on 280 collared wolves in Yellowstone National Park, we assessed the effect of wolf density, prey abundance and population structure, as well as winter severity, on age-specific survival in two areas (prey-rich vs. prey-poor) of the national park. We further analysed cause-specific mortality and explored the factors driving intraspecific aggression in the prey-rich northern area of the park., Overall, survival rates decreased during the study. In northern Yellowstone, density dependence regulated adult survival through an increase in intraspecific aggression, independent of prey availability. In the interior of the park, adult survival was less variable and density-independent, despite reduced prey availability. There was no effect of prey population structure in northern Yellowstone, or of winter severity in either area. Survival was similar among yearlings and adults, but lower for adults older than 6 years., Our results indicate that density-dependent intraspecific aggression is a major driver of adult wolf survival in northern Yellowstone, suggesting intrinsic density-dependent mechanisms have the potential to regulate wolf populations at high ungulate densities. When low prey availability or high removal rates maintain wolves at lower densities, limited inter-pack interactions may prevent density-dependent survival, consistent with our findings in the interior of the park. [ABSTRACT FROM AUTHOR]

Despite the popular view that social predators live in groups because group hunting facilitates prey capture, the apparent tendency for hunting success to peak at small group sizes suggests that the formation of large groups is unrelated to prey capture. Few empirical studies, however, have tested for nonlinear relationships between hunting success and group size, and none have demonstrated why success trails off after peaking. Here, we use a unique dataset of observations of individually known wolves (Canis lupus) hunting elk (Cervus elaphus) in Yellowstone National Park to show that the relationship between success and group size is indeed nonlinear and that individuals withholding effort (free riding) is why success does not increase across large group sizes. Beyond 4 wolves, hunting success leveled off, and individual performance (a measure of effort) decreased for reasons unrelated to interference from inept hunters, individual age, or size. But performance did drop faster among wolves with an incentive to hold back, i.e., nonbreeders with no dependent offspring, those performing dangerous predatory tasks, i.e., grabbing and restraining prey, and those in groups of proficient hunters. These results suggest that decreasing performance was free riding and that was why success leveled off in groups with >4 wolves that had superficially appeared to be cooperating. This is the first direct evidence that nonlinear trends in group hunting success reflect a switch from cooperation to free riding. It also highlights how hunting success per se is unlikely to promote formation and maintenance of large groups. [ABSTRACT FROM AUTHOR]

Environmental change has been observed to generate simultaneous responses in population dynamics, life history, gene frequencies, and morphology in a number of species. But how common are such eco-evolutionary responses to environmental change likely to be? Are they inevitable, or do they require a specific type of change? Can we accurately predict eco-evolutionary responses? We address these questions using theory and data from the study of Yellowstone wolves. We show that environmental change is expected to generate eco-evolutionary change, that changes in the average environment will affect wolves to a greater extent than changes in how variable it is, and that accurate prediction of the consequences of environmental change will probably prove elusive. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

1. Large body size hinders locomotor performance in ways that may lead to trade-offs in predator foraging ability that limit the net predatory benefit of larger size. For example, size-related improvements in handling prey may come at the expense of pursuing prey and thus negate any enhancement in overall predatory performance due to increasing size. 2. This hypothesis was tested with longitudinal data from repeated observations of 94 individually known wolves ( Canis lupus) hunting elk ( Cervus elaphus) in Yellowstone National Park, USA. Wolf size was estimated from an individually based sex-specific growth model derived from body mass measurements of 304 wolves. 3. Larger size granted individual wolves a net predatory advantage despite substantial variation in its effect on the performance of different predatory tasks; larger size improved performance of a strength-related task (grappling and subduing elk) but failed to improve performance of a locomotor-related task (selecting an elk from a group) for wolves > 39 kg. 4. Sexual dimorphism in wolf size also explained why males outperformed females in each of the three tasks considered (attacking, selecting, and killing). 5. These findings support the generalization that bigger predators are overall better hunters, but they also indicate that increasing size ultimately limits elements of predatory behaviour that require superior locomotor performance. We argue that this could potentially narrow the dietary niche of larger carnivores as well as limit the evolution of larger size if prey are substantially more difficult to pursue than to handle. [ABSTRACT FROM AUTHOR]

Current techniques for remotely monitoring wildlife lack the capability to survey a wide area or to transmit data in real time. We addressed these technology gaps by developing and testing a new video and telemetry system for remotely sampling wildlife abundance, distribution, and behavior across large open areas. The system consisted of 2 pan-tilt-zoom video cameras equipped with 20-200X lens, and an automated telemetry scanner and data logger. All components were charged by wind and solar power and located on a hilltop overlooking an open valley (23 km²) in Yellowstone National Park, USA. A satellite up-link to the internet transmitted data in real time to the University of Minnesota-St. Paul and relayed commands from undergraduate students who controlled the cameras and systematically scanned the area at 2- hour intervals 6 times/day (0800-2000 hr) 7 days/week for about 20 consecutive weeks (Dec-Jun). During each scan, students recorded presence, activity, and location of bison (Bison bison), coyote (Canis latrans), grizzly bear (Ursus arctos), red fox (Vulpes vulpes), and wolf (Canis lupus). The telemetry system continuously scanned frequencies of 100 tagged wolves, including 4 members of the one resident pack. We determined wolf presence in real time by viewing the incoming data stream, or we assessed presence later after off-loading the data logger. Matched pairs of simultaneous observations taken by remote and on-site observers during a 13-day double-sampling period were highly correlated (r = 0.71-0.94), but remote observations were biased toward larger, more visible mammals (e.g., bison) and tended to underestimate their abundance. Nevertheless, the system was deployed longer than was practical for on-site observers and was, therefore, useful for detecting long-term, fine-scale trends such as daily changes in bison numbers as a function of snow depth. [ABSTRACT FROM AUTHOR]

Because some native ungulates have lived without top predators for generations, it has been uncertain whether runaway predation would occur when predators are newly restored to these systems. We show that landscape features and vegetation, which influence predator detection and capture of prey, shape large-scale patterns of predation in a newly restored predator–prey system. We analysed the spatial distribution of wolf ( Canis lupus) predation on elk ( Cervus elaphus) on the Northern Range of Yellowstone National Park over 10 consecutive winters. The influence of wolf distribution on kill sites diminished over the course of this study, a result that was likely caused by territorial constraints on wolf distribution. In contrast, landscape factors strongly influenced kill sites, creating distinct hunting grounds and prey refugia. Elk in this newly restored predator–prey system should be able to mediate their risk of predation by movement and habitat selection across a heterogeneous risk landscape. [ABSTRACT FROM AUTHOR]

Presents information on a study which investigated wolf predation on elk in Yellowstone National Park during winter conditions. Methodology; Results of the study; Discussion.

The removal or addition of a predator in an ecosystem can trigger a trophic cascade, whereby the predator indirectly influences plants and/or abiotic processes via direct effects on its herbivore prey. A trophic cascade can operate through a density‐mediated indirect effect (DMIE), where the predator reduces herbivore density via predation, and/or through a trait‐mediated indirect effect (TMIE), where the predator induces an herbivore trait response that modifies the herbivore's effect on plants. Manipulative experiments suggest that TMIEs are an equivalent or more important driver of trophic cascades than are DMIEs. Whether this applies generally in nature is uncertain because few studies have directly compared the magnitudes of TMIEs and DMIEs on natural unmanipulated field patterns. A TMIE is often invoked to explain the textbook trophic cascade involving wolves (Canis lupus), elk (Cervus canadensis), and aspen (Populus tremuloides) in northern Yellowstone National Park. This hypothesis posits that wolves indirectly increase recruitment of young aspen into the overstory primarily through reduced elk browsing in response to spatial variation in wolf predation risk rather than through reduced elk population density. To test this hypothesis, we compared the effects of spatiotemporal variation in wolf predation risk and temporal variation in elk population density on unmanipulated patterns of browsing and recruitment of young aspen across 113 aspen stands over a 21‐year period (1999–2019) in northern Yellowstone National Park. Only 2 of 10 indices of wolf predation risk had statistically meaningful effects on browsing and recruitment of young aspen, and these effects were 8–28 times weaker than the effect of elk density. To the extent that temporal variation in elk density was attributable to wolf predation, our results suggest that the wolf–elk–aspen trophic cascade was primarily density‐mediated rather than trait‐mediated. This aligns with the alternative hypothesis that wolves and other actively hunting predators with broad habitat domains cause DMIEs to dominate whenever prey, such as elk, also have a broad habitat domain. For at least this type of predator–prey community, our study suggests that risk‐induced trait responses can be abstracted or ignored while still achieving an accurate understanding of trophic cascades. [ABSTRACT FROM AUTHOR]

We know much about pathogen evolution and the emergence of new disease strains, but less about host resistance and how it is signaled to other individuals and subsequently maintained. The cline in frequency of black-coated wolves (Canis lupus) across North America is hypothesized to result from a relationship with canine distemper virus (CDV) outbreaks. We tested this hypothesis using cross-sectional data from wolf populations across North America that vary in the prevalence of CDV and the allele that makes coats black, longitudinal data from Yellowstone National Park, and modeling. We found that the frequency of CDV outbreaks generates fluctuating selection that results in heterozygote advantage that in turn affects the frequency of the black allele, optimal mating behavior, and black wolf cline across the continent. [ABSTRACT FROM AUTHOR]