Prey risk allocation in a grazing ecosystem.

Understanding the behaviorally mediated indirect effects of predators in ecosystems requires knowledge of predator-prey behavioral interactions. In predator-ungulate-plant systems, empirical research quantifying how predators affect ungulate group sizes and distribution, in the context of other influential variables, is particularly needed. The risk allocation hypothesis proposes that prey behavioral responses to predation risk depend on background frequencies of exposure to risk, and it can be used to make predictions about predator-ungulate-plant interactions. We determined non-predation variables that affect elk (Cervus elaphus) group sizes and distribution on a winter range in the Greater Yellowstone Ecosystem (GYE) using logistic and log-linear regression on surveys of 513 1-km2 areas conducted over two years. Employing model selection techniques, we evaluated risk allocation and other a priori hypotheses of elk group size and distributional responses to wolf (Canis lupus) predation risk while accounting for influential non-wolf-predation variables. We found little evidence that wolves affect elk group sizes, which were strongly influenced by habitat type and hunting by humans. Following predictions from the risk allocation hypothesis, wolves likely created a more dynamic elk distribution in areas that they frequently hunted, as elk tended to move following wolf encounters in those areas. This response should dilute elk foraging pressure on plant communities in areas where they are frequently hunted by wolves. We predict that this should decrease the spatial heterogeneity of elk impacts on grasslands in areas that wolves frequently hunt. We also predict that this should decrease browsing pressure on heavily browsed woody plant stands in certain areas, which is supported by recent research in the GYE.     

Elk decision-making rules are simplified in the presence of wolves

The risk of predation drives many behavioral responses in prey. However, few studies have directly tested whether predation risk alters the way other variables influence prey behavior. Here we use information theory (Akaike’s Information Criterion, AICc) in a novel way to test the hypothesis that the decision-making rules governing elk behavior are simplified by the presence of wolves. With elk habitat use as the dependent variable, we test whether the number of independent variables (i.e., the size of the models) that best predict this behavior differ when wolves are present vs absent. Thus, we use AICc scores simply to determine the number of variables to which elk respond when making decisions. We measured habitat use using 2,288 locations from GPS collars on 14 elk, over two winters (14 elk winters), in the Gallatin Canyon portion of the Greater Yellowstone Ecosystem. We found that the use of three major habitat components (grass, conifer, sage) was sensitive to many variables on days that wolves were locally absent, with the best models (ΔAICc≤2) averaging 7.4 parameters. In contrast, habitat use was sensitive to few variables on days when wolves were present: the best models averaged only 2.5 parameters. Because fewer variables affect elk behavior in the presence of wolves, we conclude that elk use simpler decision-making rules in the presence of wolves. This simplification of decision-making rules implies that predation risk imposes selection pressures that do not allow prey to respond to other pressures in ways that they otherwise would. If the affected processes are important, then this indirect effect of predation is likely to be important.     

Are wolves saving Yellowstone's aspen? A landscape-level test of a behaviorally mediated trophic cascade

Behaviorally mediated trophic cascades (BMTCs) occur when the fear of predation among herbivores enhances plant productivity. Based primarily on systems involving small-bodied predators, BMTCs have been proposed as both strong and ubiquitous in natural ecosystems. Recently, however, synthetic work has suggested that the existence of BMTCs may be mediated by predator hunting mode, whereby passive (sit-and-wait) predators have much stronger effects than active (coursing) predators. One BMTC that has been proposed for a wide-ranging active predator system involves the reintroduction of wolves (Canis lupus) to Yellowstone National Park, USA, which is thought to be leading to a recovery of trembling aspen (Populus tremuloides) by causing elk (Cervus elaphus) to avoid foraging in risky areas. Although this BMTC has been generally accepted and highly popularized, it has never been adequately tested. We assessed whether wolves influence aspen by obtaining detailed demographic data on aspen Stands using tree rings and by monitoring browsing levels in experimental elk exclosures arrayed across a gradient of predation risk for three years. Our study demonstrates that the historical failure of aspen to regenerate varied widely among stands (last recruitment year ranged from 1892 to 1956), and our data do not indicate an abrupt cessation of recruitment. This pattern of recruitment failure appears more consistent with a gradual increase in elk numbers rather than a rapid behavioral shift in elk foraging following wolf extirpation. In addition, our estimates of relative survivorship of young browsable aspen indicate that aspen are not currently recovering in Yellowstone, even in the presence of a large wolf population. Finally, in an experimental test of the BMTC hypothesis we found that the impacts of elk browsing on aspen demography are not diminished in sites where elk are at higher risk of predation by wolves. These findings suggest the need to further evaluate how trophic cascades are mediated by predator-prey life history and ecological context.     

Cover Caption

Cover: During winter in the interior of Yellowstone National Park most elk migrate from deep snow while many bison do not, existing near thermal areas and on wind-blown ridges. Wolves still attempt to kill these bison and often spend hours making a kill. Wolf-bison systems used to dominate in North America prior to European settlement but now only Yellowstone and Wood Buffalo National Parks have intact wolf-bison systems. See pp. 1105-1116. Photo by Doug Smith, National Park Service.     

Willow on Yellowstone's northern range: evidence for a trophic cascade?

Reintroduction of wolves (Canis lupus) to Yellowstone National Park in 1995-1996 has been argued to promote a trophic cascade by altering elk (Cervus elaphus) density, habitat-selection patterns, and behavior that, in turn, could lead to changes within the plant communities used by elk. We sampled two species of willow (Salix boothii and S. geyeriana) on the northern winter range to determine whether (1) there was quantitative evidence of increased willow growth following wolf reintroduction, (2) browsing by elk affected willow growth, and (3) any increase in growth observed was greater than that expected by climatic and hydrological factors alone, thereby indicating a trophic cascade caused by wolves. Using stem sectioning techniques to quantify historical growth patterns we found an approximately twofold increase in stem growth-ring area following wolf reintroduction for both species of willow. This increase could not be explained by climate and hydrological factors alone; the presence of wolves on the landscape was a significant predictor of stem growth above and beyond these abiotic factors. Growth-ring area was positively correlated with the previous year's ring area and negatively correlated with the percentage of twigs browsed from the stem during the winter preceding growth, indicating that elk browse impeded stem growth. Our results are consistent with the hypothesis of a behaviorally mediated trophic cascade on Yellowstone's northern winter range following wolf reintroduction. We suggest that the community-altering effects of wolf restoration are an endorsement of ecological-process management in Yellowstone National Park.     

Desert pastoralists’ negative and positive effects on rare wildlife in the Gobi

In arid regions of the developing world, pastoralists and livestock commonly inhabit protected areas, resulting in human–wildlife conflict. Conflict is inextricably linked to the ecological processes shaping relationships between pastoralists and native herbivores and carnivores. To elucidate relationships underpinning human–wildlife conflict, we synthesized 15 years of ecological and ethnographic data from Ikh Nart Nature Reserve in Mongolia's Gobi steppe. The density of argali (Ovis ammon), the world's largest wild sheep, at Ikh Nart was among the highest in Mongolia, yet livestock were >90% of ungulate biomass and dogs >90% of large‐carnivore biomass. For argali, pastoral activities decreased food availability, increased mortality from dog predation, and potentially increased disease risk. Isotope analyses indicated that livestock accounted for >50% of the diet of the majority of gray wolves (Canis lupus) and up to 90% of diet in 25% of sampled wolves (n = 8). Livestock composed at least 96% of ungulate prey in the single wolf pack for which we collected species‐specific prey data. Interviews with pastoralists indicated that wolves annually killed 1–4% of Ikh Nart's livestock, and pastoralists killed wolves in retribution. Pastoralists reduced wolf survival by killing them, but their livestock were an abundant food source for wolves. Consequently, wolf density appeared to be largely decoupled from argali density, and pastoralists had indirect effects on argali that could be negative if pastoralists increased wolf density (apparent competition) or positive if pastoralists decreased wolf predation (apparent facilitation). Ikh Nart's argali population was stable despite these threats, but livestock are increasingly dominant numerically and functionally relative to argali. To support both native wildlife and pastoral livelihoods, we suggest training dogs to not kill argali, community insurance against livestock losses to wolves, reintroducing key native prey species to hotspots of human–wolf conflict, and developing incentives for pastoralists to reduce livestock density.     

Indirect effects and traditional trophic cascades: a test involving wolves, coyotes, and pronghorn

The traditional trophic cascades model is based on consumer resource interactions at each link in a food chain. However, trophic-level interactions, such as mesocarnivore release resulting from intraguild predation, may also be important mediators of cascades. From September 2001 to August 2004, we used spatial and seasonal heterogeneity in wolf distribution and abundance in the southern Greater Yellowstone Ecosystem to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus), accounts for high rates of coyote predation on pronghorn (Antilocapra americana) fawns observed in some areas. Results of this ecological perturbation in wolf densities, coyote densities, and pronghorn neonatal survival at wolf-free and wolf-abundant sites support the existence of a species-level trophic cascade. That wolves precipitated a trophic cascade was evidenced by fawn survival rates that were four-fold higher at sites used by wolves. A negative correlation between coyote and wolf densities supports the hypothesis that interspecific interactions between the two species facilitated the difference in fawn survival. Whereas densities of resident coyotes were similar between wolf-free and wolf-abundant sites, the abundance of transient coyotes was significantly lower in areas used by wolves. Thus, differential effects of wolves on solitary coyotes may be an important mechanism by which wolves limit coyote densities. Our results support the hypothesis that mesopredator release of coyotes contributes to high rates of coyote predation on pronghorn fawns, and demonstrate the importance of alternative food web pathways in structuring the dynamics of terrestrial systems.     

Covariates affecting spatial variability in bison travel behavior in Yellowstone National Park.

Understanding mechanisms influencing the movement paths of animals is essential for comprehending behavior and accurately predicting use of travel corridors. In Yellowstone National Park (USA), the effects of roads and winter road grooming on bison (Bison bison) travel routes and spatial dynamics have been debated for more than a decade. However, no rigorous studies have been conducted on bison spatial movement patterns. We collected 121 380 locations from 14 female bison with GPS collars in central Yellowstone to examine how topography, habitat type, roads, and elevation affected the probability of bison travel year-round. We also conducted daily winter bison road use surveys (2003-2005) to quantify how topography and habitat type influenced spatial variability in the amount of bison road travel. Using model comparison techniques, we found the probability of bison travel and spatial distribution of travel locations were affected by multiple topographic and habitat type attributes including slope, landscape roughness, habitat type, elevation, and distances to streams, foraging areas, forested habitats, and roads. Streams were the most influential natural landscape feature affecting bison travel, and results suggest the bison travel network throughout central Yellowstone is spatially defined largely by the presence of streams that connect foraging areas. Also, the probability of bison travel was higher in regions of variable topography that constrain movements, such as in canyons. Pronounced travel corridors existed both in close association with roads and distant from any roads, and results indicate that roads may facilitate bison travel in certain areas. However, our findings suggest that many road segments used as travel corridors are overlaid upon natural travel pathways because road segments receiving high amounts of bison travel had similar landscape features as natural travel corridors. We suggest that most spatial patterns in bison road travel are a manifestation of general spatial travel trends. Our research offers novel insights into bison spatial dynamics and provides conceptual and analytical frameworks for examining movement patterns of other species.     

Shifting prey selection generates contrasting herbivore dynamics within a large-mammal predator-prey web

Shifting prey selection has been identified as a mechanism potentially regulating predator-prey interactions, but it may also lead to different outcomes, especially in more complex systems with multiple prey species available. We assessed changing prey selection by lions, the major predator for 12 large herbivore species in South Africa's Kruger National Park. The database was provided by records of found carcasses ascribed to kills by lions assembled over 70 years, coupled with counts of changing prey abundance extending over 30 years. Wildebeest and zebra constituted the most favored prey species during the early portion of the study period, while selection for buffalo rose in the south of the park after a severe drought increased their vulnerability. Rainfall had a negative influence on the proportional representation of buffalo in lion kills, but wildebeest and zebra appeared less susceptible to being killed under conditions of low rainfall. Selection by lions for alternative prey species, including giraffe, kudu, waterbuck, and warthog, was influenced by the changing relative abundance and vulnerability of the three principal prey species. Simultaneous declines in the abundance of rarer antelope species were associated with a sharp increase in selection for these species at a time when all three principal prey species were less available. Hence shifting prey selection by lions affected the dynamics of herbivore populations in different ways: promoting contrasting responses by principal prey species to rainfall variation, while apparently being the main cause of sharp declines by alternative prey species under certain conditions. Accordingly, adaptive responses by predators, to both the changing relative abundance of the principal prey species, and other conditions affecting the relative vulnerability of various species, should be taken into account to understand the interactive dynamics of multispecies predator-prey webs.     

Recolonizing wolves and mesopredator suppression of coyotes: impacts on pronghorn population dynamics.

Food web theory predicts that the loss of large carnivores may contribute to elevated predation rates and, hence, declining prey populations, through the process of mesopredator release. However, opportunities to test predictions of the mesopredator release hypothesis are rare, and the extent to which changes in predation rates influence prey population dynamics may not be clear due to a lack of demographic information on the prey population of interest. We utilized spatial and seasonal heterogeneity in wolf distribution and abundance to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus) throughout much of the United States, contributes to high rates of neonatal mortality in ungulates. To test this hypothesis, we contrasted causes of mortality and survival rates of pronghorn (Antilocapra americana) neonates captured at wolf-free and wolf-abundant sites in western Wyoming, USA, between 2002 and 2004. We then used these data to parameterize stochastic population models to heuristically assess the impact of wolves on pronghorn population dynamics due to changes in neonatal survival. Coyote predation was the primary cause of mortality at all sites, but mortality due to coyotes was 34% lower in areas utilized by wolves (P < 0.001). Based on simulation modeling, the realized population growth rate was 0.92 based on fawn survival in the absence of wolves, and 1.06 at sites utilized by wolves. Thus, wolf restoration is predicted to shift the trajectory of the pronghorn population from a declining to an increasing trend. Our results suggest that reintroductions of large carnivores may influence biodiversity through effects on prey populations mediated by mesopredator suppression. In addition, our approach, which combines empirical data on the population of interest with information from other data sources, demonstrates the utility of using simulation modeling to more fully evaluate ecological theories by moving beyond estimating changes in vital rates to analyses of population-level impacts.     

Response of predators to loss and fragmentation of prey habitat: a review of theory

Despite extensive empirical research and previous reviews, no clear patterns regarding the effects of habitat loss and fragmentation on predator-prey interactions have emerged. We suggest that this is because empirical researchers do not design their studies to test specific hypotheses arising from the theoretical literature. In fact, theoretical work is almost completely ignored by empirical researchers, perhaps because it may be inaccessible to them. The purpose of this paper is to review theoretical work on the effects of habitat loss and fragmentation on predator-prey interactions. We provide a summary of clear, testable theoretical predictions for empirical researchers. To test one or more of these predictions, an empiricist will need certain information on the predator and prey species of interest. This includes: (1) whether the predator is a specialist on one prey species or feeds on many kinds of prey (omnivore and generalist); (2) whether the predator is restricted to the same habitat type as the focal prey (specialist), can use a variety of habitats but has higher survival in the prey habitat (omnivore), or lives primarily outside of the focal prey's habitat (generalist); (3) whether prey-only patches have lower prey extinction rates than predator-prey patches; and (4) whether the prey emigrate at higher rates from predator-prey patches than from prey-only patches. Empiricists also need to be clear on whether they are testing a prediction about habitat loss or habitat fragmentation and need to conduct empirical studies at spatial scales appropriate for testing the theoretical prediction(s). We suggest that appropriate use of the theoretical predictions in future empirical research will resolve the apparent inconsistencies in the empirical literature on this topic.     

Evaluating the impact of caribou habitat restoration on predator and prey movement

Fragmentation of the boreal forest by linear features, including seismic lines, has destabilized predator–prey dynamics, resulting in the decline of woodland caribou (Rangifer tarandus caribou) populations. Restoration of human-altered habitat has therefore been identified as a critical management tool for achieving self-sustaining woodland caribou populations. However, only recently has testing of the response of caribou and other wildlife to restoration activities been conducted. Early work has centered around assessing changes in wildlife use of restored seismic lines. We evaluated whether restoration reduces the movement rates of predators and their associated prey, which is expected to decrease predator hunting efficiency and ultimately reduce caribou mortality. We developed a new method for using cameras to measure fine-scale movement by measuring speed as animals traveled between cameras in an array. We used our method to quantify speed of caribou, moose (Alces alces), bears (Ursus americanus), and wolves (Canis lupus) on treated (restored) and untreated seismic lines. Restoration treatments reduced travel speeds along seismic lines of wolves by 1.38 km/h, bears by 0.55 km/h, and caribou by 1.57 km/h, but did not reduce moose travel speeds. Reduced predator and caribou speeds on treated seismic lines are predicted to decrease encounter rates between predators and caribou and thus lower caribou kill rates. However, further work is needed to determine whether reduced movement rates result in reduced encounter rates with prey, and ultimately reduced caribou mortality.     

The importance of considering multiple interacting species for conservation of species at risk

Conservation of species at risk of extinction is complex and multifaceted. However, mitigation strategies are typically narrow in scope, an artifact of conservation research that is often limited to a single species or stressor. Knowledge of an entire community of strongly interacting species would greatly enhance the comprehensiveness and effectiveness of conservation decisions. We investigated how camera trapping and spatial count models, an extension of spatial-recapture models for unmarked populations, can accomplish this through a case study of threatened boreal woodland caribou (Rangifer tarandus caribou). Population declines in caribou are precipitous and well documented, but recovery strategies focus heavily on control of wolves (Canis lupus) and pay less attention to other known predators and apparent competitors. Obtaining necessary data on multispecies densities has been difficult. We used spatial count models to concurrently estimate densities of caribou, their predators (wolf, black bear [Ursus americanus], and coyote [Canis latrans]), and alternative prey (moose [Alces alces] and white-tailed deer [Odocoileus virginianus]) from a camera-trap array in a highly disturbed landscape within northern Alberta's Oil Sands Region. Median densities were 0.22 caribous (95% Bayesian credible interval [BCI] = 0.08–0.65), 0.77 wolves (95% BCI = 0.26–2.67), 2.39 moose (95% BCI = 0.56–7.00), 2.64 coyotes (95% BCI = 0.45–6.68), and 3.63 black bears (95% BCI = 1.25–8.52) per 100 km². (The white-tailed deer model did not converge.) Although wolf densities were higher than densities recommended for caribou conservation, we suggest the markedly higher black bear and coyote densities may be of greater concern, especially if government wolf control further releases these species. Caribou conservation with a singular focus on wolf control may leave caribou vulnerable to other predators. We recommend a broader focus on the interacting species within a community when conserving species.     

Fine-scale habitat modeling of a top marine predator: do prey data improve predictive capacity?

Predators and prey assort themselves relative to each other, the availability of resources and refuges, and the temporal and spatial scale of their interaction. Predictive models of predator distributions often rely on these relationships by incorporating data on environmental variability and prey availability to determine predator habitat selection patterns. This approach to predictive modeling holds true in marine systems where observations of predators are logistically difficult, emphasizing the need for accurate models. In this paper, we ask whether including prey distribution data in fine-scale predictive models of bottlenose dolphin (Tursiops truncatus) habitat selection in Florida Bay, Florida, U.S.A., improves predictive capacity. Environmental characteristics are often used as predictor variables in habitat models of top marine predators with the assumption that they act as proxies of prey distribution. We examine the validity of this assumption by comparing the response of dolphin distribution and fish catch rates to the same environmental variables. Next, the predictive capacities of four models, with and without prey distribution data, are tested to determine whether dolphin habitat selection can be predicted without recourse to describing the distribution of their prey. The final analysis determines the accuracy of predictive maps of dolphin distribution produced by modeling areas of high fish catch based on significant environmental characteristics. We use spatial analysis and independent data sets to train and test the models. Our results indicate that, due to high habitat heterogeneity and the spatial variability of prey patches, fine-scale models of dolphin habitat selection in coastal habitats will be more successful if environmental variables are used as predictor variables of predator distributions rather than relying on prey data as explanatory variables. However, predictive modeling of prey distribution as the response variable based on environmental variability did produce high predictive performance of dolphin habitat selection, particularly foraging habitat.     

The causes and consequences of partial prey consumption by wolves preying on moose

For a wide range of taxa, partial prey consumption (PPC) is a frequent occurrence. PPC may arise from physiological constraints to gut capacity or digestive rate. Alternatively, PPC may represent an optimal foraging strategy. Assessments that clearly distinguish between these causes are rare and have been conducted only for invertebrate species that are ambush predators with extra-intestinal digestion (e.g., wolf spiders). We present the first strong test for the cause of PPC in a cursorial vertebrate predator with intestinal digestion: wolves (Canis lupus) feeding on moose (Alces alces). Previous theoretical assessments indicate that if PPC represents an optimal foraging strategy and is not caused by physiological limitations, then mean carcass utilization is negatively correlated with mean kill rate and the utilization of individual carcasses is uncorrelated with time between kills. Wolves exhibit exactly this pattern. We explore how the typical portrayal of PPC by wolves has been not only misleading but also detrimental to conservation by promoting negative attitudes toward wolves.     

War Zones and Game Sinks in Lewis and Clark's West

The journals of Lewis and Clark reveal a major difference in the taxa, numbers, and behavior of megafauna on either side of the Rocky Mountains in western North America. Two prior events set the stage for what Lewis and Clark would find. The first was the extinction around 13,000 years ago of two-thirds of the native megafauna of the American West. The second was the effects on Indians of deadly new diseases and new technologies brought by Europeans in the post-Columbian era. Populations of large animals, which were preferred prey for native people, were not immune to European influence. Along the Columbia River corridor west of the Rockies, tens of thousands of people lived in a game sink. Here Lewis and Clark's party found too few animals to live off the land by hunting. They adapted poorly to the local diet of fish and roots offered by the Nez Perce and bought dogs and horses to sustain themselves. To the east, uninhabited lands along the Upper Missouri and the Yellowstone rivers supported an abundance of wild game, especially bison, elk, deer, pronghorn, and wolves. This game source occupied part of a buffer zone of 120,000 km² probed by various Indian war parties, some of them armed with muskets. William Clark recognized the relationship and near the end of their journey he wrote that they found large numbers of large animals in the land between nations that were at war. Both the abundance of game in buffer or war zones and scarcity of big game in sinks have been misinterpreted as a natural or typical condition. Although efforts to restore ecosystems to what is described in early journals may have merit, they are aimed at a flickering target. Long before these journals were written, the land had been stripped of most of its native megafauna through human influence. In the absence of humans, we predict that much larger populations of bison, elk, deer, and wolves would have ranged the West than were reported in historic documents.     

Predator-prey spatial game as a tool to understand the effects of protected areas on harvester-wildlife interactions

No-take reserves are sometimes implemented for sustainable population harvesting because they offer opportunities for animals to spatially avoid harvesters, whereas harvesters can benefit in return from the reserve spillover. Here, we used the framework of predator-prey spatial games to understand how protected areas shape spatial interactions between harvesters and target species and determine animal mortality. In these spatial games, the "predator" searches for "prey" and matches their habitat use, unless it meets spatial constraints offering the opportunity for prey to avoid the mortality source. However, such prey refuges could attract predators in the surroundings, which questions the potential benefits for prey. We located, in the Geneva Basin (France), hunting dogs and wild boar Sus scrofa L. during hunting seasons with global positioning systems and very-high-frequency collars. We quantified how the proximity of the reserve shaped the matching between both habitat uses using multivariate analyses and linked these patterns to animals' mortality with a Cox regression analysis. Results showed that habitat uses by both protagonists disassociated only when hunters were spatially constrained by the reserve. In response, hunters increased hunting efforts near the reserve boundary, which induced a higher risk exposure for animals settled over the reserve. The mortality of adult wild boar decreased near the reserve as the mismatch between both habitat uses increased. However the opposite pattern was determined for younger individuals that suffered from the high level of hunting close to the reserve. The predator-prey analogy was an accurate prediction of how the protected area modified spatial relationships between harvesters and target species. Prey-searching strategies adopted by hunters around reserves strongly impacted animal mortality and the efficiency of the protected area for this harvested species. Increasing reserve sizes and/or implementing buffer areas with harvesting limitations can dampen this edge effect and helps harvesters to benefit durably from source populations of reserves. Predator-prey spatial games therefore provide a powerful theoretical background for understanding wildlife-harvester spatial interactions and developing substantial application for sustainable harvesting.     

How humans shape wolf behavior in Banff and Kootenay National Parks, Canada

This paper describes the conceptualization and implementation of an agent-based model to investigate how varying levels of human presence could affect elements of wolf behavior, including highway crossings; use of areas in proximity to roads and trails; size of home ranges; activities, such as hunting, patrolling, resting, and feeding pups; and survival of individuals in Banff and Kootenay National Parks, Canada. The model consists of a wolf module as the primary component with five packs represented as cognitive agents, and grizzly bear, elk, and human modules that represent dynamic components of the environment. A set of environmental data layers was used to develop a friction model that serves as a base map representing the landscape over which wolves moved. A decision model was built to simulate the sequence of wolf activities. The model was implemented in a Java Programming Language using RePast, an agent-based modeling library. Six months of wolf activities were simulated from April 16 to October 15 (i.e., a season coherent with regard to known wolf behaviors), and calibrated with GPS data from wolf radiocollars (n = 15) deployed from 2002 to 2004. Results showed that the simulated trajectories of wolf movements were correlated with the observed trajectories (Spearman's rho 0.566, P < 0.001); other critical behaviors, such as time spent at the den and not traveling were also correlated. The simulations revealed that wolf movements and behaviors were noticeably affected by the intensity of human presence. The packs’ home ranges shrank and wolves crossed highways less frequently with increased human presence. In an extreme example, a wolf pack whose home range is traversed by a high-traffic-volume highway was extirpated due to inability to hunt successfully under a scenario wherein human presence levels were increased 10-fold. The modeling prototype developed in this study may serve as a tool to test hypotheses about human effects on wolves and on other mammals, and guide decision-makers in designing management strategies that minimize impacts on wolves and on other species functionally related to wolves in the ecosystem.     

Behavioral response races, predator-prey shell games, ecology of fear, and patch use of pumas and their ungulate prey

The predator-prey shell game predicts random movement of prey across the landscape, whereas the behavioral response race and landscape of fear models predict that there should be a negative relationship between the spatial distribution of a predator and its behaviorally active prey. Additionally, prey have imperfect information on the whereabouts of their predator, which the predator should incorporate in its patch use strategy. I used a one-predator-one-prey system, puma (Puma concolor)-mule deer (Odocoileus hemionus) to test the following predictions regarding predator-prey distribution and patch use by the predator. (1) Pumas will spend more time in high prey risk/low prey use habitat types, while deer will spend their time in low-risk habitats. Pumas should (2) select large forage patches more often, (3) remain in large patches longer, and (4) revisit individual large patches more often than individual smaller ones. I tested these predictions with an extensive telemetry data set collected over 16 years in a study area of patchy forested habitat. When active, pumas spent significantly less time in open areas of low intrinsic predation risk than did deer. Pumas used large patches more than expected, revisited individual large patches significantly more often than smaller ones, and stayed significantly longer in larger patches than in smaller ones. The results supported the prediction of a negative relationship in the spatial distribution of a predator and its prey and indicated that the predator is incorporating the prey's imperfect information about its presence. These results indicate a behavioral complexity on the landscape scale that can have far-reaching impacts on predator-prey interactions.     

Behavioral responses of bison and elk in Yellowstone to snowmobiles and snow coaches.

Managers of public lands are charged with protecting some of our most important natural resources and ecosystems, while providing for their use and enjoyment by visitors. Almost one million visitors entered Yellowstone National Park by motorized means on snowmobiles (87%) or snow coaches (13%) during 1992-2003. Most vehicles toured the central portion of the park where bison (Bison bison) and elk (Cervus elaphus) concentrate in geothermal areas. We sampled >6500 interactions between groups of these species and groups of snowmobiles and snow coaches (collectively, OSV, over-snow vehicles) during five winters (1999-2000, 2002-2004). Multinomial logits models were used to identify conditions leading to behavioral responses. Elk responded three times as often (52%) as bison (19%) during interactions with groups of snowmobiles and snow coaches due to increased vigilance responses (elk, 44%; bison, 10%). However, the frequency of higher-intensity movement responses by bison and elk were similar (6-7% travel, 1-2% flight, <1% defense) and relatively low compared to other studies of ungulates and snowmobile disturbance. The likelihood of active responses by bison and elk increased significantly if animals were on or near roads, groups were smaller, or humans approached. The likelihood of an active response by bison decreased within winters having the largest visitation, suggesting some habituation to snowmobiles and snow coaches. There was no evidence that snowmobile use during the past 35 years affected the population dynamics or demography of bison or elk. Thus, we suggest that regulations restricting levels and travel routes of over-snow vehicles (OSVs) were effective at reducing disturbances to bison and elk below a level that would cause measurable fitness effects. We recommend park managers consider maintaining OSV traffic levels at or below those observed during our study. Regardless, differing interpretations of the behavioral and physiological response data will continue to exist because of the diverse values and beliefs of the many constituencies of Yellowstone.