Influence of Landscape Characteristics on Hunter Space Use and Success

Sport hunting of ungulates is a predominant recreational pursuit and the primary tool for managing their populations in North America and beyond, given its influence on ungulate distributions, social organization, and population performance. Similarly, land management, such as motorized vehicle access, influences ungulate distributions during and outside hunting seasons. Although research on ungulate responses to hunting and land use is widespread, knowledge gaps persist about space use of hunters and what landscape features discriminate among hunt types and between successful and unsuccessful hunters. We used telemetry location data from hunters (n = 341) to estimate space use from 2008–2013 during 3 types of controlled, 5-day hunts for antlered mule deer (Odocoileus hemionus) and elk (Cervus canadensis) in northeastern Oregon, USA: archery elk, rifle deer, and rifle elk. To evaluate space use, we developed utilization distributions for each hunter, created core areas (50% contours) for groups of hunters, and derived several metrics of space-use overlap between successful and unsuccessful hunters. We also modeled predictors of space use using resource utilization functions with beta regression and stepwise model building. Hunter space use was compressed, with even the largest core area (unsuccessful rifle elk hunters) encompassing <16% (1,178 ha) of the area. We found strong similarities in space use of rifle hunters compared to archers, and core areas of successful hunters were markedly smaller than those of unsuccessful hunters (e.g., = 104 ha vs. 681 ha, respectively, for archers). Percentage cover and distance from open roads were the most consistent covariates in the 6 final models (successful vs. unsuccessful for each of 3 hunts) but with different signs. For example, predicted use of archery and rifle elk hunters increased with cover but decreased for rifle deer hunters. Although the same covariates were in the final models for unsuccessful and successful rifle elk hunters, their negligible spatial overlap suggested they sought those features in different locales, a pattern also documented for rifle deer hunters. Our models performed well (Spearman's rank correlation coefficients = 0.99 for 5 of 6 models), reflecting their utility for managing hunters and landscapes. Our results suggest that strategic management of open roads and forest cover can benefit managers seeking to balance hunter opportunity and satisfaction with harvest objectives, especially for species of special concern such as mule deer, and that differences in space use among hunter groups should be accounted for in hunting season designs. © 2021 The Wildlife Society. This article has been contributed to by US Government employees and their work is in the public domain in the USA.     

Effects of hunter access and habitat security on elk habitat selection in landscapes with a public and private land matrix

Traditional elk habitat management on public land has focused on providing security habitat for bull elk during the hunting season to provide for both adequate hunter opportunity and bull survival. This paradigm has given less consideration to adult female elk habitat use, patterns of adjacent land ownership, and hunter access. This paradigm also was developed when elk population sizes were much smaller in many areas. In many Rocky Mountain states, the focus of elk population management has recently shifted to reducing or maintaining elk population sizes, necessitating a better understanding of the implications of security habitat management, as well as patterns of adjacent land ownership and hunter access, on adult female elk. We addressed this need by testing the hypotheses that during the hunting season: 1) adult female elk selection for areas prohibiting or limiting hunter access is stronger than elk selection for publicly owned and managed elk security habitat, 2) these effects occur during the archery hunting period and intensify during the rifle hunting period, and 3) the effects of hunter access on selection are consistent among herds that occupy landscapes characterized by a matrix of public and private lands. We used global position system locations collected from 82 females in 2 different Greater Yellowstone Ecosystem (GYE) elk herds to evaluate effects of hunter access, security habitat as defined by the Hillis paradigm, and other landscape attributes on adult female elk resource selection during the pre-hunting, archery, rifle, and post-hunting periods. We found that female elk selection for areas restricting public hunting access was stronger than selection for security habitat in both study areas, and that the density of roads open to motorized use was the strongest predictor of elk distribution. Increases in selection for areas that restricted hunting access occurred during the rifle hunting period, and we did not find consistent evidence these movements were triggered by the archery hunting period. Our results provide evidence that in landscapes characterized by a matrix of public and privately owned lands, traditional concepts of elk security habitat need to be expanded to also include areas that restrict hunter access to plan for elk population management that is regulated through adult female harvest. Future efforts should investigate whether elk use of areas that restrict hunter access are flexible behavioral responses to hunting risk, or if these behaviors are passed from generation to generation such that a learned pattern of private land use becomes the normal movement pattern rather than a short-term behavioral response. Published 2013. This article is a U.S. Government work and is in the public domain in the USA.     

Hunting‐mediated predator facilitation and superadditive mortality in a European ungulate

Predator‐prey theory predicts that in the presence of multiple types of predators using a common prey, predator facilitation may result as a consequence of contrasting prey defense mechanisms, where reducing the risk from one predator increases the risk from the other. While predator facilitation is well established in natural predator‐prey systems, little attention has been paid to situations where human hunters compete with natural predators for the same prey. Here, we investigate hunting‐mediated predator facilitation in a hunter‐predator‐prey system. We found that hunter avoidance by roe deer (Capreolus capreolus) exposed them to increase predation risk by Eurasian lynx (Lynx lynx). Lynx responded by increasing their activity and predation on deer, providing evidence that superadditive hunting mortality may be occurring through predator facilitation. Our results reveal a new pathway through which human hunters, in their role as top predators, may affect species interactions at lower trophic levels and thus drive ecosystem processes.     

‘Impact hunters’ catalyse cooperative hunting in two wild chimpanzee communities

Even when hunting in groups is mutually beneficial, it is unclear how communal hunts are initiated. If it is costly to be the only hunter, individuals should be reluctant to hunt unless others already are. We used 70 years of data from three communities to examine how male chimpanzees ‘solve’ this apparent collective action problem. The ‘impact hunter’ hypothesis proposes that group hunts are sometimes catalysed by certain individuals that hunt more readily than others. In two communities (Kasekela and Kanyawara), we identified a total of five males that exhibited high hunt participation rates for their age, and whose presence at an encounter with red colobus monkeys increased group hunting probability. Critically, these impact hunters were observed to hunt first more often than expected by chance. We argue that by hunting first, these males dilute prey defences and create opportunities for previously reluctant participants. This by-product mutualism can explain variation in group hunting rates within and between social groups. Hunting rates declined after the death of impact hunter FG in Kasekela and after impact hunter MS stopped hunting frequently in Kanyawara. There were no impact hunters in the third, smaller community (Mitumba), where, unlike the others, hunting probability increased with the number of females present at an encounter with prey.     

Behavioural Responses of European Roe Deer to Temporal Variation in Predation Risk

In natural environments, predation risk varies over time. The risk allocation hypothesis predicts that prey is expected to adjust key anti‐predator behaviours such as vigilance to temporal variation in risk. We tested the predictions of the risk allocation hypothesis in a natural environment where both a species‐rich natural predator community and human hunters are abundant and where the differences in seasonal and circadian activity between natural and anthropogenic predators provided a unique opportunity to quantify the contributions of different predator classes to anti‐predator behaviour. Whereas natural predators were expected to show similar levels of activity throughout the seasons, hunter activity was high during the daytime during a clearly defined hunting season. According to the risk allocation hypothesis, vigilance should then be higher during the hunting season and during daytime hours than during the non‐hunting season and night‐time hours. Roe deer (Capreolus capreolus) on the edge of Bia?owie?a Primeval Forest in Eastern Poland displayed vigilance behaviour consistent with these predictions. The behavioural response of roe deer to temporarily varying predation risks emphasises the behavioural plasticity of this species and suggests that future studies of anti‐predator behaviour need to incorporate circadian variation in predation pressure as well as risk gradients of both natural and anthropogenic predators.     

Demographic influences on the hunting behavior of chimpanzees

We investigated hunting in an unusually large community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Aspects of predation were recorded with respect to the prey, the predators, and hunting episodes. During 23 months of observation, the Ngogo chimpanzees caught 128 prey items from four primate and three ungulate species. Chimpanzees preyed selectively on immature red colobus primarily during group hunts, with adult males making the majority of kills. Party size and composition were significant predictors of the probability that chimpanzees would hunt and of their success during attempts. Chimpanzees were more likely to hunt red colobus if party size and the number of male hunters were large; party size and the number of male hunters were also significantly larger in successful compared with unsuccessful hunts. The Ngogo chimpanzees did not appear to hunt cooperatively, but reciprocal meat-sharing typically took place after kills. Hunts occurred throughout the year, though there was some seasonality as displayed by periodic hunting binges. The extremely high success rate and large number of kills made per successful hunt are the two most striking aspects of predation by the Ngogo chimpanzees. We compare currently available observations of chimpanzee hunting behavior across study sites and conclude that the large size of the Ngogo community contributes to their extraordinary hunting success. Demographic differences between groups are likely to contribute to other patterns of interpopulation variation in chimpanzee predation. Am J Phys Anthropol 109:439–454, 1999. © 1999 Wiley-Liss, Inc.     

Evaluating Indirect Effects of Hunting on Mule Deer Spatial Behavior

Mule deer (Odocoileus hemionus) are widely hunted throughout western North America and are experiencing population declines across much of their range. Consequently, understanding the direct and indirect effects of hunting is important for management of mule deer populations. Managers can influence deer mortality rates through changes in hunting season length or authorized tag numbers. Little is known, however, about how hunting can affect site fidelity patterns and subsequent habitat use and movement patterns of mule deer. Understanding these patterns is especially important for adult females because changes in behavior may influence their ability to acquire resources and ultimately affect their productivity. Between 2008 and 2013, we obtained global positioning system locations for 42 adult female deer at the Starkey Experimental Forest and Range in northeast Oregon, USA, during 5-day control and treatment periods in which hunters were absent (pre-hunt), present but not actively hunting (scout and post-hunt), and actively hunting male mule deer (hunt) on the landscape. We estimated summer home ranges and 5-day use areas during pre-hunt and hunt periods and calculated overlap metrics across home ranges and use areas to assess site fidelity within and across years. We used step selection functions to evaluate whether female mule deer responded to human hunters by adjusting fine-scale habitat selection and movement patterns during the hunting season compared to the pre-hunt period. Mule deer maintained site fidelity despite disturbance by hunters with 72 ± 4% (SE) within-year overlap between summer home ranges and hunt use areas and 54 ± 7% inter-annual overlap among pre-hunt use areas and 56 ± 7% among hunt use areas. Mule deer diurnal movement rates, when hunters are active on the landscape, were higher during the hunting period versus pre-hunt or scout periods. In contrast, nocturnal movement rates, when hunters are inactive on the landscape, were similar between hunting and non-hunting periods. Additionally, during the hunt, female mule deer hourly movements increased in areas with high greenness values, indicating that mule deer spent less time in areas with more vegetative productivity. Female mule deer maintained consistent habitat selection patterns before and during hunts, selecting areas that offered more forest canopy cover and high levels of vegetative productivity. Our results indicate that deer at Starkey are adopting behavioral strategies in response to hunters by increasing their movement rates and selecting habitat in well-established ranges. Therefore, considering site fidelity behavior in management planning could provide important information about the spatial behavior of animals and potential energetic costs incurred, especially by non-target animals during hunting season. © 2020 The Wildlife Society.     

Do prey select for vacant hunting domains to minimize a multi‐predator threat?

Many ecosystems contain sympatric predator species that hunt in different places and times. We tested whether this provides vacant hunting domains, places and times where and when predators are least active, that prey use to minimize threats from multiple predators simultaneously. We measured how northern Yellowstone elk (Cervus elaphus) responded to wolves (Canis lupus) and cougars (Puma concolor), and found that elk selected for areas outside the high‐risk domains of both predators consistent with the vacant domain hypothesis. This enabled elk to avoid one predator without necessarily increasing its exposure to the other. Our results demonstrate how the diel cycle can serve as a key axis of the predator hunting domain that prey exploit to manage predation risk from multiple sources. We argue that a multi‐predator, spatiotemporal framework is vital to understand the causes and consequences of prey spatial response to predation risk in environments with more than one predator.     

Autumn resource selection by female elk in a recently burned landscape in western Montana

Wildfire activity across the western United States has increased in recent decades, with wildfires burning at a higher severity and larger scale. The effect of wildfires on forest structure and wildlife habitat is largely influenced by wildfire severity; however, few studies have evaluated the effects of wildfire severity on resource selection of ungulates, particularly during hunting seasons, when knowledge of resource selection is essential for making informed management decisions. To fill this knowledge gap, we fit resource selection probability functions for female elk (Cervus canadensis) in years 2 and 3 post-wildfire to evaluate the effects of wildfire severity and other environmental and anthropogenic factors on elk resource selection during 4 autumn periods with varying levels of hunter pressure (prehunt, archery-only, backcountry rifle, and rifle). The probability of female elk selecting low-severity burned forests during the prehunt, archery-only, backcountry rifle, and rifle periods was 0.99 (95% credible interval [CrI] = 0.98–1.00), 0.99 (CrI = 0.97–1.00), 0.99 (CrI = 0.99–1.00), and 0.0010 (CrI = 0.00067–0.0015]), respectively, and did not strongly differ from the probability of selecting high-severity burned forests. During the prehunt period, elk also selected areas with greater forage quality and areas farther from open roads. Elk selected similar resources during the archery period, and selected areas with higher hunter pressure. Elk started leaving hunting districts that had higher snowpack (i.e., snow water equivalent; β = −0.84, CrI = −0.96–−0.72) and allowed rifle hunting (β = −5.39, CrI = −5.80–−4.97) but still selected areas with higher hunter pressure (β = 0.92, CrI = 0.78–1.07) during the backcountry rifle period. During the rifle period, elk continued avoiding areas with high snowpack (β = −3.96, CrI = −4.22–−3.71) and started selecting areas with lower hunter pressure (β = −1.71, CrI = −1.79–−1.64) and lower canopy cover. Overall, wildfire affected elk distributions in early autumn 2 and 3 years after fire in our study area, with limited differences in resource selection between wildfire severity categories. By late autumn, hunter pressure and snowpack were the primary factors influencing elk distribution, and wildfire had little influence on selection. When estimating wildfire effects on elk movements during autumn and establishing appropriate hunting regulations, managers should consider the hunting season, hunter pressure, timing and amount of snowpack, location of traditional winter range, and the seasonal elk range burned, as all these factors may contribute to how elk use the landscape in autumn.     

Bark Beetle-Affected Forests Provide Elk Only a Marginal Refuge from Hunters

For nearly 2 decades, the forests of the Rocky Mountains in the United States experienced a bark-beetle (Dendoctronus ponderosae) epidemic. The number of dead and falling trees from this epidemic likely will affect how elk (Cervus canadensis) and hunters use the forest and their interactions. Downed trees potentially create a component of refuge habitat that could affect the effectiveness of hunting to regulate abundance of growing elk populations. We evaluated how forests affected by bark beetles in south-central Wyoming, USA, influenced resource selection of 50 female elk and 374 hunters in 2012–2016, and interactions between elk and hunters. We employed global positioning system (GPS) technology on elk and hunters, and developed a satellite-derived land classification specifically depicting beetle-affected forests. We tested the predictions that elk would increase use of beetle-killed areas from summer to the hunting season, and that hunters would avoid beetle-killed areas regardless of elk use. Elk increased use of beetle-killed areas during hunting seasons as did hunters during the archery season; however, during the rifle season, hunters avoided beetle-killed areas. Nevertheless, during the rifle season, areas of beetle-kill with a high probability of elk occurrence dampened the aversion hunters had towards beetle-killed areas. Therefore, in contrast to our expectations, forests that have been altered by the bark-beetle epidemic may only function marginally as a refuge for elk. Our study area was at the beginning of the tree-fall phase (i.e., ~3–7 yr after peak infestation) of the bark-beetle epidemic; thus, future research efforts should focus on how a continued increase in downed trees will influence interactions between elk and hunters. At current levels of tree fall, however, beetle-kill should not limit the ability of managers to regulate elk herds through harvest. © 2020 The Wildlife Society.     

Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators

Predator avoidance depends on prey being able to discern how risk varies in space and time, but this is made considerably more complicated if risk is simultaneously present from multiple predators. This is the situation for an increasing number of mammalian prey species, as large carnivores recover or are reintroduced in ecosystems on several continents. Roe deer Capreolus capreolus in southern Norway illustrate a case in which prey face two predators with contrasting patterns of predation risk. They face a catch‐22 situation: spatially avoiding the risk from one predator (lynx Lynx lynx in dense habitat) implies exposure to the other (hunters in open habitat). Using GPS‐data from 29 roe deer, we tested for daily and seasonal variation in roe deer selection for habitat with respect to the habitats’ year‐round average risk level. Generally, roe deer altered their habitat selection between night and day in a pattern consistent with being able to avoid predicted risk from the nocturnal lynx during night and predicted risk from human hunters during day. However, seasonal variation in habitat selection only partially corresponded with the predicted seasonal variation in risk. Whereas roe deer avoided areas with high risk from hunters more strongly during the hunting season than in other seasons, there was a lack of selection towards areas and time periods lowering the risk of lynx predation during winter. It seems likely that the risk of starvation and thermal stress constrain roe deer habitat selection during this energetically challenging season with cold temperatures, snow cover and limited natural forage. The habitat selection pattern of roe deer fits thus only partly with the two contrasting risk gradients they face. Adjusting risk‐avoidance behavior temporally can be an adaptive response in the case of several predators whose predation patterns differ in space and time.     

The economic value of publicly accessible deer hunting land

Access to lands for hunting is widely recognized as critical to sustaining and developing participation in hunting. Deer hunters comprise a substantial portion of recreational hunters in the United States, and numerous stakeholders benefit from the deer management services provided by hunters on public lands. We estimated the economic value of publicly accessible deer hunting land in Michigan using economic models of hunting destinations for hunters in the archery season and for hunters in the firearm season. We found that the amounts of state-owned hunting land, federally owned hunting land, and publicly accessible, privately owned Commercial Forest Act land are significant (P < 0.001) and positive determinants of hunting destinations. The annual economic value of Michigan's publicly accessible hunting land to archery and firearm deer hunters, beyond hunter expenditures, was estimated to be over $80 million. Access to state-owned land accounted for approximately $50 million in annual economic value to Michigan deer hunters. The average per-acre value of publicly accessible hunting land is greatest in the Southern Lower Peninsula of Michigan, where public access is scarce and the majority of hunters reside. Understanding the economic values hunters derive from publicly accessible hunting lands will enable managers and policy makers to make better decisions when considering policy options that involve reducing or increasing public access. © 2011 The Wildlife Society.     

Influence of habitat features and hunter behavior on white-tailed deer harvest

Sport hunting may help in controlling cervid populations over large areas. As with natural predators, several environmental factors can influence sport harvest. A better understanding of the environmental variables that limit the efficiency of sport hunting could provide guidelines for more efficient wildlife management using hunting. We studied white-tailed deer (Odocoileus virginianus) hunting on a high deer density island where hunting was the sole form of predation. Our objective was to study the behavior of sport hunters and determine the habitat characteristics (e.g., abundance of deer forage, visibility of the deer from the hunter's point of view, and accessibility of the territory to hunters) that are associated with a successful harvest. We collected movements and harvest site location data from 477 hunters equipped with handheld Global Positioning System (GPS) units. Harvest sites were visited and characterized, along with a paired random site, to determine the environmental conditions associated with a successful hunt. We also developed a model to predict the daily number of deer seen by hunters considering weather conditions, hunter characteristics (e.g., age, experience), and date of hunting. We used the mean number of deer seen per hunter per day as a relative index of local density in each hunted territory. At both the site and landscape scales, the combination of visibility and access had a positive effect on the distribution of harvested deer. Habitat types with less visual obstruction from vegetation enabled hunters to see more deer in a given day. At the site scale, harvested deer were located in areas with a lower density of access routes compared to areas where hunters travelled throughout the day. Using an innovative approach of studying hunter behavior with GPS technology, digital maps, and questionnaires, we highlighted the factors associated with hunter success. Our study suggests that habitat characteristics could be modified to increase harvest by improving accessibility and visibility near roads. Creating openings in mature and regenerating forest near access roads could make sport hunting a more efficient management tool, but the potential impact of increased forage availability in forest openings should not be overlooked. © 2012 The Wildlife Society.     

Reduction and Maintenance of a White-Tailed Deer Herd in Central Massachusetts

Abstract: Controlled public hunts have been used in a variety of settings to reduce overabundant white-tailed deer (Odocoileus virginianus) herds. We present the results of a large-scale (160 km²) controlled hunt at Quabbin Reservation (QR) in central Massachusetts, USA. The QR was divided into 5 hunt zones. Hunting was initiated in each zone from 1991 to 1994 and continued through 2004. The management goal was to achieve posthunt deer densities of4 deer/km². Initial estimated deer densities in each zone ranged from 11.4 deer/km² to 27.6 deer/km². The management goals were reached in each zone after 2-4 years of hunting. Posthunt populations were maintained at or below the goal even though total hunter effort was reduced. Hunters were not required to harvest antlerless deer, but antlerless deer comprised 55-83% of the harvest each year. We simulated the effects of 5 years without hunting on deer populations. The simulated deer population exceeded management goals after 2 years. Our results demonstrate that controlled public deer hunts can effectively reduce deer populations and maintain them at desired levels over large areas with minimal hunter restrictions. Managers should prepare stakeholders during the hunt planning process for the need to continue overall harvest rates of >30% during the maintenance phase of a deer management program.     

Big game hunting in New Zealand: per capita effort,harvest and expenditure in 2011–2012

Recreational big game hunters make a significant contribution to conservation through kills of deer, pigs, chamois and tahr. New opportunities for managing recreational hunting through the proposed Game Animal Council underscore the need to understand the implications of potential changes in recreational hunting participation and harvests. Based on a survey of hunters' recall over a year, hunters averaged 15.63 (SEM = 0.58) big game hunts per year, spending 30.53 (SEM = 0.85) days hunting and killing 8.92 (SEM = 0.69) big game animals. Hunters commonly targeted several species on a single hunt, with highly skewed distributions for hunter effort and kills. Mean monthly expenditure on big game hunting items was $296.78 (SEM = $8.95). Results demonstrate that big game hunting is a significant activity in New Zealand, but this varies considerably among hunters with a small number responsible for the vast majority of kills. These are important considerations for future big game hunting management.     

Drivers of Bushmeat Hunting and Perceptions of Zoonoses in Nigerian Hunting Communities

Bushmeat hunting threatens biodiversity and increases the risk of zoonotic pathogen transmission. Nevertheless, limited information exists on patterns of contact with wildlife in communities that practice bushmeat hunting, especially with respect to social drivers of hunting behavior. We used interview responses from hunters and non-hunters in rural hunting communities in Nigeria to: 1) quantify contact rates with wildlife, 2) identify specific hunting behaviors that increase frequency of contact, 3) identify socioeconomic factors that predispose individuals to hunt, and 4) measure perceptions of risk. Participants engaged in a variety of behaviors that increased contact with wild animals, including: butchering to sell (37%), being injured (14%), using body parts for traditional medicine (19%), collecting carcasses found in forests and/or farms (18%), and keeping as pets (16%). Hunters came into contact with wildlife significantly more than non-hunters, even through non-hunting exposure pathways. Participants reported hunting rodents (95%), ungulates (93%), carnivores (93%), primates (87%), and bats (42%), among other prey. Reported hunting frequencies within taxonomic groups of prey were different for different hunting behaviors. Young age, lower education level, larger household size, having a father who hunts, and cultural group were all associated with becoming a hunter. Fifty-five percent of respondents were aware that they could contract diseases from wild animals, but only 26% of these individuals reported taking protective measures. Overall, hunters in this setting frequently contact a diversity of prey in risky ways, and the decision to become a hunter stems from family tradition, modified by economic necessity. Conservation and public health interventions in such settings may be most efficient when they capitalize on local knowledge and target root socio-economic and cultural drivers that lead to hunting behavior. Importantly, interventions that target consumption alone will not be sufficient; other drivers and modes of interaction with wildlife must also be considered.     

Living and dying in a multi‐predator landscape of fear: roe deer are squeezed by contrasting pattern of predation risk imposed by lynx and humans

The theory of predation risk effects predicts behavioral responses in prey when risk of predation is not homogenous in space and time. Prey species are often faced with a tradeoff between food and safety in situations where food availability and predation risk peak in the same habitat type. Determining the optimal strategy becomes more complex if predators with different hunting mode create contrasting landscapes of risk, but this has rarely been documented in vertebrates. Roe deer in southeastern Norway face predation risk from lynx, as well as hunting by humans. These two predators differ greatly in their hunting methods. The predation risk from lynx, an efficient stalk‐and‐ambush predator is expected to be higher in areas with dense understory vegetation, while predation risk from human hunters is expected to be higher where visual sight lines are longer. Based on field observations and airborne LiDAR data from 71 lynx predation sites, 53 human hunting sites, 132 locations from 15 GPS‐marked roe deer, and 36 roe deer pellet locations from a regional survey, we investigated how predation risk was related to terrain attributes and vegetation classes/structure. As predicted, we found that increasing cover resulted in a contrasting lower predation risk from humans and higher predation risk from lynx. Greater terrain ruggedness increased the predation risk from both predators. Hence, multiple predators may create areas of contrasting risk as well as double risk in the same landscape. Our study highlights the complexity of predator–prey relationship in a multiple predator setting. Synthesis In this study of risk effects in a multi‐predator context, LiDAR data were used to quantify cover in the habitat and relate it to vulnerability to predation in a boreal forest. We found that lynx and human hunters superimpose generally contrasting landscapes of fear on a common prey species, but also identified double‐risk zones. Since the benefit of anti‐predator responses depends on the combined risk from all predators, it is necessary to consider complete predator assemblages to understand the potential for and occurrence of risk effects across study systems.     

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

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

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

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

Underestimating the frequency,strength and cost of antipredator responses with data from GPS collars: an example with wolves and elk

Field studies that rely on fixes from GPS‐collared predators to identify encounters with prey will often underestimate the frequency and strength of antipredator responses. These underestimation biases have several mechanistic causes. (1) Step bias: The distance between successive GPS fixes can be large, and encounters that occur during these intervals go undetected. This bias will generally be strongest for cursorial hunters that can rapidly cover large distances (e.g., wolves and African wild dogs) and when the interval between GPS fixes is long relative to the duration of a hunt. Step bias is amplified as the path travelled between successive GPS fixes deviates from a straight line. (2) Scatter bias: Only a small fraction of the predators in a population typically carry GPS collars, and prey encounters with uncollared predators go undetected unless a collared group‐mate is present. This bias will generally be stronger for fission–fusion hunters (e.g., spotted hyenas, wolves, and lions) than for highly cohesive hunters (e.g., African wild dogs), particularly when their group sizes are large. Step bias and scatter bias both cause underestimation of the frequency of antipredator responses. (3) Strength bias: Observations of prey in the absence of GPS fix from a collared predator will generally include a mixture of cases in which predators were truly absent and cases in which predators were present but not detected, which causes underestimation of the strength of antipredator responses. We quantified these biases with data from wolves and African wild dogs and found that fixes from GPS collars at 3‐h intervals underestimated the frequency and strength of antipredator responses by a factor >10. We reexamined the results of a recent study of the nonconsumptive effects of wolves on elk in light of these results and confirmed that predation risk has strong effects on elk dynamics by reducing the pregnancy rate.