Are migrant and resident elk (Cervus elaphus) exposed to similar forage and predation risk on their sympatric winter range?

Partially migratory populations, where one portion of a population conducts seasonal migrations (migrants) while the other remains on a single range (residents), are common in ungulates. Studies that assess trade-offs between migratory strategies typically compare the amount of predation risk and forage resources migrants and residents are exposed to only while on separate ranges and assume both groups intermix completely while on sympatric ranges. Here we provide one of the first tests of this assumption by comparing the amount of overlap between home ranges of GPS-collared migrant and resident elk and fine-scale exposure to wolf predation risk and forage biomass at telemetry locations on a sympatric winter range in west-central Alberta, Canada. Overlap between migrant and resident home ranges increased throughout the winter, and both groups were generally intermixed and exposed to equal forage biomass. During the day, both migrants and residents avoided predation risk by remaining in areas far from timber with high human activity, which wolves avoided. However, at night wolves moved onto the grasslands close to humans and away from timber. Resident elk were consistently closer to areas of human activity and further from timber than migrants, possibly because of a habituation to humans. As a result, resident elk were exposed to higher night-time predation risk than migrants. Our study does not support the assumption that migrant and resident elk are exposed to equal predation risk on their sympatric range when human presence alters predation risk dynamics and habituation to humans is unequal between migratory strategies.     

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.     

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.     

Beyond the encounter: Predicting multi‐predator risk to elk (Cervus canadensis) in summer using predator scats

1. There is growing evidence that prey perceive the risk of predation and alter their behavior in response, resulting in changes in spatial distribution and potential fitness consequences. Previous approaches to mapping predation risk across a landscape quantify predator space use to estimate potential predator‐prey encounters, yet this approach does not account for successful predator attack resulting in prey mortality. An exception is a prey kill site that reflects an encounter resulting in mortality, but obtaining information on kill sites is expensive and requires time to accumulate adequate sample sizes.
2. We illustrate an alternative approach using predator scat locations and their contents to quantify spatial predation risk for elk (Cervus canadensis) from multiple predators in the Rocky Mountains of Alberta, Canada. We surveyed over 1300 km to detect scats of bears (Ursus arctos/U. americanus), cougars (Puma concolor), coyotes (Canis latrans), and wolves (C. lupus). To derive spatial predation risk, we combined predictions of scat‐based resource selection functions (RSFs) weighted by predator abundance with predictions that a predator‐specific scat in a location contained elk. We evaluated the scat‐based predictions of predation risk by correlating them to predictions based on elk kill sites. We also compared scat‐based predation risk on summer ranges of elk following three migratory tactics for consistency with telemetry‐based metrics of predation risk and cause‐specific mortality of elk.
3. We found a strong correlation between the scat‐based approach presented here and predation risk predicted by kill sites and (r = .98, p < .001). Elk migrating east of the Ya Ha Tinda winter range were exposed to the highest predation risk from cougars, resident elk summering on the Ya Ha Tinda winter range were exposed to the highest predation risk from wolves and coyotes, and elk migrating west to summer in Banff National Park were exposed to highest risk of encountering bears, but it was less likely to find elk in bear scats than in other areas. These patterns were consistent with previous estimates of spatial risk based on telemetry of collared predators and recent cause‐specific mortality patterns in elk.
4. A scat‐based approach can provide a cost‐efficient alternative to kill sites of quantifying broad‐scale, spatial patterns in risk of predation for prey particularly in multiple predator species systems.

     

Multiscale wolf predation risk for elk: does migration reduce risk?

While migration is hypothesized to reduce predation risk for ungulates, there have been few direct empirical tests of this hypothesis. Furthermore, few studies examined multiscale predation risk avoidance by migrant ungulates, yet recent research reveals that predator–prey interactions occur at multiple scales. We test the predation risk reduction hypothesis at two spatial scales in a partially migratory elk (Cervus elaphus) population by comparing exposure of migrant and resident elk to wolf (Canis lupus) predation risk. We used GPS and VHF telemetry data collected from 67 migrant and 44 resident elk over the summers of 2002–2004 in and adjacent to Banff National Park (BNP), Canada. We used wolf GPS and VHF telemetry data to estimate predation risk as a function of the relative probability of wolf occurrence weighted by a spatial density model that adjusted for varying pack sizes. We validated the predation risk model using independent data on wolf-killed elk, and showed that combining wolf presence and spatial density best predicted where an elk was likely to be killed. Predation risk on summer ranges of migrant elk was reduced by 70% compared to within resident elk summer ranges. Because wolves avoided areas near high human activity, however, fine-scale selection by resident elk for areas near high human activity reduced their predation risk exposure to only 15% higher than migrants, a difference significant in only one of three summers. Finally, during actual migration, elk were exposed to 1.7 times more predation risk than residents, even though migration was rapid. Our results support the hypothesis that large-scale migrations can reduce predation. However, we also show that where small-scale spatial variation in predation risk exists, nonmigratory elk may equally reduce predation risk as effectively as migrants under some circumstances.     

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

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

Ecosystem Scale Declines in Elk Recruitment and Population Growth with Wolf Colonization: A Before-After-Control-Impact Approach

The reintroduction of wolves (Canis lupus) to Yellowstone provided the unusual opportunity for a quasi-experimental test of the effects of wolf predation on their primary prey (elk – Cervus elaphus) in a system where top-down, bottom-up, and abiotic forces on prey population dynamics were closely and consistently monitored before and after reintroduction. Here, we examined data from 33 years for 12 elk population segments spread across southwestern Montana and northwestern Wyoming in a large scale before-after-control-impact analysis of the effects of wolves on elk recruitment and population dynamics. Recruitment, as measured by the midwinter juvenile∶female ratio, was a strong determinant of elk dynamics, and declined by 35% in elk herds colonized by wolves as annual population growth shifted from increasing to decreasing. Negative effects of population density and winter severity on recruitment, long recognized as important for elk dynamics, were detected in uncolonized elk herds and in wolf-colonized elk herds prior to wolf colonization, but not after wolf colonization. Growing season precipitation and harvest had no detectable effect on recruitment in either wolf treatment or colonization period, although harvest rates of juveniles∶females declined by 37% in wolf-colonized herds. Even if it is assumed that mortality due to predation is completely additive, liberal estimates of wolf predation rates on juvenile elk could explain no more than 52% of the total decline in juvenile∶female ratios in wolf-colonized herds, after accounting for the effects of other limiting factors. Collectively, these long-term, large-scale patterns align well with prior studies that have reported substantial decrease in elk numbers immediately after wolf recolonization, relatively weak additive effects of direct wolf predation on elk survival, and decreased reproduction and recruitment with exposure to predation risk from wolves.     

A Review of Environmental Factors Affecting Elk Winter Diets

Abstract: Decades of research have produced substantial data on elk (Cervus elaphus) diets in winter, when foraging conditions are most likely to affect population dynamics. Using data from 72 studies conducted in western North America between 1938 and 2002, we collated data on elk diets and environmental variables. We used these data to quantify diet selection by elk and to test whether variation in elk diets is associated with habitat type, winter severity, period of winter, human hunting, and study method. Graminoids (grasses and grass-like plants such as sedges) dominated elk diets and consistently occurred at a higher proportion in the diet than in elk foraging habitats, indicating preference. Forbs commonly made up ≤5% of the diet, with no evidence for preference; we conclude that forb use is largely incidental to grazing for graminoids. Browse was consumed in proportion to its availability, implying that the amount of browse in the diet was primarily determined by habitat use rather than selection. Comparing the diets of elk and sympatric ruminants, elk consistently selected graminoids more strongly than sympatric ruminants with the exception of bison (Bison bison), suggesting that elk are not environmentally forced to adopt the graminoid-biased diet that they normally select. The proportion of open meadows and grasslands on winter ranges was strongly and positively associated with graminoid consumption by elk. The proportion of graminoids in the diet was significantly lower in elk experiencing severe winter conditions or predation risk from human hunting. The period of winter (early, middle, and late) had only small effects on elk diets, as did the method by which the diet was determined. Overall, variation in elk diets is well-explained by a consistent tendency to select graminoids if available, modified by winter habitat type, predation risk, and winter severity, which can constrain habitat selection and access to grazing opportunities. To fully understand variation in foraging behavior, biologists should recognize these broad patterns when interpreting resource selection data. Managers should recognize that inconspicuous behavioral responses to environmental stimuli can alter the diet in ways that probably carry nutritional consequences.     

Testing the risk of predation hypothesis: the influence of recolonizing wolves on habitat use by moose

Considered as absent throughout Scandinavia for >100 years, wolves (Canis lupus) have recently naturally recolonized south-central Sweden. This recolonization has provided an opportunity to study behavioral responses of moose (Alces alces) to wolves. We used satellite telemetry locations from collared moose and wolves to determine whether moose habitat use was affected by predation risk based on wolf use distributions. Moose habitat use was influenced by reproductive status and time of day and showed a different selection pattern between winter and summer, but there was weak evidence that moose habitat use depended on predation risk. The seemingly weak response may have several underlying explanations that are not mutually exclusive from the long term absence of non-human predation pressure: intensive harvest by humans during the last century is more important than wolf predation as an influence on moose behavior; moose have not adapted to recolonizing wolves; and responses may include other behavioral adaptations or occur at finer temporal and spatial levels than investigated.     

Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

Behavioral responses of male elk to hunting risk

Prey respond to predation risk with a range of behavioral tactics that can vary based on space use and hunting mode of the predator. Unlike other predators, human hunters are often more spatially and temporally restricted, which creates a period of short-duration, high-intensity predation risk for prey. Consequently, identifying the roles different hunting modes (i.e., archery and rifle), hunts for targeted and non-targeted species, and landscape features play in altering spatial and temporal responses of prey to predation risk by humans is important for effective management of harvested populations. From 2009 to 2016, we used a large-scale experiment including 50 animal-years of location data from 38 unique male elk (Cervus canadensis) to quantify changes in movement and resource selection in response to hunters during 3 separate 5-day controlled hunts for antlered males (elk archery, deer [Odocoileus spp.] rifle, and elk rifle) at the Starkey Experimental Forest and Range in northeast Oregon, USA. We evaluated competing hypotheses regarding elk responses to varying levels of prey risk posed by the different hunt types. We predicted that the strength of elk behavioral responses would increase with perceived hunter lethality (i.e., weak response to elk archery but similar response to elk and deer rifle hunts) and that prey response would be closely associated with hunter activity within the diel cycle (greater during diurnal than nocturnal hours) and across hunting seasons. Elk responses were strongest during diurnal hours when hunters were active on the landscape and were generally more pronounced during both rifle hunts than during the archery hunt (supporting our perceived lethality hypothesis). Male elk avoided open roads across all periods except during nocturnal hours of the breeding season and alternated between avoidance of areas with high canopy cover during nocturnal hours and selection during diurnal hours. In combination these patterns led to distinct distributional changes of male elk from pre-hunt to hunt periods. Patterns of male elk selection highlight the importance of managing for heterogeneous landscapes to meet a variety of habitat, harvest, hunter satisfaction, and escapement objectives.     

Winter predation patterns of wolves in Northwestern Wyoming

Wolf (Canis lupus) diets and potential effects on prey have been a prominent subject of interest to wildlife researchers and managers since reintroduction into Yellowstone National Park, Wyoming, USA, in 1995 and 1996. Post-reintroduction, wolves expanded south and recolonized areas in the southern Yellowstone ecosystem. Elk (Cervus elaphus) in this area are supplementally fed during winter (Dec–Mar) at state-managed feedgrounds, resulting in high-density congregations of elk. From December to March 2000–2007, we determined the winter predation patterns of wolves by examining the remains of 289 wolf kills on 3 state-managed feedgrounds and adjacent winter range near Jackson, Wyoming. During winters 2002–2005, we also monitored the movements of radio-collared elk on feedgrounds to describe the response of elk to the presence of wolf kills. Thirty-seven percent (n = 106) of kills were located on elk feedgrounds where elk composition included 49% calves, 42% adult females, 5% adult males, and 5% unknown. Sixty-three percent (n = 183) of kills were located on winter range adjacent to feedgrounds and prey species consisted of 90% elk (38% calves, 35% adult females, 24% adult males, 2% unknown), 9% moose (Alces alces; 13% calves, 69% adult females, 6% adult males, 1% unknown), 1% mule deer (Odocoileus hemionus; 1 fawn, 1 adult female), and 0.5% adult female bison (Bison bison). Mean age of elk killed on feedgrounds was 4.2 years (range = 0–20) and 4.6 years (range = 0–23) on winter range. Calves were selected more than available in most years with female elk killed less than expected. Adult males were killed more than expected in 2005–2007. Eighty-eight percent (n = 198) of the time elk remained on the feedground even when wolves made a kill. Less commonly, elk left the feedground, gathered in larger herds on adjacent feedgrounds absent of wolves, and returned within a few days (6%, n = 13) or left the feedground for another feedground and did not return for the rest of the winter (6%; n = 14). Elk were less likely to leave feedgrounds in the presence of a wolf kill when there were more elk on that feedground. Elk left feedgrounds with greater topography and tree cover (Alkali and Fish Creek) and gathered on the flat, open feedgrounds (Patrol Cabin) more frequently than they left flat, open feedgrounds for feedgrounds with greater topography and tree cover. Our results indicate wolves in our study area primarily preyed on elk and exhibited a strong preference for elk calves. High-density concentrations of elk on feedgrounds will continue to be an attractant for wolves. Although elk leave feedgrounds for reasons other than wolf presence, any displacement of elk from feedgrounds due to wolves will be temporary. State managers have the ability to alter management strategies (e.g., increasing wolf harvest, phasing out elk feeding, increasing the intensity of elk feeding) in an effort to affect predator-prey relationships. © 2019 The Wildlife Society.     

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

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

Temporal variation in site fidelity: scale-dependent effects of forage abundance and predation risk in a non-migratory large herbivore

Large herbivores are typically confronted by considerable spatial and temporal variation in forage abundance and predation risk. Although animals can employ a range of behaviours to balance these limiting factors, scale-dependent movement patterns are expected to be an effective strategy to reduce predation risk and optimise foraging opportunities. We tested this prediction by quantifying site fidelity of global positioning system-collared, non-migratory female elk (Cervus canadensis manitobensis) across multiple nested temporal scales using a long-established elk–wolf (Canis lupus) system in Manitoba, Canada. Using a hierarchical analytical approach, we determined the combined effect of forage abundance and predation risk on variation in site fidelity within four seasons across four nested temporal scales: monthly, biweekly, weekly, daily. Site fidelity of female elk was positively related to forage-rich habitat across all seasons and most temporal scales. At the biweekly, weekly and daily scales, elk became increasingly attached to low forage habitat when risk was high (e.g. when wolves were close or pack sizes were large), which supports the notion that predator-avoidance movements lead to a trade-off between energetic requirements and safety. Unexpectedly, predation risk at the monthly scale increased fidelity, which may indicate that elk use multiple behavioural responses (e.g. movement, vigilance, and aggregation) simultaneously to dilute predation risk, especially at longer temporal scales. Our study clearly shows that forage abundance and predation risk are important scale-dependent determinants of variation in site fidelity of non-migratory female elk and that their combined effect is most apparent at short temporal scales. Insight into the scale-dependent behavioural responses of ungulate populations to limiting factors such as predation risk and forage variability is essential to infer the fitness costs incurred.     

Effects of wolf pack size and winter conditions on elk mortality

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

Mobility of moose—comparing the effects of wolf predation risk,reproductive status,and seasonality

In a predator–prey system, prey species may adapt to the presence of predators with behavioral changes such as increased vigilance, shifting habitats, or changes in their mobility. In North America, moose (Alces alces) have shown behavioral adaptations to presence of predators, but such antipredator behavioral responses have not yet been found in Scandinavian moose in response to the recolonization of wolves (Canis lupus). We studied travel speed and direction of movement of GPS‐collared female moose (n = 26) in relation to spatiotemporal differences in wolf predation risk, reproductive status, and time of year. Travel speed was highest during the calving (May–July) and postcalving (August–October) seasons and was lower for females with calves than females without calves. Similarly, time of year and reproductive status affected the direction of movement, as more concentrated movement was observed for females with calves at heel, during the calving season. We did not find support for that wolf predation risk was an important factor affecting moose travel speed or direction of movement. Likely causal factors for the weak effect of wolf predation risk on mobility of moose include high moose‐to‐wolf ratio and intensive hunter harvest of the moose population during the past century.     

Mothers' Movements: Shifts in Calving Area Selection by Partially Migratory Elk

Loss of migratory behavior or shifts in migratory ranges are growing concerns to wildlife managers. How ungulates prioritize safety from predators at the expense of high-quality foraging opportunities during calving may be key to understanding these shifts and long-term reproductive success. We compared trade-offs in selection for forage and predation risk by elk (Cervus canadensis) following 3 migratory tactics (western and eastern migration and resident) during 2 time periods in a declining (by almost 70% from 2002–2016), partially migratory elk population adjacent to Banff National Park in Alberta, Canada. We hypothesized that regardless of migratory tactic, maternal elk would show stronger trade-offs between high-quality foraging associated with higher predation risk and forage resources of lower-quality yet lower risk on calving ranges than on ranges used during summer because of vulnerability of their offspring. Additionally, we hypothesized these trade-offs would occur at high (2002–2006) and low (2013–2016) elk population sizes. We used a machine-learning algorithm to predict dates of parturition based on global positioning system (GPS) movements of elk equipped with vaginal implants (n = 60) and predictions were within 1.43 ± 0.85 (SE) days of the known date. We applied the model to an additional 58 GPS-collared elk without vaginal implants. Based on changes in localized movements, we defined calving areas as the 26 days post-parturition and compared habitat characteristics of calving areas to 10 similar-sized areas centered on random locations during summer for the same individual in a latent selection framework. Across the 2 time periods, parturition occurred from 8 May–11 July with median parturition dates differing among migratory tactics and residents shifting towards an earlier parturition date in the later period. All elk, regardless of migratory tactic and time period, selected calving areas with greater forage resources than were available on areas used during summer, with no evidence for greater selection of areas that reduced predation risk at the expense of higher-quality foraging. Calving season selection for areas with abundant forage exposed western migrants to high risk of bear (Ursus spp.) predation, residents to high risk of wolf (Canis lupus) predation, and eastern migrants to low risk of bear and wolf predation. Patterns in exposure to predation risk during calving between migratory tactics were consistent with the recent decline in western migrants and increase in eastern migrants, implying that conditions on calving areas contributed to observed changes in the number of elk following these tactics. © 2021 The Wildlife Society.     

Neonatal mortality of elk driven by climate, predator phenology and predator community composition

1.?Understanding the interaction among predators and between predation and climate is critical to understanding the mechanisms for compensatory mortality. We used data from 1999 radio-marked neonatal elk (Cervus elaphus) calves from 12 populations in the north-western United States to test for effects of predation on neonatal survival, and whether predation interacted with climate to render mortality compensatory. 2.?Weibull survival models with a random effect for each population were fit as a function of the number of predator species in a community (3-5), seven indices of climatic variability, sex, birth date, birth weight, and all interactions between climate and predators. Cumulative incidence functions (CIF) were used to test whether the effects of individual species of predators were additive or compensatory. 3.?Neonatal elk survival to 3 months declined following hotter previous summers and increased with higher May precipitation, especially in areas with wolves and/or grizzly bears. Mortality hazards were significantly lower in systems with only coyotes (Canis latrans), cougars (Puma concolor) and black bears (Ursus americanus) compared to higher mortality hazards experienced with gray wolves (Canis lupus) and grizzly bears (Ursus horribilis). 4.?In systems with wolves and grizzly bears, mortality by cougars decreased, and predation by bears was the dominant cause of neonatal mortality. Only bear predation appeared additive and occurred earlier than other predators, which may render later mortality by other predators compensatory as calves age. Wolf predation was low and most likely a compensatory source of mortality for neonatal elk calves. 5.?Functional redundancy and interspecific competition among predators may combine with the effects of climate on vulnerability to predation to drive compensatory mortality of neonatal elk calves. The exception was the evidence for additive bear predation. These results suggest that effects of predation by recovering wolves on neonatal elk survival, a contentious issue for management of elk populations, may be less important than the composition of the predator community. Future studies would benefit by synthesizing overwinter calf and adult-survival data sets, ideally from experimental studies, to test the roles of predation in annual compensatory and additive mortality of elk.     

Changes in Elk Resource Selection and Distributions Associated With a Late-Season Elk Hunt

ABSTRACT Changes in resource selection associated with human predation risk may alter elk distributions and availability for harvest. We used Global Positioning System data collected from telemetered female elk (Cervus elaphus) to evaluate effects of refuges (areas where hunting was prohibited), spatial variation in hunting risk, and landscape attributes on resource selection within an established Greater Yellowstone Area, USA, winter range. We also evaluated elk distributions during and outside of a late-season hunting period. Refuge areas and landscape attributes such as habitat type and snow water equivalents (SWE) affected resource selection. Elk selection for flat grasslands increased as SWE increased, likely because these areas were windswept, leaving grasses exposed for foraging. Elk distributions differed during hunting and no-hunting periods. During the hunting period, elk shifted to privately owned refuge areas and the estimated odds of elk occupying refuge areas more than doubled. Risk-driven changes in resource selection resulted in reduced availability of elk for harvest. Elk selection for areas where hunting is prohibited presents a challenge for resource managers that use hunting as a tool for managing populations and influences grazing patterns on private ranchlands.     

Complex tactics in a dynamic large herbivore–carnivore spatiotemporal game

The spatiotemporal game between predators and prey is a fundamental process governing their distribution dynamics. Players may adopt different tactics as the associated costs and benefits change through time. Yet few studies have investigated the potentially simultaneous and dynamic nature of movement tactics used by both players. It is particularly unclear to what extent perceived predation risk mediates the fine‐scale distribution of large and dangerous prey, which are mostly driven by bottom–up, resource‐related processes. We built habitat use and movement models based on 10 years of monitoring GPS‐collared grey wolves Canis lupus and plains bison Bison bison bison in Prince Albert National Park, Canada, to investigate the predator–large prey game in a multi‐prey system. Bison did not underuse patches of high‐quality vegetation at any time during the seasonal cycle even though wolves were selectively patrolling these areas. Rather, in at least one season, bison engaged in complex tactics comprised of proactive responses to the long‐term distribution (risky places) and reactive responses to the immediate proximity (risky times) of their opponent. In summer–autumn, bison reduced the time spent in food‐rich patches as both the long‐term use and the immediate proximity of wolves increased. By demonstrating that wolf distribution triggers patch abandonment by bison, we provide a key element in support of the shell game hypothesis – where prey move constantly to avoid predators attempting to anticipate their location. In winter, a season of relatively high energetic stress, bison no longer abandoned food‐rich patches as predation risk increased, while no bison responses to wolves were observed in spring–summer. Our work demonstrates the highly dynamic and complex nature of the predator–large prey spatiotemporal game, a key trait‐mediated mechanism by which trophic interactions structure ecological communities.