Raven scavenging favours group foraging in wolves

Wolves, Canis lupus, routinely live in large packs that include unrelated individuals and mature offspring. Studies show that individual wolves that live in large packs suffer reduced foraging returns. Therefore, group hunting and group living (sociality) in wolves is generally thought to be favoured by indirect fitness gains accrued through kin-directed altruism. However, we show that kin-directed altruism cannot account for groups that include mature offspring or unrelated individuals. We also present an analysis that incorporates a previously ignored feature of wolf foraging ecology, namely the loss of food to scavenging ravens, Corvus corax. By accounting for this process, we show that individuals in large packs do indeed accrue foraging advantages. In the hypothetical absence of this scavenging pressure, an individual would maximize its rate of prey acquisition, and minimize its risk of energetic shortfall, by foraging with just one other individual. However, incorporating the effect of scavenging by ravens leads to a dramatic increase in the predicted group size. Our analysis indicates that per capita gains are highest in the largest observed packs. The greater food-sharing costs in a larger pack are more than offset by smaller losses to scavengers and increased rates of prey acquisition. Thus, in contrast with previous interpretations, the selfish benefits of social foraging appear to contribute to the maintenance of sociality in wolves after all. We explore whether such benefits favour group living in various social carnivores that hunt large prey and are thus vulnerable to scavenging.     

Scrounging Tactics in Free-Ranging Ravens, Corvus corax

Social foraging allows individuals to scrounge, i.e. to exploit the food others have made available. The conditions promoting scrounging as an alternative foraging tactic have yet received limited attention. We presently examine whether ravens, as opportunistic scavengers, adjust their foraging tactics according to the potential costs involved in accessing a particular food source. We observed wild ravens foraging in a game park, at the enclosures of wolves, Canis lupus, and wild boars, Sus scrofa. Wolves may aggressively defend their food and even kill ravens, whereas wild boars do not. When co‐feeding with wolves, the ravens showed higher scrounging rates than with wild boars. Only at the wolves, they tended to specialize either on scrounging or on getting food directly from the site. However, scrounging techniques differed in relation to the state of food depletion. Early on, after food became available, the ravens most frequently displaced others from food, whereas towards the end, stealing, solicited sharing, and cache raiding became prevalent. These techniques differed in their profitability and their use was related to the scroungers’ age, social status and affiliative relationships. This suggests that ecological conditions, such as co‐feeding with potential predators, may influence the individuals’ decision whether or not to scrounge in competition for food. Social conditions, on the other hand, may affect the way how to get at food possessed by others and may thus, to a large extent, determine the profitability of scrounging.     

Comparison of Mexican Wolf and Coyote Diets in Arizona and New Mexico

ABSTRACT Interactions between wolves (Canis lupus) and coyotes (C. latrans) can have significant impacts on their distribution and abundance. We compared diets of recently translocated Mexican wolves (C. l. baileyi) with diets of resident coyotes in Arizona and New Mexico, USA. We systematically collected scats during 2000 and 2001. Coyote diet was composed mostly of mammalian species, followed by vegetation and insects. Elk (Cervus elaphus) was the most common item in coyote scats. Mexican wolf diet had a higher proportion of large mammals and fewer small mammals than coyote diet; however, elk was also the most common food item in Mexican wolf scats. Our results suggest that Mexican wolf diet was more similar to coyote diet than previously reported, but coyotes had more seasonal variation. Considering results in other areas, we expect that Mexican wolves will have a negative impact on coyotes through direct mortality and possibly competition. Reintroduction of Mexican wolves may have great impacts on communities by changing relationships among other predators and their prey.     

Influence of Group Size on the Success of Wolves Hunting Bison

An intriguing aspect of social foraging behaviour is that large groups are often no better at capturing prey than are small groups, a pattern that has been attributed to diminished cooperation (i.e., free riding) in large groups. Although this suggests the formation of large groups is unrelated to prey capture, little is known about cooperation in large groups that hunt hard-to-catch prey. Here, we used direct observations of Yellowstone wolves (Canis lupus) hunting their most formidable prey, bison (Bison bison), to test the hypothesis that large groups are more cooperative when hunting difficult prey. We quantified the relationship between capture success and wolf group size, and compared it to previously reported results for Yellowstone wolves hunting elk (Cervus elaphus), a prey that was, on average, 3 times easier to capture than bison. Whereas improvement in elk capture success levelled off at 2–6 wolves, bison capture success levelled off at 9–13 wolves with evidence that it continued to increase beyond 13 wolves. These results are consistent with the hypothesis that hunters in large groups are more cooperative when hunting more formidable prey. Improved ability to capture formidable prey could therefore promote the formation and maintenance of large predator groups, particularly among predators that specialize on such prey.     

Resource dispersion and consumer dominance: scavenging at wolf- and hunter-killed carcasses in Greater Yellowstone, USA

The Greater Yellowstone Ecosystem in the northern Rocky Mountains provides the context for a natural experiment to investigate the response of consumers to resources with differing spatial and temporal dispersion regimes. Grey wolves (Canis lupus) and human hunters both provide resource subsidies to scavengers by provisioning them with the remains of their kills. Carrion from hunter kills is highly aggregated in time and space whereas carrion from wolf kills is more dispersed in both time and space. We estimated the total amount of carrion consumed by each scavenger species at both wolf and hunter kills over 4 years. Species with large feeding radii [bald eagles (Haliaeetus leucocephalus) and ravens (Corvus corax)], defined as the area over which a consumer can efficiently locate and integrate resources, dominated consumption at the highly aggregated hunter kills whereas competitively dominant species [coyotes (Canis latrans)] dominated at the more dispersed wolf kills. In addition, species diversity and the evenness of carrion consumption between scavengers was greater at wolf kills than at hunter kills while the total number of scavengers at hunter kills exceeded those at wolf kills. From a community perspective, the top–down effect of predation is likely to be stronger in the vicinity of highly aggregated resource pulses as species with large feeding radii switch to feeding on alternative prey once the resource pulse subsides.     

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.     

Spatio-temporal segregation of facultative avian scavengers at ungulate carcasses

Spatial and/or temporal segregation of resource use are mechanisms that may allow coexistence between potential competitors. Spatial and temporal patterns of carrion use were studied in the main avian scavengers of Sierra Espuña Natural Park (SE, Spain). We monitored the use of ungulate carcasses provided by hunting during winter and summer of 2005–2006. Non-breeding ravens exploited aggregated carcasses and depleted the resource in a few days while golden eagles used scattered carcasses over a longer consumption period. Moreover, non-breeding ravens exploited carcasses when golden eagles were less active. Almost all available hunter kills were exploited by golden eagles and/or ravens during winter whereas a high number of carcasses were not used during the breeding season, suggesting strong competition in a period of low food availability. Thus carrion arrangement, seasonal behaviour and competitive interactions could be affecting the foraging patterns of these two species. Interestingly, intraspecific aggregation of the smaller species and dominance of the larger one may act on the use of shared resources and trigger segregation mechanisms. Our results support that differences in the spatial and temporal patterns of resource use may allow resource partitioning between two species, thus facilitating their coexistence in sympatric areas.     

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.

     

Interference competition between wolves and coyotes during variable prey abundance

Interference competition occurs when two species have similar resource requirements and one species is dominant and can suppress or exclude the subordinate species. Wolves (Canis lupus) and coyotes (C. latrans) are sympatric across much of their range in North America where white‐tailed deer (Odocoileus virginianus) can be an important prey species. We assessed the extent of niche overlap between wolves and coyotes using activity, diet, and space use as evidence for interference competition during three periods related to the availability of white‐tailed deer fawns in the Upper Great Lakes region of the USA. We assessed activity overlap (Δ) with data from accelerometers onboard global positioning system (GPS) collars worn by wolves (n = 11) and coyotes (n = 13). We analyzed wolf and coyote scat to estimate dietary breadth (B) and food niche overlap (α). We used resource utilization functions (RUFs) with canid GPS location data, white‐tailed deer RUFs, ruffed grouse (Bonasa umbellus) and snowshoe hare (Lepus americanus) densities, and landscape covariates to compare population‐level space use. Wolves and coyotes exhibited considerable overlap in activity (Δ = 0.86–0.92), diet (B = 3.1–4.9; α = 0.76–1.0), and space use of active and inactive RUFs across time periods. Coyotes relied less on deer as prey compared to wolves and consumed greater amounts of smaller prey items. Coyotes exhibited greater population‐level variation in space use compared to wolves. Additionally, while active and inactive, coyotes exhibited greater selection of some land covers as compared to wolves. Our findings lend support for interference competition between wolves and coyotes with significant overlap across resource attributes examined. The mechanisms through which wolves and coyotes coexist appear to be driven largely by how coyotes, a generalist species, exploit narrow differences in resource availability and display greater population‐level plasticity in resource use.     

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.     

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.     

Assessing differential prey selection patterns between two sympatric large carnivores

Several conceptual models describing patterns of prey selection by predators have been proposed, but such models rarely have been tested empirically, particularly with terrestrial carnivores. We examined patterns of prey selection by sympatric wolves ( Canis lupus ) and cougars ( Puma concolor ) to determine i) if both predators selected disadvantaged prey disproportionately from the prey population, and ii) if the specific nature and intensity of prey selection differed according to disparity in hunting behavior between predator species. We documented prey characteristics and kill site attributes of predator kills during winters 1999–2001 in Idaho, and located 120 wolf-killed and 98 cougar-killed ungulates on our study site. Elk ( Cervus elephus ) were the primary prey for both predators, followed by mule deer ( Odocoileus hemionus ). Both predators preyed disproportionately on elk calves and old individuals; among mule deer, wolves appeared to select for fawns, whereas cougars killed primarily adults. Nutritional status of prey, as determined by percent femur marrow fat, was consistently poorer in wolf-killed prey. We found that wolf kills occurred in habitat that was more reflective of the entire study area than cougar kills, suggesting that the coursing hunting behavior of wolves likely operated on a larger spatial scale than did the ambush hunting strategy of cougars. We concluded that the disparity in prey selection and hunting habitat between predators probably was a function of predator-specific hunting behavior and capture success, where the longer prey chases and lower capture success of wolf packs mandated a stronger selection for disadvantaged prey. For cougars, prey selection seemed to be limited primarily by prey size, which could be a function of the solitary hunting behavior of this species and the risks associated with capturing prime-aged prey.     

Can't bear the competition: Energetic losses from kleptoparasitism by a dominant scavenger may alter foraging behaviors of an apex predator

The interspecific interactions of apex predators are integral to the function of ecological communities, but most studies have focused on understanding their top down effects. Kleptoparasitism (the stealing of procured food) by dominant scavengers can have negative effects on populations and behaviors of apex predators. We captured 7 pumas (Puma concolor) and fitted them with GPS collars to investigate potential kill sites (n = 352), some of which we monitored with camera traps (n = 58). We analyzed whether observed kleptoparasitism by American black bears (Ursus americanus) affected puma energetics and foraging behavior. We found that black bears were the most frequent scavenger of puma kills (72.4%), and we documented bears scavenging puma kills during every month. The top model for bear detection of puma kills included prey size, temperature, and canopy cover, with bears more likely to scavenge from adult black-tailed deer (Odocoileus hemionus columbianus) carcasses in warmer temperatures and under dense canopy cover. When black bear scavenging occurred, pumas spent 22% less time at their kill and incurred energetic losses. In response, pumas shortened their inter-kill intervals by 1.3 days thus increasing their kill rates. Our results demonstrate how a dominant scavenger directly mediates the foraging behavior of an apex predator. These results suggest that community interactions do not necessarily start at the top in top-down systems, and the effects of predators on prey populations can only be understood within their respective ecological communities.     

Biomass Flow and Scavengers Use of Carcasses after Re-Colonization of an Apex Predator

Background

Reestablishment of apex predators influences the availability and distribution of biomass for scavengers and can therefore be an important agent for structuring species communities. We studied how the re-colonization of the Scandinavian Peninsula by wolves (Canis lupus) affected the amount and temporal variation in use of moose (Alces alces) carcasses.

Methodology/Principal Findings

We compared the availability of biomass from remains at wolf kills with those killed by hunters, vehicle collisions and natural death. Movement-triggered cameras monitored patterns of use on wolf kills and remains from hunter harvest by scavengers (n = 15 276) in relation to time of year, available carcass biomass, time since the death of the moose and presence of wolves. Remains from hunter harvest were the largest food source for scavengers both within wolf territories (57%) and in areas without wolves (81%). The total annual biomass available were similar in areas with (25 648 kg) and without (24 289 kg) wolves. Presence of wolves lowered the peak biomass available from hunter harvest in October (20%) and increased biomass available during December to August (38–324% per month). The probability of scavengers being present decreased faster with time at remains from hunter harvest compared to wolf kills and both the probability of being present and the number of visits by scavengers to wolf kills increased as the amount of biomass available on the carcass increased.

Conclusions/Significance

Wolves reduced the seasonal variation of biomass from moose carcasses and most important increased it during spring. Scavengers also visited wolf kills most frequently during spring when most scavenging species have young, which may lead to an increase in survival and/or reproductive success of scavengers within wolf territories. This applies both for abundant scavenging species that were the most frequent visitors at wolf kills and threatened scavengers with lower visit frequency.     

Size‐assortative choice and mate availability influences hybridization between red wolves (Canis rufus) and coyotes (Canis latrans)

Anthropogenic hybridization of historically isolated taxa has become a primary conservation challenge for many imperiled species. Indeed, hybridization between red wolves (Canis rufus) and coyotes (Canis latrans) poses a significant challenge to red wolf recovery. We considered seven hypotheses to assess factors influencing hybridization between red wolves and coyotes via pair‐bonding between the two species. Because long‐term monogamy and defense of all‐purpose territories are core characteristics of both species, mate choice has long‐term consequences. Therefore, red wolves may choose similar‐sized mates to acquire partners that behave similarly to themselves in the use of space and diet. We observed multiple factors influencing breeding pair formation by red wolves and found that most wolves paired with similar‐sized conspecifics and wolves that formed congeneric pairs with nonwolves (coyotes and hybrids) were mostly female wolves, the smaller of the two sexes. Additionally, we observed that lower red wolf abundance relative to nonwolves and the absence of helpers increased the probability that wolves consorted with nonwolves. However, successful pairings between red wolves and nonwolves were associated with wolves that maintained small home ranges. Behaviors associated with territoriality are energetically demanding and behaviors (e.g., aggressive interactions, foraging, and space use) involved in maintaining territories are influenced by body size. Consequently, we propose the hypothesis that size disparities between consorting red wolves and coyotes influence positive assortative mating and may represent a reproductive barrier between the two species. We offer that it may be possible to maintain wild populations of red wolves in the presence of coyotes if management strategies increase red wolf abundance on the landscape by mitigating key threats, such as human‐caused mortality and hybridization with coyotes. Increasing red wolf abundance would likely restore selection pressures that increase mean body and home‐range sizes of red wolves and decrease hybridization rates via reduced occurrence of congeneric pairs.     

Inter-specific territoriality in a Canis hybrid zone: spatial segregation between wolves, coyotes, and hybrids

Gray wolves (Canis lupus) and coyotes (Canis latrans) generally exhibit intraspecific territoriality manifesting in spatial segregation between adjacent packs. However, previous studies have found a high degree of interspecific spatial overlap between sympatric wolves and coyotes. Eastern wolves (Canis lycaon) are the most common wolf in and around Algonquin Provincial Park (APP), Ontario, Canada and hybridize with sympatric gray wolves and coyotes. We hypothesized that all Canis types (wolves, coyotes, and hybrids) exhibit a high degree of spatial segregation due to greater genetic, morphologic, and ecological similarities between wolves and coyotes in this hybrid system compared with western North American ecosystems. We used global positioning system telemetry and probabilistic measures of spatial overlap to investigate spatial segregation between adjacent Canis packs. Our hypothesis was supported as: (1) the probability of locating wolves, coyotes, and hybrids within home ranges ( $\bar{x}$  = 0.05) or core areas ( $\bar{x}$  < 0.01) of adjacent packs was low; and (2) the amount of shared space use was negligible. Spatial segregation did not vary substantially in relation to genotypes of adjacent packs or local environmental conditions (i.e., harvest regulations or road densities). We provide the first telemetry-based demonstration of spatial segregation between wolves and coyotes, highlighting the novel relationships between Canis types in the Ontario hybrid zone relative to areas where wolves and coyotes are reproductively isolated. Territoriality among Canis may increase the likelihood of eastern wolves joining coyote and hybrid packs, facilitate hybridization, and could play a role in limiting expansion of the genetically distinct APP eastern wolf population.     

Traumatic, degenerative, and developmental lesions in wolves and coyotes from Saskatchewan.

A retrospective review was done of traumatic and osseous lesions in 241 wolves (Canis lupus) and 316 coyotes (Canis latrans) necropsied at the University of Saskatchewan between 1971 and 1990. Most lesions were the result of interspecific conflict. The most frequently occurring lesion in wolves was fracture of one or more bones, primarily ribs. Lesions were healed in most cases and appeared to be compatible with injuries caused by prey animals. One wolf, found dead, died as a result of thoracic trauma. Limb and skull fractures were less common. Fractures were uncommon in coyotes. The most frequent injuries in coyotes were related to gunshot wounds. Four coyotes had been killed but not eaten by wolves. One wolf had been killed and another attacked by wolves. Porcupine (Erethizon dorsatum) quills contributed to the death of a wolf and two coyotes. Degenerative joint disease, involving the spinal column and limb joints, was found in a few individuals of both species. A coyote had severe anomalies of the spinal column and a wolf had anomalous external genitalia.     

Differential risk effects of wolves on wild versus domestic prey have consequences for conservation

Predators play integral roles in shaping ecosystems through cascading effects to prey and vegetation. Such effects occur when prey species alter their behavior to avoid predators, a phenomenon called the risk effects of predators. Risk effects of wild predators such as wolves are well documented for wild prey, but not for free ranging domestic animals such as cattle despite their importance for ecosystem function and conservation. We compared risk effects of satellite‐collared wolves (n = 16) on habitat selection by global‐positioning‐system‐collared elk (n = 10) and cattle (n = 31). We calculated resource selection functions (RSFs) in periods before, during and after wolf visits in elk home ranges or cattle pastures. The habitat variables tested included: distance to roads and trails, terrain ruggedness, food‐quality and distance to forest. When wolves were present, elk stayed closer to forest cover and selected less for high‐quality‐food habitat. Thus, the risk effects of wolf presence on elk produced a change in the tradeoff between food and cover selection. Cattle responded by avoiding high‐quality‐food habitat and selecting areas closer to roads and trails (where people likely provided security), but these effects manifested only after wolves had left. Artificial selection in cattle may have attenuated natural anti‐predator behaviors. The effects of predators on ecosystems are likely different when mediated through risk effects on domestic compared to wild animals. Furthermore, predator control in response to livestock predation, an important conservation issue, may produce broad ecosystem effects triggered by decrease of an important predator species. Conservation planners should consider these effects where domestic herbivores are dominant species in the ecosystem.     

Diurnal and nocturnal functional response of juvenile Notonecta maculata considered as a consequence of shifting predation behaviour

Presence or absence of light is considered to significantly affect predation within freshwater communities. In relation to light regime, the scope of the present study is to identify proximate factors accounting for different rates of predation in juvenile Notonecta maculata. It has been shown that foraging of juvenile N. maculata on Daphnia magna is reduced under dark conditions. These differences are accounted for by the presence/absence of light and are not regulated by any endogenous circadian rhythm. Direct observation of predation components revealed that in the dark, daphnid prey is detected at shorter distances, thus lowering the rate of encounter which finally results in a decreasing number of prey items eaten. Handling time was found to significantly increase during dark periods. Since the amount of food ingested per prey item increases to a certain extent with increasing handling time, it is suggested that the decreasing number of daphnids consumed is compensated by gaining a higher amount of food per item during a lengthier, more thorough, handling of the prey. A mechanistic model that describes the foraging process along a general predation cycle was parameterized based on the results of direct observations, instead of using classic functional response curves. This allows the comparison of model output to independent functional response data. A good correlation between observed and predicted data confirms the assumption that the reduction of the nocturnal predation rate is caused by shifting predation behaviour and indicates that the crucial light/dark differences in the foraging of N. maculata are considered in the approach.     

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.