Variation in predation risk and vole feeding behaviour: a field test of the risk allocation hypothesis

Many prey animals experience temporal variation in the risk of predation and therefore face the problem of allocating their time between antipredator efforts and other activities like feeding and breeding. We investigated time allocation of prey animals that balanced predation risk and feeding opportunities. The predation risk allocation hypothesis predicts that animals should forage more in low- than in high-risk situations and that this difference should increase with an increasing attack ratio (i.e. difference between low- and high-risk situations) and proportion of time spent at high risk. To test these predictions we conducted a field test using bank voles (Clethrionomys glareolus) as a prey and the least weasel (Mustela nivalis nivalis) as a predator. The temporal pattern and intensity of predation risk were manipulated in large outdoor enclosures and the foraging effort and patch use of voles were measured by recording giving-up densities. We did not observe any variation in feeding effort due to changes in the level of risk or the proportion of time spent under high-risk conditions. The only significant effect was found when the attack ratio was altered: the foraging effort of voles was higher in the treatment with a low attack ratio than in the treatment with a high attack ratio. Thus the results did not support the predation risk allocation hypothesis and we question the applicability of the hypothesis to our study system. We argue that the deviation between the observed pattern of feeding behaviour of bank voles and that predicted by the predation risk allocation hypothesis was mostly due to the inability of voles to accurately assess the changes in the level of risk. However, we also emphasise the difficulties of testing hypotheses under outdoor conditions and with mammals capable of flexible behavioural patterns.     

Does Temporal Variation in Predation Risk Influence the Intensity of Antipredator Responses? A Test of the Risk Allocation Hypothesis

Temporal variation in predation risk may fundamentally influence antipredator responses of prey animals. To maximize lifetime fitness, prey must be able to optimize energy gain and minimize predation risk, and responses to current levels of risk may be influenced by background levels of risk. A ‘risk allocation’ model has recently been proposed to predict the intensity of antipredator responses that should occur as predation risk varies over time. Prey animals from high‐risk environments should respond to predators with relatively low intensities of antipredator behaviour because long periods of antipredator behaviour may result in unacceptable decreases in levels of foraging activity. Moreover, animals that are under frequent risk should devote more energy to foraging during brief pulses of safety compared with animals under infrequent attack. In this study, we experimentally tested the risk allocation hypothesis. We exposed juvenile rainbow trout, Oncorhynchus mykiss, to three levels of risk (high, moderate and low) crossed with two levels of temporal variation (exposed to risk three times a day and once a day). In accordance with the model, we found that trout exposed to risky situations more frequently responded with significantly less intense antipredator behaviour than trout exposed to risk infrequently. The intensity of response of trout exposed to moderate risk three times a day decreased to levels similar to situations of no risk. However, in contrast to the second prediction of the model, animals under frequent risk were not more active during periods of safety compared with animals under infrequent risk. Although behaviour in the face of predation risk was dependent on the broader temporal context in which risk varied, the specific predictions of the risk allocation model were only partly supported.     

The predation risks of interspecific eavesdropping: weasel–vole interactions

Competing species benefit from eavesdropping on each other's signals by learning about shared resources or predators. But conspicuous signals are also open to exploitation by eavesdropping predators and should also pose a threat to other sympatric prey species. In western Finland, sibling voles Microtus rossiameridionalis and field voles M. agrestis compete for food and space, and both species rely upon scent marks for intraspecific communication. Both vole species are prey to a range of terrestrial scent hunting predators such as least weasels, however, the competitively superior sibling voles are taken preferentially. We tested in large out‐door enclosures whether field voles eavesdrop on the signals of its competitor, and whether they behave as though this eavesdropping carries a risk of predation. We presented field voles with scent marks from unknown conspecifics and sibling voles and measured their visitation, activity and scent marking behaviours at these scents under high (weasel present) and low (weasel absent) predation risk. Field voles readily visited both field and sibling vole scents under both high and low predation risk; however their activity at sibling vole scent marks declined significantly under increased predation risk. In contrast, predation risk did not affect field voles’ activity at conspecific scents. Thus, field voles were compelled to maintain eavesdropping on heterospecific scents under an increased risk of predation, however they compensated for this additional risk by reducing their activity at these risky scents. Scent marking rates declined significantly under high predation risk. Our results therefore reveal a hidden complexity in the use of social signals within multi‐species assemblages that is clearly sensitive to the potential for increased predation risk. The predation risks of interspecific eavesdropping demonstrated here represents a significant generalisation of the concept of associational susceptibility.     

Heterogeneous landscapes and the role of refuge on the population dynamics of a specialist predator and its prey

How, and where, a prey species survives predation by a specialist predator during low phases of population fluctuations or a cycle, and how the increase phase of prey population is initiated, are much-debated questions in population and theoretical ecology. The persistence of the prey species could be due mainly to habitats that act as refuges from predation and/or due to anti-predatory behaviour of individuals. We present models for the former conjecture in two (and three) habitat systems with a specialist predator and its favoured prey. The model is based on dispersal of prey between habitats with high reproductive output but high risk of predation, and less productive habitats with relatively low risk of predation. We illustrate the predictions of our model using parameters from one of the most intriguing vertebrate predator–prey systems, the multi-annual population cycles of boreal voles and their predators. We suggest that cyclic population dynamics could result from a sequence of extinction and re–colonization events. Field voles (Microtus agrestis), a key vole species in the system, can be hunted to extinction in their preferred meadow habitat, but persist in sub-optimal wet habitats where their main predator, the least weasel (Mustela nivalis nivalis) has a low hunting efficiency. Re–colonization of favourable habitats would occur after the predator population crashes. At the local scale, the model suggests that the periodicity and amplitude of population cycles can be strongly influenced by the relative availability of risky and safe habitats for the prey. Furthermore, factors like intra-guild predation may lead to reduced predation pressure on field voles in sub-optimal habitats, which would act as a refuge for voles during the low phase of their population cycles. Elasticity analysis suggested that our model is quite robust to changes in most parameters but sensitive to changes in the population dynamics of field voles in the optimal grassland habitat, and to the maximum predation rate of weasels.     

Predation on two vole species by a shared predator: antipredatory response and prey preference

In prey communities with shared predators, variation in prey vulnerability is a key factor in shaping community dynamics. Conversely, the hunting efficiency of a predator depends on the prey community structure, preferences of the predator and antipredatory behavioural traits of the prey. We studied experimentally, under seminatural field conditions, the preferences of a predator and the antipredatory responses of prey in a system consisting of two Myodes species of voles, the grey-sided vole (M. rufocanus Sund.) and the bank vole (M. glareolus Schreb.), and their specialist predator, the least weasel (Mustela nivalis nivalis L.). To quantify the preference of the weasels, we developed a new modelling framework that can be used for unbalanced data. The two vole species were hypothesised to have different habitat-dependent vulnerabilities. We created two habitats, open and forest, to provide different escape possibilities for the voles. We found a weak general preference of the weasels for the grey-sided voles over the bank voles, and a somewhat stronger preference specifically in open habitats. The weasels clearly preferred male grey-sided voles over females, whereas in bank voles, there was no difference. The activity of voles changed over time, so that voles increased their movements immediately after weasel introduction, but later adjusted their movements to times of lowered predation risk. Females that were more active had an elevated mortality risk, whereas in the case of males, the result was the opposite. We conclude that, in vulnerability to predation, the species- or habitat-specific characteristics of these prey species are playing a minor role compared to sex-specific characteristics.     

Microhabitat use and behavior of voles under weasel and raptor predation risk: predator facilitation?

An example of predator facilitation is that a microhabitat shiftin a prey species induced by one predator increases the probabilityof the prey falling victim to other predators. Least weasels(Mustela nivalis) hunt in dense plant cover, whereas kestrels(Falco tinnunculus) hunt in habitats with sparse plant cover.Field voles (Microtus agrestis), the main food of weasels andkestrels, prefer open country with a high grass layer. We simulateda multipredator environment in an aviary (3.0 x 4.8 x 2.2 m)to find out whether predator facilitation plays a role in theinteractions between voles, small mustelids, and raptors. Ineach replicate, we placed a field vole in a pen including sidesof high and low grass layers (cover and open). In a predator-freesituation, voles preferred cover but shifted to open when aweasel was introduced to cover. In the presence of a kestrel,voles occupied cover and decreased their mobility. In the presenceof a weasel plus a kestrel, voles behaved as under the kestrelrisk alone. Therefore, in these aviary circumstances, volesperceived the kestrel risk as greater than the weasel risk.Predator facilitation in the assemblage of predators subsistingon rodent prey may contribute to the crash of the four-yearvole cycle: microhabitat shift due to an avoidance of weaseljaws may drive voles to raptor talons.     

Predator (<Emphasis Type="Italic">Carcinus maenas</Emphasis>) nonconsumptive limitation of prey (<Emphasis Type="Italic">Nucella lapillus</Emphasis>) feeding depends on prey density and predator cue type

Predators often have nonconsumptive effects (NCEs) on prey. For example, upon detection of predator cues, prey can reduce feeding activities to hamper being detected by predators. Previous research showed that waterborne chemical cues from green crabs (Carcinus maenas, predator) limit the dogwhelk (Nucella lapillus, prey) consumption of barnacles regardless of dogwhelk density, even though individual predation risk for dogwhelks decreases with conspecific density. Such NCEs might disappear with dogwhelk density if dogwhelks feed on mussels, as mussel stands constitute better antipredator refuges than barnacle stands. Through a laboratory experiment, we effectively found that crab chemical cues limit the per-capita consumption of mussels by dogwhelks at low dogwhelk density but not at high density. The combination of tactile and chemical cues from crabs, however, limited the dogwhelk consumption of mussels at both dogwhelk densities. The occurrence of such NCEs at both dogwhelk densities could have resulted from tactile cues indicating a stronger predation risk than chemical cues alone. Overall, the present study reinforces the notions that prey evaluate conspecific density when assessing predation risk and that predator cue type affects their perception of risk.     

Mammalian predator–prey interaction in a fragmented landscape: weasels and voles

The relationship between predators and prey is thought to change due to habitat loss and fragmentation, but patterns regarding the direction of the effect are lacking. The common prediction is that specialized predators, often more dependent on a certain habitat type, should be more vulnerable to habitat loss compared to generalist predators, but actual fragmentation effects are unknown. If a predator is small and vulnerable to predation by other larger predators through intra-guild predation, habitat fragmentation will similarly affect both the prey and the small predator. In this case, the predator is predicted to behave similarly to the prey and avoid open and risky areas. We studied a specialist predator’s, the least weasel, Mustela nivalis nivalis, spacing behavior and hunting efficiency on bank voles, Myodes glareolus, in an experimentally fragmented habitat. The habitat consisted of either one large habitat patch (non-fragmented) or four small habitat patches (fragmented) with the same total area. The study was replicated in summer and autumn during a year with high avian predation risk for both voles and weasels. As predicted, weasels under radio-surveillance killed more voles in the non-fragmented habitat which also provided cover from avian predators during their prey search. However, this was only during autumn, when the killing rate was also generally high due to cold weather. The movement areas were the same for both sexes and both fragmentation treatments, but weasels of both sexes were more prone to take risks in crossing the open matrix in the fragmented treatment. Our results support the hypothesis that habitat fragmentation may increase the persistence of specialist predator and prey populations if predators are limited in the same habitat as their prey and they share the same risk from avian predation.     

Fixed vs. Random Temporal Predictability of Predation Risk: An Extension of the Risk Allocation Hypothesis

Predation is a strong selective force acting on prey animals. Predation is by nature highly variable in time; however, this aspect of predation risk has traditionally been overlooked by behavioural ecologists. Lima and Bednekoff proposed the predation risk allocation hypothesis (RAH), predicting how temporal variation in predation risk drives prey antipredator behaviours. This model is based on the concept that prey adaptively allocate their foraging and antipredator efforts across high‐ and low‐risk situations, depending on the duration of high‐ vs. low‐risk situations and the relative risk associated with each of them. An unstudied extension of the RAH is the effect of predictability of predation risk. A predictable risk should lead to prey displaying minimal vigilance behaviours during predictable low‐risk periods and the strongest antipredator behaviours during risky periods. Conversely, an unpredictable predation risk should result in prey displaying constant vigilance behaviour, with suboptimal foraging rates during periods of safety but antipredator behaviours of lower intensity during periods of risk. We tested this extension of the RAH using convict cichlids exposed to high‐risk alarm cues at two frequencies of risk (1× vs. 3×) per day, on either a fixed or random schedule for 5 d. We then tested the fish for a response to high‐risk cues (alarm cues) and to low‐risk cues (disturbance resulting from the introduction of distilled water). Our study supports previous results on the effects of risk frequency and cue intensity on cichlid behaviour. We failed to show an effect of risk predictability on the behavioural responses of cichlids to high‐risk alarm cues, but predictability did influence responses to low‐risk cues. We encourage further studies to test the effect of predictability in other systems.     

Behavioral and neuroendocrine response of Brandt's voles,Lasiopodomys brandtii,to odors of different species

Predator odors are nonintrusive natural stressors of high ethological relevance. Behavioral and hormonal responses and changes in the expression of medial hypothalamic c-fos mRNA were examined in Brandt's voles (Lasiopodomys brandtii) after exposure to feces of a domestic cat (Felis catus), weasel (Mustela sibirica), snake (Xenopeltis hainanensis), goat (Capra aegagrus), and distilled water (control). One hundred voles were tested in the defensive withdrawal apparatus. Voles showed an aversion to the predator odors by showing significant high levels of flight-related behaviors, more freezing behavior, and less and higher grooming behavior when they exposed to weasel and cat feces and more vigilant rearing compared to goat and control groups. Adrenocorticotropic hormone and corticosterone serum levels significantly increased when voles were exposed to the predator odors compared to voles exposed to goat and control odors. A significant high c-fos mRNA level indicates a strong predator odor-induced activation of the medial hypothalamus of the tested voles. Our results display significant fear responses of Brandt's voles exposed to predator odors and that they can skillfully differentiate between odors from different species with odors from sympatric predators triggering the highest responses.     

Antipredator responses of aphids to parasitoids change as a function of aphid physiological state

Under predation risk, prey may prioritize antipredator behaviours and sacrifice feeding. However, energetically constrained animals may choose to sacrifice or change antipredator responses and accept relatively greater risk in order to secure food. In this last case, the antipredator tactics chosen must balance safety and feeding in such a way that costs are minimized and benefits maximized. We studied the antipredator behaviour of pea aphid, Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae) subjected to different periods of food deprivation, against the parasitoid Aphidius ervi (Hymenoptera: Braconidae). As the energetic internal stress of aphids increased, the predominant antipredator response changed from walking away and dropping to kicking behaviour, and parasitization avoidance decreased. Parasitoids did not show preference between food-deprived and nonfood-deprived aphids. Dropping and walking away reduced parasitization from 50 to 33%. These results support the hypothesis that the antipredator behaviour of an aphid changes as a function of internal stress. By performing less costly behaviour such as kicking under energetically constrained conditions, aphids seem to minimize their probability of energy shortfall. Given that aphid antipredator behaviour is a function of nutritional state, its occurrence under natural conditions may match host quality spatial distribution. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Multiple predators induce risk reduction in coexisting vole species

Large predators may affect the hunting efficiency of smaller ones directly by decreasing their numbers, or indirectly by altering their behaviour. Either way this may have positive effects on the density of shared prey. Using large outdoor enclosures, we experimentally studied whether the presence of the Tengmalm's owl Aegolius funereus affects the hunting efficiency of the smallest member of the vole-eating predator guild, the least weasel Mustela nivalis, as measured by population responses of coexisting prey species, the field vole Microtus agrestis and the sibling vole M. levis . We compared the density and survival probability of vole populations exposed to no predation, weasel predation or combined predation by a weasel and an owl. The combined predation of both owl and weasel did not result in obvious changes in the density of sibling and field vole populations compared to the control populations without predators, while predation by least weasel alone decreased the densities of sibling voles and induced a similar trend in field vole densities. Survival of field voles was not affected by predator treatment while sibling vole survival was lower in predator treated populations than in control populations. Our results suggest that weasels are intimidated by avian predators, but without changing the effects of predators on competitive situations between the two vole species. Non-lethal effects of intraguild predation therefore will not necessarily change competitive interactions between shared prey species.     

Variation in the diet composition of a generalist predator, the red fox, in relation to season and density of main prey

Diet composition of a generalist predator, the red fox (Vulpes vulpes) in relation to season (winter or summer) and abundance of multi-annually cyclic voles was studied in western Finland from 1983 to 1995. The proportion of scats (PS; a total of 58 scats) including each food category was calculated for each prey group. Microtus voles (the field vole M. agrestis and the sibling vole M. rossiaemeridionalis) were the main prey group of foxes (PS = 0.55) and they frequently occurred in the scats both in the winter and summer (PSs 0.50 and 0.62, respectively). There was a positive correlation between the PSs of Microtus voles in the winter diet of foxes and the density indices of these voles in the previous autumn. Other microtine rodents (the bank vole Clethrionomys glareolus, the water vole Arvicola terrestris and the muskrat Ondatra zibethicus) were consumed more in winter than in summer. The unusually high small mustelid predation by red foxes (PS = approx. 0.10) in our study area gives qualitative support for the hypothesis on the limiting impact of mammalian predators on least weasel and stoat populations. None of the important prey groups was preyed upon more at low than at high densities of main prey (Microtus voles). This is consistent with the notion that red foxes are generalist predators that tend to opportunistically subsist on many prey groups. Among these prey groups, particularly hares and birds (including grouse), were frequently used as food by foxes.     

Fungal-Mediated Multitrophic Interactions - Do Grass Endophytes in Diet Protect Voles from Predators?

Plant-associated micro-organisms such as mycotoxin-producing endophytes commonly have direct negative effects on herbivores. These effects may be carried over to natural enemies of the herbivores, but this has been rarely explored. We examined how feeding on Neotyphodium endophyte infected (E+) and endophyte free (E−) meadow ryegrass (Scherodonus pratensis) affects body mass, population size and mobility of sibling voles (Microtus levis), and whether the diet mediates the vulnerability of voles to least weasel (Mustela nivalis nivalis) predation. Because least weasels are known to be olfactory hunters, we also examined whether they are able to distinguish olfactory cues of voles fed on E+ and E− diets. Neither body mass of voles nor population size differed between diets. However, contrary to our prediction, least weasels preyed more often on voles fed with E− grass than on voles fed with E+ grass. The mobility of voles fed on E+ grass was reduced compared to voles fed on E− grass, but this effect was unrelated to risk of predation. Least weasels appeared unable to distinguish between excrement odours of voles between the two treatments. Our results suggest that consumption of endophytic grass is not directly deleterious to sibling voles. What''s more, consumption of endophytes appears to be advantageous to voles by reducing risk of mammalian predation. Our study is thus the first to demonstrate an effect of plant-associated microbial symbionts on herbivore-predator interactions in vertebrate communities.     

Survival Through Bottlenecks of Vole Cycles: Refuge or Chance Events?

In small rodent populations with wide-amplitude fluctuations and low-density bottlenecks, the individuals that survive through the bottlenecks may gain major fitness advantages as they will be the founders of the following population expansion. Most hypotheses assume that there exists a physical or behavioural refuge from increased predation risk, and that the survivors are most likely individuals adapted to use such refuges. A recent hypothesis suggests that survival probability is habitat-dependent so that some otherwise sub-optimal habitats provide a spatial refuge from predation risk by the main predator(s). We used spatially replicated long-term (1981–2004) trapping and tracking data of voles (field vole Microtus agrestis and sibling vole M. rossiaemeridionalis) and their main predators (weasel Mustela nivalis and stoat M. erminea) to test predictions based on this hypothesis. We did not find support for the hypothesis. We did not find marked phase-dependent differences in the habitat-level distribution of Microtus voles. Habitat types with low Microtus vole abundance had, on average, comparable predator activity than the main Microtus vole habitats, indicating that there were no habitat-level refuges from predators. There appeared to be no permanent site-level refuges: the spatial distribution of voles varied from one bottleneck to another. This suggests that survival through bottlenecks is at least partly determined by chance events. We propose that in this kind of systems, where relatively short-lived prey are hunted by nomadic or widely ranging predators, short-term anti-predator responses may increase survival prospects as efficiently as more costly anti-predator adaptations, and there is no apparent need to maintain special adaptations to bottleneck situations that occur at infrequent intervals.Co-ordinating editor: J. Tuomi     

Optimal foraging of sticklebacks on swarming prey

Hungry sticklebacks, Gasterosteus aculeatus, preferentially attacked the densest region of a swarm of water fleas, but with decreasing attack readiness they increasingly preferred less dense regions. Such a hunger dependent change in feeding preference has not yet been dealt with by optimal foraging theory. A model, which assumes that high swarm densities provide high feeding rates (because of small inter-prey distances) but also high costs of confusion, predicts that a predator should always choose the lowest prey density in which it can achieve a feeding rate sufficient to satisfy its hunger. Some predictions of the model were experimentally verified. Hungry fish have a higher feeding rate in a high prey density than in a lower density and less hungry fish have a higher rate in a low density than in a high density.     

Behavioural Correlations May Cause Partial Support for the Risk Allocation Hypothesis in Damselfly Larvae

Prey animals are often confronted with situations that differ in predation risk. According to the risk allocation hypothesis, prey animals should adaptively allocate antipredator behaviour in accordance with the magnitude and frequency of those risk situations. According to the first prediction prey animals should increase foraging in the safe situations and decrease foraging in the dangerous situations as these situations become relatively more dangerous. The second prediction is that with increased time spent in the dangerous situations, progressively more foraging effort is shown in both the dangerous and safe situations, especially in the safer ones. Prey animals may, however, show maladaptive behaviour due to behavioural correlations across risk situations. Here we test for the first time both predictions generated by the risk allocation hypothesis while considering behavioural correlations. We reared larvae of the damselfly Ischnura elegans, from the egg stage, under five rearing risk conditions: (i) in isolation, (ii) in the presence of conspecific larvae, (iii) in the presence of one fish, (iv) in the presence of two fish, and (v) in the presence of two fish for 50% of the time. For each rearing risk condition, we scored their behaviour in the absence and in the presence of fish. In accordance with the first prediction, in the absence of a predator, larvae reared under increasing risk conditions increased their level of foraging. In accordance with the second prediction, in the absence of a predator, larvae that were more frequently exposed to fish during rearing, increased foraging. However, opposite to the predictions from the risk allocation hypothesis, foraging increased both with increasing rearing risk, and with increased predator exposure frequency. The observed positive behavioural correlation of foraging activity across test situations with and without fish, may generate the combination of adaptive patterns in the absence of fish and the maladaptive patterns in the presence of fish. Former studies of the risk allocation hypothesis also found, at best, mixed support, and we hypothesize that behavioural correlations across risk situations, if present, will likely cause partial deviations from model predictions.     

Tadpoles of the bronze frog (Rana temporalis) assess predation risk before evoking antipredator defense behavior

Predation threat-associated behavioral response was studied in Rana temporalis tadpoles to discover the importance of predators’ visual and chemical cues (kairomones and diet-derived metabolites of consumed prey) in evoking antipredator behavior. The caged predators (dragonfly larvae) fed on prey tadpoles or insects (Notonecta spp.) and water conditioned with the predators provided the threat stimuli to the tadpole prey. The predators’ visual cues were ineffective in evoking antipredator behaviors in the tadpole prey. However, exposure to caged tadpole-fed predators or water conditioned with tadpole-fed predators elicited predator avoidance behavior in the tadpoles; they stayed away from the predators, significantly reduced swimming activity (swimming time and distance traveled), and increased burst speed. Interestingly, exposure to water conditioned with starved predators did not elicit any antipredator behavior in the prey. Further, the antipredator responses of predator-experienced tadpoles were significantly greater than those exhibited by predator-na?ve tadpoles. The study shows that R. temporalis tadpoles assess predation threat based exclusively on chemical cues emanating from the predators’ dietary metabolites and that the inclusion of conspecific prey items in the diet of the predators is perceived as a threat. The study also shows that antipredator behavior in these tadpoles is innate and is enhanced during subsequent encounters with the predators.     

The role of Apodemus mice and Microtus voles in the diet of the Tengmalm’s owl in Central Europe

Based on a long-term dataset (1999–2010), we investigated how the availability of main prey affects the breeding density and food ecology of the Tengmalm’s owl (Aegolius funereus) in the Czech Republic. In particular, we assessed the role of Microtus voles and Apodemus mice in the diet, based on the main predictions of the optimal diet theory that the diet composition depends on the availability of the main prey. We found that (i) the Tengmalm’s owl exhibited no numerical response to the availability of Microtus voles and Apodemus mice in the field; (ii) the availability of Apodemus mice in the field positively affected their proportion in the diet (26 %), and despite a high proportion of Microtus voles in the owls’ diet (47 %), no relationship was found between their availability in the field and proportion in the diet; (iii) the proportion of Apodemus mice was negatively correlated to the proportion of Microtus voles, Sorex shrews and birds in the diet, but no similar relationship was detected for Microtus voles; (iv) the reproductive output of Tengmalm’s owls was positively correlated to the proportion of Apodemus mice in the diet, as well as to Apodemus mice and Microtus vole availability in the field; and (v) diet diversity and diet overlap were not significantly affected by the abundance of Apodemus mice and Microtus voles. Therefore, the validity of these main optimal diet theory predictions was not confirmed, especially for Microtus vole prey, due to an opportunistic choice between Apodemus mice and Microtus voles. We suggest that the reproductive output of nocturnal raptors in Central Europe may be less dependent on Microtus vole supply than that of their northern counterparts.     

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.