Use of winter habitat by roe deer at a northern latitude where Eurasian lynx are present

Winter climate at northern latitudes is a challenge to small-bodied ungulates, and they modify behaviour to save energy and to increase the likelihood of survival. Also, the ongoing expansion of large carnivores in several European countries can lead to the recovery of (potentially energetically costly) anti-predator behaviours. In an area recently recolonized by Eurasian lynx Lynx lynx , we snow-tracked radio-collared roe deer Capreolus capreolus in order to investigate their bedding and feeding behaviour during winter, and assess how environmental factors affect their habitat use. We also tested the prediction that roe deer use more open sites than locally available in areas with a stalking predator such as the lynx. Our results showed that both bed sites and foraging sites had more cover, compared with random sites. Most of the variation in canopy cover and in the distance and foraging sites between bed sites and foraging sites was explained by prevailing weather. As the winter progressed, the presumed depletion of fat reserves promoted the use of more canopy cover at foraging sites by night, less by day and a decrease in the distance between beds, foraging sites and human activities. Males used artificial feeding sites less often and bedded further from humans than females. The data fit the hypothesis of tighter energy budgets for family groups (females with fawns) or that males are more cautious towards humans. There was no support for the hypothesis that roe deer used more open habitat than locally available in order to reduce their vulnerability to lynx predation. Owing to severe winter conditions and the danger of starvation, roe deer seem to be forced to accept a high risk when predators are present, not changing their main pattern of habitat use from comparative areas where predators are absent.     

Predator and heterospecific stimuli alter behaviour in cattle

Wild and domestic ungulates modify their behaviour in the presence of olfactory and visual cues of predators but investigations have not exposed a domestic species to a series of cues representing various predators and other ungulate herbivores. We used wolf (Canis lupus), mountain lion (Puma concolor), and mule deer (Odocoileus hemionus) stimuli (olfactory and visual), and a control (no stimuli) to experimentally test for differences in behaviour of cattle (Bos taurus) raised in Arizona. We measured (1) vigilance, (2) foraging rates, (3) giving up density (GUD) of high quality foods and (4) time spent in high quality forage locations in response to location of stimuli treatments. In general, we found a consistent pattern in that wolf and deer treatments caused disparate results in all 4 response variables. Wolf stimuli significantly increased cattle vigilance and decreased cattle foraging rates; conversely, deer stimuli significantly increased cattle foraging rate and increased cattle use of high quality forage areas containing stimuli. Mountain lion stimuli did not significantly impact any of the 4 response variables. Our findings suggest that domestic herbivores react to predatory stimuli, can differentiate between stimuli representing two predatory species, and suggest that cattle may reduce antipredatory behaviour when near heterospecifics.     

Behavioural and spatial adaptation of the Eurasian lynx to a decline in prey availability

The distribution and abundance of food resources is a major factor influencing animal populations. I studied the effect of a roe and red deer population decline on diet composition, home range size and foraging pattern in the Eurasian lynxLynx lynx (Linnaeus, 1758) in the Bia?owie?a Primeval Forest (BPF), eastern Poland. The population of cervids in BPF experienced a nearly two-fold reduction in size from 1991 through 2006 due to severe hunting pressure between 1991 and 1996. Comparison of published data on lynx diet during the high abundance of ungulates with new data obtained for the low abundance period showed that despite a significant decline in their availability, cervids (roe and red deer) continued to form the majority of the diet of lynx, with roe deer being most preferred in both periods. Home range sizes of lynx showed a tendency to increase with declining prey densities, as indicated by relative percentage increases in average yearly home range sizes amongst different sex/age groups. In response to lower availability of their main prey, lynx increased their daily straight-line movement distances by 44% and doubled the ranges covered in 5-day periods. This illustrated that, with declining prey abundance, the lynx increased their hunting efforts by either spending more time actively searching for prey or continuing foraging even after a successful hunt. Spatial analysis of the distribution of ungulates and lynx indicated that deer were evenly distributed throughout lynx ranges in BPF and spatial proximity of the predator to prey sites did not play an important role in the efficiency of hunting. Lynx may adapt to changing prey availability by increasing search effort, but this was not sufficient to prevent the negative influences of the prey decline on the lynx population. Prey depletion has an immediate effect on lynx spatial organization and, in consequence, on their density. This information has to be considered in prioritizing lynx conservation measures and management of ungulates.     

What Cues Do Ungulates Use to Assess Predation Risk in Dense Temperate Forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.     

Seeing a Ghost? Vigilance and Its Drivers in a Predator‐free World

Vigilance is a key to the early detection of predators, but may be costly if it impairs foraging efficiency. Hence, we would expect vigilance to be suppressed and/or counter‐selected in predator‐free environments, although this might depend on the environmental drivers influencing perceived predation risk. We studied vigilance in two populations of Sitka black‐tailed deer (Odocoileus hemionus sitkensis) on Haida Gwaii (Canada) which have not been exposed to predators since they colonized the study islands approx. 60 yr ago. In this context, anti‐predator behavior should not have any obvious current benefit. Moreover, its maintenance should be particularly costly in our study populations because these deer have depleted their food resources and, thus, anti‐predator behaviors should interfere with time spent searching for scarce resources. We used bait stations equipped with camera traps to assess vigilance under standardized feeding conditions. We expected to observe lower vigilance levels than those observed elsewhere in locations with predators. We investigated how vigilance varied in relation to the amount of bait, the level of visibility, and between day and night. During the day, deer spent, on average, 14% of their time in overt vigilance during foraging bouts, a level similar to, although in the lower range of, values reported at sites where predators are present. Levels of vigilance were lower at night, and decreased with increasing visibility, but not during the day. Deer were less vigilant when bait availability was high, but only when visibility was also high. We discuss why the maintenance of vigilance is here best explained by the ghosts of predators past, and how, at the temporal scale of a few generations, the ecological factors driving vigilance levels might override the absence of significant risk from large predators.     

To scan or not to scan? Occurrence of the group‐size effect in a seasonally nongregarious forager

Group‐living requires a compromise between safety and direct/indirect costs for individuals. The larger is the group, the greater is the collective vigilance, leading to a greater net food intake per forager because of the time saved individually from scanning behaviour. In turn, individual alertness usually decreases with increasing group size (“group‐size effect”). Information on the occurrence of group‐size effect is still unclear. Previous studies have shown that it may fail to occur or even reverse, for example when costs of interference between conspecifics are high. In turn, assessing whether the group‐size effect would occur in weakly or seasonally gregarious species may help to understand its drivers. We evaluated the occurrence and the extent of group‐size effect in a seasonally nongregarious herbivore, the roe deer Capreolus capreolus. We examined the roles of sex/age class and season as drivers of vigilance behaviour. In roe deer, the group‐size effect did not depend on sex/age class: time spent foraging increased with increasing group size; in turn, vigilance increased with decreasing group size, in all sex/age classes. Females with fawns were the most vigilant sex/age class, thus revealing the cost of offspring protection. Accordingly, the higher spring vigilance levels of does could be related to reproductive costs (e.g., defence of newborn fawns). Conversely, the greater summer vigilance of bucks could result from patrolling/defence of territories. Both adult males and females also showed the higher vigilance in winter, likely because of an increase in the perception of predation risk and/or, possibly, hormones linked to an increase in intolerance of conspecifics, in males. However, the group‐size effect occurred in all the seasons, for adult males and females. Our findings suggest that foraging benefits provided by the group‐size effect may have overcome costs of group‐living, even in a weakly gregarious forager.     

Relationship between prey availability and population dynamics of the Eurasian lynx and its diet in northern Belarus

The diet and population dynamics of the Eurasian lynxLynx lynx Linnaeus, 1758 as well as an index of its main prey abundance were studied in transitional mixed forests of northern Belarus in 1985-2004. Monitoring of the lynx population and its main potential prey (the mountain hareLepus timidus, and the roe deerCapreolus capreolus) was done by snow-tracking. Also, abundance of tetraonids (Tetraonidae) was monitored by sight count. Hare numbers were fairly stable during the study period, whereas density of the roe deer population markedly increased, and tetraonids decreased. Composition of the lynx diet was stable seasonally. Lynx fed mostly on hares, roe deer and birds (usually tetraonids) year-round. However, the share of roe deer in lynx diet increased significantly during the period of its higher abundance and the share of tetraonids decreased with their decreasing numbers. There was also a remarkable increase of lynx population, which followed that of the roe deer, despite the pronounced decline of tetraonids. The results of the study emphasised the importance of roe deer as a prey of the Eurasian lynx.     

Effectiveness of odour repellents on red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) and roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>): a field test

Chemical repellents are promoted as a method to reduce ungulate–vehicle collisions and ungulate browsing damages to agricultural and forestry resources. We tested the effectiveness of two odour repellents (Mota FL and Wolf Urine) on the foraging behaviour and area avoidance of free-ranging roe deer (Capreolus capreolus) and red deer (Cervus elaphus). The effects of the repellents were assessed by comparing deer visitation rates to sand arenas before and after application of repellents and visitation rates to control arenas. Neither of the tested products reduced deer visitation rates. Rapid habituation to olfactory stimuli and lack of sensitivity to predator odours may explain the ineffectiveness of the repellents to alter the behaviour of the deer. The results indicate that the tested products have no effects on roe deer and red deer behaviour and suggest that the effectiveness of the chemical area repellents as a measure to reduce deer–vehicle collision risk and browsing damages is questionable.     

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

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

Large Impact of Eurasian Lynx Predation on Roe Deer Population Dynamics

The effects of predation on ungulate populations depend on several factors. One of the most important factors is the proportion of predation that is additive or compensatory respectively to other mortality in the prey, i.e., the relative effect of top-down and bottom-up processes. We estimated Eurasian lynx (Lynx lynx) kill rate on roe deer (Capreolus capreolus) using radio-collared lynx. Kill rate was strongly affected by lynx social status. For males it was 4.85 ± 1.30 S.E. roe deer per 30 days, for females with kittens 6.23 ± 0.83 S.E. and for solitary females 2.71 ± 0.47 S.E. We found very weak support for effects of prey density (both for Type I (linear) and Type II (non-linear) functional responses) and of season (winter, summer) on lynx kill rate. Additionally, we analysed the growth rate in a roe deer population from 1985 to 2005 in an area, which lynx naturally re-colonized in 1996. The annual roe deer growth rate was lower after lynx re-colonized the study area, but it was also negatively influenced by roe deer density. Before lynx colonized the area roe deer growth rate was λ = 1.079 (± 0.061 S.E.), while after lynx re-colonization it was λ = 0.94 (± 0.051 S.E.). Thus, the growth rate in the roe deer population decreased by Δλ = 0.14 (± 0.080 S.E.) after lynx re-colonized the study area, which corresponded to the estimated lynx predation rate on roe deer (0.11 ± 0.042 S.E.), suggesting that lynx predation was mainly additive to other mortality in roe deer. To conclude, this study suggests that lynx predation together with density dependent factors both influence the roe deer population dynamics. Thus, both top-down and bottom-up processes operated at the same time in this predator-prey system.     

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

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

Vulnerability of Domestic Sheep to Lynx Depredation in Relation to Roe Deer Density

Abstract The Eurasian lynx (Lynx lynx) causes large loss of free-ranging domestic sheep in Norway. We tested whether the observed higher kill rates by male lynx than female lynx were related to an association between the availability of the main natural prey, as measured by roe deer (Capreolus capreolus) habitat suitability, presence of sheep, and habitat selection of male and female lynx. We found that lynx selected areas with high roe deer suitability during summer and winter. Moreover, during summer, compared to male lynx, females had greater selection for roe deer areas and a stronger avoidance for sheep grazing areas, which suggests that previously observed differences in kill rates between male and female lynx can be attributed to sex-specific habitat use during summer. The connection between lynx habitat use and roe deer also was reflected in a positive relationship between the roe deer suitability of a sheep grazing area and the total loss of lambs, which suggests that livestock, rather than being actively selected, are mainly killed by lynx incidentally when encountered during other lynx activities (e.g., searching for natural prey species). Therefore, any management practice that separates lynx and sheep, such as concentrating livestock into small patches or less preferred habitats, may reduce depredation.     

What shapes Eurasian lynx distribution in human dominated landscapes: selecting prey or avoiding people?

In the multi-use landscape of southern Norway, the distribution of lynx is likely to be determined both by the abundance of their favoured prey – the roe deer – and the risk associated with the presence of humans because most lynx mortalities are caused by humans (recreational harvest, poaching, vehicle collisions). We described the distribution of the reproductive portion of the lynx population based on snow-track observations of females with dependent kittens collected over 10  yr (1997–2006) in southern Norway. We used the ecological-niche factor analysis to examine how lynx distribution was influenced by roe deer, human activity, habitat type, environmental productivity and elevation. Our first prediction that lynx should be found in areas of relatively high roe deer abundance was supported. However, our second prediction that lynx should avoid human activity was rejected, and lynx instead occupied areas more disturbed in average than those available (with the exception of the most densely occupied areas). Lynx, however, avoided the most disturbed areas and our third prediction of a trade-off between abundance of prey and avoidance of human activity was supported. On the one hand, roe deer in the most disturbed areas benefit to a large extent from current human land use practices, potentially allowing them to escape predation from lynx. On the other hand, the situation is not so favourable for the predators who are restricted in competition refuges with medium to low prey densities. The consequence is that lynx conservation will have to be achieved in a human modifed environment where the potential for a range of conflicts and high human-caused mortality will remain a constant threat.     

Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest

Human harvest is the most important mortality factor for wild ungulates in Europe and can affect several aspects of ungulate biology. There is a growing concern about possible negative side effects of human harvest. To better understand the differences between human and natural mortality, we compared the extent, age and sex structure, nutritional condition, spatial and temporal distribution of human harvest, and natural predation by the Eurasian lynx Lynx lynx on the European roe deer Capreolus capreolus, the most abundant wild ungulate in Europe. Compared to the human harvest, lynx were less likely to kill fawns and yearlings than adults, and among adult deer, lynx were more likely to kill females. The proportion of roe deer with fat-depleted bone marrow was higher among lynx prey than among harvested animals. Average lynx kill rate was estimated to 47.8 roe deer per year, and lynx predation was considerably lower than the human harvest in the same area. While human harvest increased with higher roe deer density, lynx predation was similar across the gradient of roe deer densities. Comparison with other countries indicated that differences between human harvest and natural mortality of ungulates vary considerably in different parts of Europe. Variation in hunting practices and, even more importantly, carnivore predation may have an important role in buffering unwanted side effects of harvest of wild ungulates.     

Patterns of Lynx Predation at the Interface between Protected Areas and Multi-Use Landscapes in Central Europe

In Central Europe, protected areas are too small to ensure survival of populations of large carnivores. In the surrounding areas, these species are often persecuted due to competition with game hunters. Therefore, understanding how predation intensity varies spatio-temporally across areas with different levels of protection is fundamental. We investigated the predation patterns of Eurasian lynx (Lynx lynx) on roe deer (Capreolus capreolus) and red deer (Cervus elaphus) in both protected areas and multi-use landscapes of the Bohemian Forest Ecosystem. Based on 359 roe and red deer killed by 10 GPS-collared lynx, we calculated the species-specific annual kill rates and tested for effects of season and lynx age, sex and reproductive status. Because roe and red deer in the study area concentrate in unprotected lowlands during winter, we modeled spatial distribution of kills separately for summer and winter and calculated-the probability of a deer killed by lynx and-the expected number of kills for areas with different levels of protection. Significantly more roe deer (46.05–74.71/year/individual lynx) were killed than red deer (1.57–9.63/year/individual lynx), more deer were killed in winter than in summer, and lynx family groups had higher annual kill rates than adult male, single adult female and subadult female lynx. In winter the probability of a deer killed and the expected number of kills were higher outside the most protected part of the study area than inside; in summer, this probability did not differ between areas, and the expected number of kills was slightly larger inside than outside the most protected part of the study area. This indicates that the intensity of lynx predation in the unprotected part of the Bohemian Forest Ecosystem increases in winter, thus mitigation of conflicts in these areas should be included as a priority in the lynx conservation strategy.     

Foraging of lynxes in a managed boreal-alpine environment

Foraging of Eurasian lynxes Lynx lynx was studied with telemetry and snow tracking in central Norway. In all habitats and at all seasons, medium-sized ungulates (roe deer Capreolus capreolus , reindeer Rangifer tarandus and domestic sheep Ovis aries) dominated the diet (81% of ingested biomass estimated from faeces). Mountain hares Lepus timidus and galliform birds comprised the remainder of the diet (15% and 3%, respectively). Lynxes with different life history status did not differ in prey choice, but adult males utilised carcasses of ungulate prey considerably less (16% of the edible parts) than did females with offspring (80%) and subadults (58%.). Forest habitats in lowlands and adjacent to cultivated fields were the most favourable foraging habitats (indexed as the prey encounter rate per km lynx track) primarily owing to the presence of roe deer. Two family groups tracked in winter killed 0.2 ungulate per day. The importance of agricultural land as a foraging habitat and the dominance of livestock in the diet in remoter areas indicate that the lynx has responded to agriculture and reindeer husbandry during the past century by switching from smaff game to ungulates.     

Density of Wild Prey Modulates Lynx Kill Rates on Free-Ranging Domestic Sheep

Understanding the factors shaping the dynamics of carnivore–livestock conflicts is vital to facilitate large carnivore conservation in multi-use landscapes. We investigated how the density of their main wild prey, roe deer Capreolus capreolus, modulates individual Eurasian lynx Lynx lynx kill rates on free-ranging domestic sheep Ovis aries across a range of sheep and roe deer densities. Lynx kill rates on free-ranging domestic sheep were collected in south-eastern Norway from 1995 to 2011 along a gradient of different livestock and wild prey densities using VHF and GPS telemetry. We used zero-inflated negative binomial (ZINB) models including lynx sex, sheep density and an index of roe deer density as explanatory variables to model observed kill rates on sheep, and ranked the models based on their AICc values. The model including the effects of lynx sex and sheep density in the zero-inflation model and the effect of lynx sex and roe deer density in the negative binomial part received most support. Irrespective of sheep density and sex, we found the lowest sheep kill rates in areas with high densities of roe deer. As roe deer density decreased, males killed sheep at higher rates, and this pattern held for both high and low sheep densities. Similarly, females killed sheep at higher rates in areas with high densities of sheep and low densities of roe deer. However, when sheep densities were low females rarely killed sheep irrespective of roe deer density. Our quantification of depredation rates can be the first step towards establishing fairer compensation systems based on more accurate and area specific estimation of losses. This study demonstrates how we can use ecological theory to predict where losses of sheep will be greatest, and can be used to identify areas where mitigation measures are most likely to be needed.     

Stress During Adolescence Shapes Performance in Adulthood: Context‐Dependent Effects on Foraging and Vigilance

Exposure to chronic stress during adolescence can shape behaviour, cognition and physiology in adulthood, but the consequences of these long‐term changes remain unclear. Prior studies reporting altered performance following exposure to stress in adolescence have generally interpreted lasting changes as impairments. However, we have recently shown that exposure to chronic unpredictable stress during adolescence (from post‐natal days 30–70) can enhance performance in a context‐dependent manner during a foraging task. Increases in foraging performance, (previously measured by the number of rewards obtained), are often associated with trade‐offs in other behaviours, such as vigilance. Here, we examined the effect of stress exposure in adolescence on adult foraging in male Sprague Dawley rats to determine (1) whether the increase in foraging performance exhibited by animals exposed to stress in adolescence is balanced by a decrease in vigilance, and (2) whether stress in adolescence alters time allocation between foraging and vigilance behaviours in low‐ and high‐threat conditions. We found no evidence of a trade‐off between foraging and vigilance; under low‐threat conditions, rats exposed to stress in adolescence spent more time being vigilant compared with unstressed rats, suggesting that exposure to stress in adolescence enhances anticipation of threat in adulthood. Under high‐threat conditions, adolescent‐stressed and unstressed rats did not differ in foraging and vigilance behaviours. Given that we have previously found that rats exposed to stress in adolescence nearly double food intake under high‐threat, and we now show that high‐performing rats do not spend more time foraging, it appears that stress exposure in adolescence may enhance foraging efficiency (food consumed/time) under high‐threat conditions rather than time allocation between foraging and competing behaviours. We also examined the relationship, at the level of the individual, between foraging performance and foraging and vigilance behaviours. We found that changes in individual foraging performance between low‐ and high‐threat conditions were independent of behavioural changes (i.e. both highly and poorly performing rats were equally active and contacted a similar number of patches). This suggests that the ability to obtain many rewards under high‐threat conditions may be related to efficiency, rather than the frequency of foraging and effort‐related behaviours.     

Patch use, apprehension, and vigilance behavior of Nubian Ibex under perceived risk of predation

Foraging theory predicts that animals will sacrifice feedingeffort in order to reduce predation risk. Once a forager choosesa habitat, it must decide how to allocate its foraging effort.Nubian Ibex are diurnal, social, cliff-dwelling herbivores.Many of their characteristics seem to have evolved as responsesto predation risk. In order to assess the effects that perceivedrisk of predation might have on foraging behavior of free-rangingNubian Ibex in the Negev Desert, Israel, we measured giving-updensities (GUDs) in artificial food patches and used them togauge apprehension level. (Apprehension can be defined as areduction in attention devoted to performing an activity asa consequence of reallocating attention to detecting or respondingto predation risk. A forager can also be vigilant. Vigilanceis often defined as time spent scanning the surroundings withthe head up.) We also quantified time budgeting using focalobservation of individual Nubian Ibex. Habitat preferences andpatch selectivity as a measure of apprehension were considered.In particular, we tested the effect of distance from refugeon GUDs, the effect of micropatch structure on selectivity,and the effect of distance from the refuge and group size onNubian Ibex vigilance level and apprehension. Nubian Ibex allocatetheir foraging effort more toward patches closer to the escapeterrain. At the same time, Nubian Ibex are more apprehensiveat intermediate distances from the cliff edge than nearer thecliff, and their use of vigilance increases with distance fromthe cliff edge. These results suggest that Nubian Ibex may switchfrom apprehension to a more extreme behavior of vigilance atgreater distances from the refuge. This study demonstrated theuse of antipredatory behaviors, apprehension, and vigilanceby a forager. Estimating apprehension and vigilance levels ofa forager simultaneously gives a more complete and accuratepicture of how the habitat is perceived by them and combinedwith measurements of GUD allow a more accurate assessment ofhabitat quality.     

Predation processes: behavioural interactions between red fox and roe deer during the fawning season

Predation by red fox (Vulpes vulpes) is the most important mortality cause for neonatal roe deer (Capreolus capreolus) in Scandinavia. With the objective of investigating how the fox finds fawns and how antipredatory behaviour of roe deer females influences choice of hunting method, I analysed observations of interactions between red fox and roe deer females. The observations were collected over 14 years in a mixed forest/agricultural landscape in Sweden. Of 49 fox–doe encounters, the doe attacked the fox in 59%. In 90% of these attacks the fox was successfully deterred. In two observations a doe saved a fawn attacked by a fox. Two hunting methods used by the fox were discerned. In 28 cases foxes searched the ground, and in 18 cases they surveyed open areas, often from a forest edge. The latter behaviour seemed more directed at fawns and was seen leading to a capture attempt. Searching seemed less efficient and also difficult to conduct due to the aggressiveness of does. A surveying sit-and-wait type of hunting method thus appeared as the most successful. The possibility to use this method could explain why roe deer fawns are more vulnerable to fox predation in open habitats.