Wildlife populations are subjected to increasing pressure linked to human activities, which introduce multiple stressors. Recently, in addition to direct effects, it has been shown that indirect (non-lethal) effects of predation risk are predominant in many populations. Predation risk is often structured in space and time, generating a heterogeneous "landscape of fear" within which animals can minimize risks by modifying their habitat use. Furthermore, for ungulates, resource quality seems to be positively correlated with humanrelated sources of risk. We studied the trade-off between access to resources of high-quality and risk-taking by contrasting habitat use of roe deer during daytime with that during nighttime for 94 roe deer in a hunted population. Our first hypothesis was that roe deer should avoid human disturbance by modifying their habitat use during daytime compared to nighttime. Our results supported this, as roe deer mainly used open fields during nighttime, but used more forested habitats during daytime, when human disturbance is higher. Moreover, we found that diel patterns in habitat use were influenced by hunting disturbance. Indeed, the roe deer decreased their use of high-crops during daytime, an important source of cover and food, during the hunting season. The proximity of roads and dwellings also affected habitat use, since roe deer used open fields during daytime to a greater extent when the distance to these sources of disturbance was higher. Hence, our results suggest that roe deer resolve the trade-off between the acquisition of high-quality resources and risk avoidance by modifying their habitat use between day and night.

The way an individual reacts to the risk of predation or disturbance may have important consequences for its immediate and future survival. Risk is likely perceived differently by individuals in relation to among-individual differences in correlated behavioral traits, that is, syndromes or personalities. Given that animals can avoid the risk of predation/disturbance through modification of their habitat use, we might expect individuals of differing behavioral types to adopt contrasting tactics of habitat use when faced with risky or stressful situations. We studied the relationship between habitat use and among-individual variation in behavioral profile in a population of roe deer. We hypothesized that an individual's habitat use tactics should be related to their capacity to cope with risky situations. In particular, we predicted that more reactive (risk-averse) roe deer, with relatively high behavioral response and high body temperature at capture, long flight distances and which were more vigilant during feeding, should use riskier open habitat less, particularly when risk is high. We found that although the use of open habitat during the day was negatively correlated with the indices of reactivity at capture, it was also positively correlated with indices of reactivity while foraging in open habitat. Furthermore, most of the behavioral parameters we measured were intercorrelated and moderately repeatable, potentially indexing personality traits. We conclude that there is substantial interindividual variability in how individuals manage risky situations which imposes constraints on how they are able to exploit high-risk habitats, suggesting the existence of a risk management syndrome in this large herbivore.

Non-consumptive eff ects of predators result from the cost of responses to perceived risk. Prey modulate risk exposure through fl exible habitat selection at multiple scales which, in interaction with landscape constraints, determines their use of risky habitats. Identifying the relative contributions of landscape constraints and habitat selection to risk exposure is a critical fi rst step towards a mechanistic understanding of non-consumptive eff ects.Here, we provide an integrative multi-scale study of roe deer spatial responses to variable hunting pressure along a landscape gradient of open habitats and dispersed refuges. Between low-risk and high-risk periods, we investigated shifts in 1) home-range location, 2) probability of using risky habitats (between-habitat scale) and 3) distance to the nearest refuge (within-habitat scale). For 2) and 3), we disentangled the contributions of landscape constraints and habitat selection to risky habitat use.We found that when risk was high, roe deer did not shift their home-range, but generally decreased their use of risky habitats, and sometimes reduced their distance to cover (particularly older animals). Th ere was a functional response in between-habitat selection, with animals living in more open landscapes responding more than those living in landscapes with more refuges. However, individuals living in more open landscapes avoided open risky habitat less. Finally, we found that among-individual variation in risk exposure was generally, but not always, minimized by habitat selection across gradients of landscape constraints.To our knowledge, this is the fi rst study simultaneously documenting prey responses to risk at the within-habitat, between-habitat and home-range scales. Our results support the view that between-habitat selection acts at a higher hierarchical level than within-habitat selection, and provide a framework for disentangling the contributions of habitat selection and landscape constraints to risk exposure. Selection cannot always compensate for landscape constraints, indicating a need for further investigation of the processes underlying habitat selection.

Humans, as super predators, can have strong effects on wildlife behaviour, including profound modifications of diel activity patterns. Subsequent to the return of large carnivores to human‐modified ecosystems, many prey species have adjusted their spatial behaviour to the contrasting landscapes of fear generated by both their natural predators and anthropogenic pressures. The effects of predation risk on temporal shifts in diel activity of prey, however, remain largely unexplored in human‐dominated landscapes. We investigated the influence of the density of lynx Lynx lynx, a nocturnal predator, on the diel activity patterns of their main prey, the roe deer Capreolus capreolus, across a gradient of human disturbance and hunting at the European scale. Based on 11 million activity records from 431 individually GPS‐monitored roe deer in 12 populations within the EURODEER network (http://eurodeer.org), we investigated how lynx predation risk in combination with both lethal and non‐lethal human activities affected the diurnality of deer. We demonstrated marked plasticity in roe deer diel activity patterns in response to spatio‐temporal variations in risk, mostly due to human activities. In particular, roe deer decreased their level of diurnality by a factor of 1.37 when the background level of general human disturbance was high. Hunting exacerbated this effect, as during the hunting season deer switched most of their activity to night‐time and, to a lesser extent, to dawn, although this pattern varied noticeably in relation to lynx density. Indeed, in the presence of lynx, their main natural predator, roe deer were relatively more diurnal. Overall, our results revealed a strong influence of human activities and the presence of lynx on diel shifts in roe deer activity. In the context of the recovery of large carnivores across Europe, we provide important insights about the effects of predators on the behavioural responses of their prey in human‐dominated ecosystems. Modifications in the temporal partitioning of ungulate activity as a response to human activities may facilitate human–wildlife coexistence, but likely also have knock‐on effects for predator–prey interactions, with cascading effects on ecosystem functioning.

1. Restricting movements to familiar areas should increase individual fitness as it provides animals with information about the spatial distribution of resources and predation risk. While the benefits of familiarity for locating resources have been reported previously, the potential value of familiarity for predation avoidance has been accorded less attention. It has been suggested that familiarity should be beneficial for anti-predator behaviour when direct cues of predation risk are unclear and do not allow prey to identify well-defined spatial refuges. However, to our knowledge, this hypothesis has yet to be tested.2. Here, we assessed how site familiarity, measured as the intensity of use of a given location, is associated with the probability of roe deer Capreolus capreolus being killed by two predators with contrasting hunting tactics, the Eurasian lynx Lynx lynx and human hunters. While risk of human hunting was confined to open habitats, risk of lynx predation was more diffuse, with no clear refuge areas.3. We estimated cause-specific mortality rates in a competing risk framework for 212 GPS-collared roe deer in two ecologically distinct areas of Central Europe to test the hypothesis that the daily risk of being killed by lynx or hunters should be lower in areas of high familiarity.4. We found strong evidence that site familiarity reduces the risk of being predated by lynx, whereas the evidence that the risk of being hunted is linked to site familiarity was weak.5. We suggest that local knowledge about small-scale differences in predation risk and information about efficient escape routes affect an individual's ability to avoid 1330 |

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