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Animals perceive human activities as risky and generally respond with fear‐induced proactive behaviors to buffer the circadian patterns of lethal and nonlethal disturbances, such as diel migrations (DMs) between risky places during safe nighttime and safer places during risky daytime. However, such responses potentially incur costs through movement or reduced foraging time, so individuals should adjust their tolerance when human activities are harmless, through habituation. Yet this is a challenging cognitive task when lethal and nonlethal risks co‐occur, forming complex landscapes of fear. The consequences of this human‐induced complexity have, however, rarely been assessed. We studied the individual DM dynamics of chamois (Rupicapra rupicapra rupicapra), 89 GPS‐tracked individual‐years, from/to trails in the French Alps in areas with co‐occurring lethal (hunting) and nonlethal (hiking and skiing) disturbances, with different intensities across seasons. We developed a conceptual framework relying on the risk‐disturbance hypothesis and habituation to predict tolerance adjustments of chamois under various disturbance contexts and across contrasted seasonal periods. Based on spatial and statistical analyses combining periodograms and multinomial logistic models, we found that DM in relation to distance to a trail was a consistent response by chamois (~85% of individuals) to avoid human disturbance during daytime, especially during the hiking and hunting periods. Such behavior revealed a low tolerance of most chamois to human activities, although there was considerable interindividual heterogeneity in DM. Interestingly, there was an increased tolerance among the most disturbed diel migrants, potentially through habituation, with chamois performing shorter DMs in areas highly disturbed by hikers. Crucially, chamois that were most human‐habituated during the hiking period remained more tolerant in the subsequent harvesting period, which could increase their risk of being harvested. In contrast, individuals less tolerant to hiking performed longer DMs when hunting risk increased, and compared to hiking, hunting exacerbated the threshold distance to trails triggering DMs. No carryover effect of hunting beyond the hunting period was observed. In conclusion, complex human‐induced landscapes of fear with co‐occurring disturbances by nature‐based tourism and hunting may shape unexpected patterns of tolerance to human activities, whereby animal tolerance could become potentially deleterious for individual survival. [ABSTRACT FROM AUTHOR]

Understanding the consequences of global change for animal movement is a major issue for conservation and management. In particular, habitat fragmentation generates increased densities of linear landscape features that can impede movements.While the influence of these features on animal movements has been intensively investigated, they may also play a key role at broader spatial scales (e.g. the home range scale) as resources, cover from predators/humans, corridors/barriers or landmarks. How space use respond to varying densities of linear features has been mostly overlooked in large herbivores, in contrast to studies done on predators. Focusing on large herbivores should provide additional insights to understand how animals solve the trade‐off between energy acquisition and mortality risk.Here, we investigated the role of anthropogenic (roads and tracks) and natural (ridges, valley bottoms and forest edges) linear features on home range features in five large herbivores. We analysed an extensive GPS monitoring database of 710 individuals across nine populations, ranging from mountain areas mostly divided by natural features to lowlands that were highly fragmented by anthropogenic features.Nearly all of the linear features studied were found at the home range periphery, suggesting that large herbivores primarily use them as landmarks to delimit their home range. In contrast, for mountain species, ridges often occurred in the core range, probably related to their functional role in terms of resources and refuge. When the density of linear features was high, they no longer occurred predominantly at the home range periphery, but instead were found across much of the home range. We suggest that, in highly fragmented landscapes, large herbivores are constrained by the costs of memorising the spatial location of key features, and by the requirement for a minimum area to satisfy their vital needs.These patterns were mostly consistent in both males and females and across species, suggesting that linear features have a preponderant influence on how large herbivores perceive and use the landscape. [ABSTRACT FROM AUTHOR]

Most habitats are distributed heterogeneously in space, forcing animals to move according to both habitat characteristics and their needs for energy and safety. Animal space use should therefore vary according to habitat characteristics, a process known as the "functional response" in habitat selection. This response has often been tested vis-à-vis the proportion of a habitat category within areas available to individuals. Measuring sought-after resources in landscape where they are continuously distributed is a challenge and we posit here that both the mean availability of a resource and its spatial variation should be measured. Accordingly, we tested for a functional response in habitat selection according to these two descriptors of the resource available for a mountain herbivore. We hypothesized that selection should decrease with mean value of resources available and increase with its spatial variation. Based on GPS data from 50 chamois females and data on the actual foodscape (i.e. distribution of edible-only biomass in the landscape), we estimated individual selection ratio (during summer months) for biomass at the home range level, comparing edible biomass in individual home ranges and the mean and standard deviation of edible biomass in their available range. Chamois being a group-living species, available accessible ranges were shared by several individuals that formed socio-spatial groups (clusters) in the population. As expected, selection ratios increased with the standard deviation of edible resources in each cluster, but unlike our prediction, was unrelated to its mean. Selection of areas richer in resources hence did not fade away when more resources were available on average, a result that may be explained by the need for this capital breeder species to accumulate fat-reserve at a high rate during summer months. Low spatial variation could limit the selection of chamois, which highlights the importance of resource distribution in the process of habitat selection. [ABSTRACT FROM AUTHOR]

Recent advances in animal ecology have enabled identification of certain mechanisms that lead to the emergence of territories and home ranges from movements considered as unbounded. Among them, memory and familiarity have been identified as key parameters in cognitive maps driving animal navigation, but have been only recently used in empirical analyses of animal movements., At the same time, the influence of landscape features on movements of numerous species and on space division in territorial animals has been highlighted. Despite their potential as exocentric information in cognitive maps and as boundaries for home ranges, few studies have investigated their role in the design of home ranges of non-territorial species., Using step selection analyses, we assessed the relative contribution of habitat characteristics, familiarity preferences and linear landscape features in movement step selection of 60 GPS-collared Mediterranean mouflon Ovis gmelini musimon × Ovis sp. monitored in southern France. Then, we evaluated the influence of these movement-impeding landscape features on the design of home ranges by testing for a non-random distribution of these behavioural barriers within sections of space differentially used by mouflon., We reveal that familiarity and landscape features are key determinants of movements, relegating to a lower level certain habitat constraints (e.g. food/cover trade-off) that we had previously identified as important for this species. Mouflon generally avoid crossing both anthropogenic (i.e. roads, tracks and hiking trails) and natural landscape features (i.e. ridges, talwegs and forest edges) while moving in the opposite direction, preferentially toward familiar areas. These specific behaviours largely depend on the relative position of each movement step regarding distance to the landscape features or level of familiarity in the surroundings. We also revealed cascading consequences on the design of home ranges in which most landscape features were excluded from cores and relegated to the peripheral areas., These results provide crucial information on landscape connectivity in a context of marked habitat fragmentation. They also call for more research on the role of landscape features in the emergence of home ranges in non-territorial species using recent methodological developments bridging the gap between movements and space use patterns. [ABSTRACT FROM AUTHOR]

The balance between food and perceived predation risk has been revealed as one of the primary drivers of animal habitat selection. However, few studies have investigated how spatiotemporal scales and movement/activity patterns shape responses to this food/cover trade-off while accounting for individual characteristics (e.g. sex) and for variation in predation risk (e.g. hunting) and in resource abundance/quality. We hence studied temporal changes in habitat selection of 30 GPS-collared females and 15 males of Mediterranean mouflon, Ovis gmelini musimon × Ovis sp., at two scales, i.e. 48 h home range selection within a subpopulation area (broad scale) and choice of movement steps (defined as the linear segment between two consecutive locations) according to activity state (fine scale), in southern France. During the hunting-free/food-abundant period, males selected at both scales the foraging habitats providing the best conditions for optimizing their future reproductive success and only selected areas perceived as safe during inactive steps. During the corresponding lambing period, and at both scales, females selected areas perceived as safe that should optimize lamb survival. They switched to the best foraging habitats only when lambs were weaned and only for active steps. By contrast, during hunting, when food was also scarce, both sexes selected home ranges with high proportions of the habitats perceived as safe, in which they performed all their activities. This result suggested that risk avoidance exceeded all the other individual and environmental factors in the hierarchy of the determinants of habitat selection during the hunting period. Coupling scale-specific habitat selection and activity patterns was hence decisive in disclosing how individuals fulfil their specific needs under seasonally changing levels of habitat attributes important for fitness. [ABSTRACT FROM AUTHOR]

Human disturbance is of growing concern owing to the increase of human activities in natural areas. Animal responses are manifold (immediate and/or delayed, short and/or long-lasting, with numerous behaviors affected) so that comprehensive studies are few. Here, we contrasted days with low or high tourism or hunting pressures to assess direct (daytime) and indirect (nighttime) responses of 66 GPS-collared Mediterranean mouflon Ovis gmelini musimon × Ovis sp. from the Caroux-Espinouse massif (southern France) in terms of movement, habitat use and daily activity. We took advantage of the fact that both human activities occurred during different periods and with different intensities in 3 contiguous areas (among which a protected area without hunting and with limited tourism) to compare their influence on mouflon behavior. Mouflon response to tourism was limited to the area where tourism pressure was intense with a decrease in diurnal activity compensated during nighttime by an increase of nocturnal activity. Hunting had marked consequences in the two hunted areas, with a similar shift in activity between day and night, a decrease in movement sinuosity during daytime by females and an increase in nocturnal use of the best foraging habitats by males, all suggesting an increase in foraging activities during nights following disturbance. The diurnal activity of mouflon living in the protected area was also modified during hunting period, but without nocturnal compensation. These findings revealed that the impact of hunting was higher than tourism, with several components of animal behavior affected. This calls for further research on hunting side-effects in terms of disturbance, especially as it occurs during both the adverse climatic season and the breeding period. [ABSTRACT FROM AUTHOR]

Temporal partitioning of daily activities between species may promote coexistence within animal communities by reducing behavioural interference, particularly when species highly overlap in the use of space and resources. Such a strategy may be used by Alpine chamois ( Rupicapra rupicapra rupicapra) when in the presence of mouflon ( Ovis gmelini musimon × Ovis sp.), an introduced highly gregarious species with a broader ecological niche, overlapping with that of chamois. Using simultaneous monitoring of 29 Global Positioning System-collared chamois and 12 mouflon, we assessed the temporal variation in activity patterns of chamois amongst two subpopulations: one without mouflon and one with mouflon, during January and August, which are the two most extreme periods of spatial overlap of mouflon with chamois distribution. Substantial differences in activity patterns between chamois and mouflon were observed (mean 13.8 ± 10.5% in January and 10.6 ± 11.6% in August). More subtle differences appeared between both subpopulations of chamois and persisted, regardless of the spatial overlap with mouflon (3.2 ± 1.8% in January and 2.6 ± 1.5% in August), thus highlighting that there is no behavioural interference from mouflon on chamois. Our findings suggest that the temporal partitioning of daily activities between chamois and mouflon, although probably a result of species-specific adaptations to environmental conditions, may contribute to their coexistence. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 621-626. [ABSTRACT FROM AUTHOR]