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Aim: Macroecological studies that require habitat suitability data for many species often derive this information from expert opinion. However, expert- based informa-tion is inherently subjective and thus prone to errors. The increasing availability of GPS tracking data offers opportunities to evaluate and supplement expert- based in-formation with detailed empirical evidence. Here, we compared expert- based habitat suitability information from the International Union for Conservation of Nature (IUCN) with habitat suitability information derived from GPS-tracking data of 1,498 individu-als from 49 mammal species.Location: Worldwide.Time period: 1998–2021.Major taxa studied: Forty- nine terrestrial mammal species.Methods: Using GPS data, we estimated two measures of habitat suitability for each individual animal: proportional habitat use (proportion of GPS locations within a habi-tat type), and selection ratio (habitat use relative to its availability). For each individual we then evaluated whether the GPS- based habitat suitability measures were in agree-ment with the IUCN data. To that end, we calculated the probability that the ranking of empirical habitat suitability measures was in agreement with IUCN's classification into suitable, marginal and unsuitable habitat types.Results: IUCN habitat suitability data were in accordance with the GPS data (> 95% probability of agreement) for 33 out of 49 species based on proportional habitat use estimates and for 25 out of 49 species based on selection ratios. In addition, 37 and 34 species had a > 50% probability of agreement based on proportional habitat use and selection ratios, respectively.Main conclusions: We show how GPS-tracking data can be used to evaluate IUCN habitat suitability data. Our findings indicate that for the majority of species in-cluded in this study, it is appropriate to use IUCN habitat suitability data in macro-ecological studies. Furthermore, we show that GPS-tracking data can be used to identify and prioritize species and habitat types for re-evaluation of IUCN habitat suitability data. expert opinion, GPS, habitat suitability, habitat type, habitat use, IUCN, mammals, movement, selection ratio, telemetry

Context The Complementary Habitat Hypothesis posits that animals access resources for different needs by moving between complementary habitats that can be seen as ‘resource composites’. These movements can occur over a range of temporal scales, from diurnal to seasonal, in response to multiple drivers such as access to food, weather constraints, risk avoidance and human disturbance. Within this framework, we hypothesised that large herbivores cope with human-altered landscapes through the alternate use of complementary habitats at both daily and seasonal scales. Objectives We tested the Complementary Habitat Hypothesis in European roe deer (Capreolus capreolus) by classifying 3900 habitat-annotated movement trajectories of 154 GPS-monitored individuals across contrasting landscapes. Methods We considered day-night alternation between open food-rich and closed refuge habitats as a measure of complementary habitat use. We first identified day–night alternation using the Individual Movement - Sequence Analysis Method, then we modelled the proportion of day–night alternation over the year in relation to population and individual characteristics. Results We found that day-night alternation is a widespread behaviour in roe deer, even across markedly different landscapes. Day–night alternation followed seasonal trends in all populations, partly linked to vegetation phenology. Within populations, seasonal patterns of open/closed habitat alternation differed between male and female adults, but not in juveniles. Conclusion Our results support the Complementary Habitat Hypothesis by showing that roe deer adjust their access to the varied resources available in complex landscapes by including different habitats within their home range, and sequentially alternating between them in response to seasonal changes and individual life history.

Deer (Cervidae) are key components of many ecosystems and estimating deer abundance or density is important to understanding these roles. Many field methods have been used to estimate deer abundance and density, but the factors determining where, when, and why a method was used, and its usefulness, have not been investigated. We systematically reviewed journal articles published during 2004–2018 to evaluate spatio-temporal trends in study objectives, methodologies, and deer abundance and density estimates, and determine how they varied with biophysical and anthropogenic attributes. We also reviewed the precision and bias of deer abundance estimation methods. We found 3,870 deer abundance and density estimates. Most estimates (58%) were for white-tailed deer (Odocoileus virginianus), red deer (Cervus elaphus), and roe deer (Capreolus capreolus). The 6 key methods used to estimate abundance and density were pedestrian sign (track or fecal) counts, pedestrian direct counts, vehicular direct counts, aerial direct counts, motion-sensitive cameras, and harvest data. There were regional differences in the use of these methods, but a general pattern was a temporal shift from using harvest data, pedestrian direct counts, and aerial direct counts to using pedestrian sign counts and motion-sensitive cameras. Only 32% of estimates were accompanied by a measure of precision. The most precise estimates were from vehicular spotlight counts and from capture–recapture analysis of images from motion-sensitive cameras. For aerial direct counts, capture–recapture methods provided the most precise estimates. Bias was robustly assessed in only 16 studies. Most abundance estimates were negatively biased, but capture–recapture methods were the least biased. The usefulness of deer abundance and density estimates would be substantially improved by 1) reporting key methodological details, 2) robustly assessing bias, 3) reporting the precision of estimates, 4) using methods that increase and estimate detection probability, and 5) staying up to date on new methods. The automation of image analysis using machine learning should increase the accuracy and precision of abundance estimates from direct aerial counts (visible and thermal infrared, including from unmanned aerial vehicles [drones]) and motion-sensitive cameras, and substantially reduce the time and cost burdens of manual image analysis.

International audience; Evolution should favour plasticity in dispersal decisions in response to spatial heterogeneity in social and environmental contexts. Sex differences in individual optimization of dispersal decisions are poorly documented in mammals, because species where both sexes commonly disperse are rare. To elucidate the sex-specific drivers governing dispersal, we investigated sex differences in condition dependence in the propensity and distance of natal dispersal in one such species, the roe deer, using fine-scale monitoring of 146 GPS-collared juveniles in an intensively monitored population in southwest France. Dispersal propensity increased with body mass in males such that 36% of light individuals dispersed, whereas 62% of heavy individuals did so, but there was no evidence for condition dependence in dispersal propensity among females. By contrast, dispersal distance increased with body mass at a similar rate in both sexes such that heavy dispersers travelled around twice as far as light dispersers. Sex differences in the strength of condition-dependent dispersal may result from different selection pressures acting on the behaviour of males and females. We suggest that females disperse prior to habitat saturation being reached, likely in relation to the risk of inbreeding. By contrast, natal dispersal in males is likely governed by competitive exclusion through male–male competition for breeding opportunities in this strongly territorial mammal. Our study is, to our knowledge, a first demonstration that condition dependence in dispersal propensity and dispersal distance may be decoupled, indicating contrasting selection pressures drive the behavioural decisions of whether or not to leave the natal range, and where to settle.

International audience; Studying the factors determining the sizes of home ranges, based on body mass, feeding style, and sociality level, is a long-standing goal at the intersection of ecology and evolution. Yet, how species-specific life history traits interact with different components of the landscape to shape differences in individual home ranges at within-population level has received much less attention. Here, we review the empirical literature on ungulates to map our knowledge of the relative effects of the key environmental drivers (resource availability, landscape heterogeneity, lethal and non-lethal risks) on the sizes of individual home ranges within a population and assess whether species' characteristics (body mass, diet, and social structure), account for observed variation in the responses of the sizes of individual home ranges to local environmental drivers. Estimating the sizes of home ranges and measuring environmental variables raise a number of methodological issues, which complicate the comparison of empirical studies. Still, from an ecological point of view, we showed that (1) a majority of papers (75%) supported the habitat productivity hypothesis, (2) the support for the influence of landscape heterogeneity was less pervasive across studies, (3) the response of cattle-type to variation in food availability was stronger than the response of moose-type, and (4) species-specific body mass or sociality level had no detectable effect on the level of support to the biological hypotheses. To our surprise, our systematic review revealed a dearth of studies focusing on the ecological drivers of the variation in the sizes of individual home ranges (only about 1% of articles that dealt with home ranges), especially in the later decade where more focus has been devoted to movement. We encourage researchers to continue providing such results with sufficient sample sizes and robust methodologies, as we still need to fully understand the link between environmental drivers and individual space use while accounting for life-history constraints.

International audience; 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.

Alpine large herbivores have developed physiological and behavioural mechanisms to cope with fluctuations in climate and resource availability that may become maladaptive under climate warming. We tested this hypothesis in female Alpine ibex (Capra ibex) by modelling annual and daily movement and activity patterns in relation to temperature, vegetation productivity and reproductive status based on bio‐logging data and climate change projections. In summer, ibex moved upslope, tracking the green wave. Ibex decreased diel activity sharply above a threshold temperature of 13–14°C, indicating thermal stress, but compensated behaviourally by foraging both earlier and later in the day, and by moving further upslope than on cooler days, especially reproductive females. This critical temperature will be exceeded three times as often under climate change projections. Under such scenarios, the altitudinal extent of the area will limit the available habitat providing thermal shelter, potentially impacting performance and population distribution of this emblematic mountain ungulate., We assessed the behavioral mechanisms of female Alpine ibex (Capra ibex) to cope with fluctuations in climate and resource availability that may become maladaptive under climate warming. Ibex optimized the use of resources along the Alpine altitudinal gradient by tracking the seasonal green wave. Further, females responded to temperatures exceeding 13–14°C by redistributing the foraging time throughout the day and moving upslope, especially reproductive individuals. Under climate change scenarios, increasingly hotter conditions will limit the possibility for behavioural compensation, especially in Alpine ranges with a limited altitudinal extent.

International audience; Although inter‐individual heterogeneity in many aspects of dispersal behaviour is widely reported, this key life‐history trait is predominantly modelled as a dichotomous state of philopatry versus dispersal. The increasing body of evidence for dispersal syndromes (i.e. a suite of correlated morphological, behavioural and life‐history traits associated with dispersal) implies substantial but, to date, undocumented individual heterogeneity in behavioural tactics during dispersal. Using a large sample (n = 154) of GPS monitored juvenile roe deer Capreolus capreolus, we evaluated among‐individual behavioural heterogeneity in dispersal tactics, and the individual and environmental drivers of these alternative tactics. We developed a sequential three‐stage decision tree based on space use stability, exploration events and the directionality of movement. We identified six discrete alternative behavioural tactics during the dispersal period which were characterised by different timing, amplitude and duration in movement: slightly less than half of the deer were sedentary, either ‘strictly philopatric’ or ‘explorers’, which subsequently settled on their natal range; around 40% dispersed (‘classic dispersal’), of which, one in six subsequently aborted, moving back to their natal range (‘aborted dispersal’); finally, around 15% expressed either a ‘progressive dispersal’ tactic, gradually moving away from their natal area to settle elsewhere, or a ‘multi‐range’ tactic. The propensity to express an alternative dispersal tactic was strongly influenced by an individual's local environment. In particular, when landscape heterogeneity, resource quality and human‐related disturbance in the natal range were low, individuals were 1) more likely to adopt the alternative tactics of either progressive dispersal or multi‐ranging, but 2) also more likely to abort their dispersal attempt. Our work indicates that natal dispersal is likely not a single uniform behaviour, but that individuals may adopt a variety of alternative movement tactics which are likely governed by different selection pressures, with potentially important impacts for population dynamics and functioning.

Summary Animals exhibit a diversity of movement tactics [ 1 ]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [ 2 ]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called “green-wave surfing” [ 3 , 4 , 5 ]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [ 6 ]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [ 7 ]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [ 8 , 9 ]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [ 10 ], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

Much research on large herbivore movement has focused on the annual scale to distinguish between resident and migratory tactics, commonly assuming that individuals are sedentary at the within-season scale. However, apparently sedentary animals may occupy a number of sub-seasonal functional home ranges (sfHR), particularly when the environment is spatially heterogeneous and/or temporally unpredictable. The roe deer (Capreolus capreolus) experiences sharply contrasting environmental conditions due to its widespread distribution, but appears markedly sedentary over much of its range. Using GPS monitoring from 15 populations across Europe, we evaluated the propensity of this large herbivore to be truly sedentary at the seasonal scale in relation to variation in environmental conditions. We studied movement using net square displacement to identify the possible use of sfHR. We expected that roe deer should be less sedentary within seasons in heterogeneous and unpredictable environments, while migratory individuals should be seasonally more sedentary than residents. Our analyses revealed that, across the 15 populations, all individuals adopted a multi-range tactic, occupying between two and nine sfHR during a given season. In addition, we showed that (i) the number of sfHR was only marginally influenced by variation in resource distribution, but decreased with increasing sfHR size; and (ii) the distance between sfHR increased with increasing heterogeneity and predictability in resource distribution, as well as with increasing sfHR size. We suggest that the multirange tactic is likely widespread among large herbivores, allowing animals to track spatio-temporal variation in resource distribution and, thereby, to cope with changes in their local environment. Migration · Residency · Sub-seasonal functional home range · Plasticity · Roe deer

International audience; 1. Dispersal is a key mechanism enabling species to adjust their geographic range to rapid global change. However, dispersal is costly and environmental modifications are likely to modify the cost–benefit balance of individual dispersal decisions, for example, by decreasing functional connectivity. 2. Dispersal costs occur during departure, transience and settlement, and are levied in terms of energy, risk, time and lost opportunity, potentially influencing individual fitness. However, to the best of our knowledge, no study has yet quantified the energetic costs of dispersal across the dispersal period by comparing dispersing and philopatric individuals in the wild. 3. Here, we employed animal‐borne biologgers on a relatively large sample (N = 105) of juvenile roe deer to estimate energy expenditure indexed using the vector of dynamic body acceleration and mobility (distance travelled) in an intensively monitored population in the south‐west of France. We predicted that energy expenditure would be higher in dispersers compared to philopatric individuals. We expected costs to be (a) particularly high during transience, (b) especially high in the more fragmented areas of the landscape and (c) concentrated during the night to avoid disturbance caused by human activity. 4. There were no differences in energy expenditure between dispersers and philopatric individuals during the pre‐dispersal phase. However, dispersers expended around 22% more energy and travelled around 63% further per day than philopatric individuals during transience. Differences in energy expenditure were much less pronounced during the settlement phase. The costs of transience were almost uniquely confined to the dawn period, when dispersers spent 23% more energy and travelled 112% further than philopatric individuals. Finally, the energetic costs of transience per unit time and the total distance travelled to locate a suitable settlement range were higher in areas of high road density. 5. Our results provide strong support for the hypothesis that natal dispersal is energetically costly and indicate that transience is the most costly part of the process, particularly in fragmented landscapes. Further work is required to link dispersal costs with fitness components so as to understand the likely outcome of further environmental modifications on the evolution of dispersal behaviour.

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.Résumé Les humains, en tant que ‘super-prédateurs’, peuvent avoir des effets importants sur le comportement de la faune sauvage, y compris des modifications profondes de leurs rythmes circadiens d'activité. A la suite du retour des grands carnivores dans les écosystèmes anthropisés, de nombreuses espèces proies ont ajusté leur comportement spatial à ces paysages de la peur contrastés, générés à la fois par les pressions liées aux risques anthropiques et à la présence de leurs prédateurs naturels. Les effets du risque de prédation sur les modifications temporelles des rythmes circadiens d'activité des proies restent cependant largement inconnus dans les écosystèmes dominés par l'homme. Ici, nous avons étudié l'influence de la densité de lynx Lynx lynx, un prédateur nocturne, sur les rythmes circadiens d'activité de leur proie principale, le chevreuil Capreolus capreolus, à travers un gradient de pressions anthropiques à l’échelle Européenne. Sur la base de plus de 11 million de données d'activité issues de 431 suivis individuels de chevreuils équipés de colliers GPS provenant de 12 populations au sein du réseau EURODEER (http://eurodeer.org), nous avons analysé comment le risque de prédation par le lynx, associé aux risques létaux et non-létaux des activités humaines, influence la diurnalité des chevreuils. Nous avons démontré une forte plasticité des rythmes circadiens d'activité des chevreuils en réponse aux variations spatio-temporelles du risque, et notamment face aux activités humaines. Plus particulièrement, les chevreuils diminuent leur degré de diurnalité d'un facteur de 1.37 lorsque le dérangement humain est important. La chasse accentue cet effet, puisque durant la saison de chasse les chevreuils basculent la plupart de leur activité de nuit, et dans une moindre mesure, durant l'aube également, bien que ce patron soit essentiellement variable en fonction de la densité de lynx. En effet, en présence de lynx, leur principal prédateur, les chevreuils sont relativement plus diurnes. Globalement, nos résultats révèlent une forte influence des activités humaines et de la présence de lynx sur l'ajustement des rythmes circadiens d'activité des chevreuils. Dans le contexte du retour des grands carnivores en Europe, notre étude apporte de nouvelles connaissances sur les effets des prédateurs sur la réponse comportementale de leur proie dans des écosystèmes anthropisés. La modification de la répartition temporelle de l'activité des ongulés en réponse aux activités humaines pourrait être un facteur facilitant la coexistence homme-faune sauvage, avec toutefois des conséquences autres sur les interactions prédateurs-proies et leurs effets en cascade sur le fonctionnement des écosystèmes.

The most common framework under which ungulate migration is studied predicts that it is driven by spatio–temporal variation in plant phenology, yet other hypotheses may explain differences within and between species. To disentangle more complex patterns than those based on single species/ single populations, we quantified migration vari-ability using two sympatric ungulate species differing in their foraging strategy, mating system and physiological constraints due to body size. We related observed variation to a set of hypotheses. We used GPS-collar data from 537 individuals in 10 roe Capreolus capreolus and 12 red deer Cervus elaphus populations spanning environmental gra-dients across Europe to assess variation in migration propensity, distance and tim-ing. Using time-to-event models, we explored how the probability of migration varied in relation to sex, landscape (e.g. topography, forest cover) and temporally-varying environmental factors (e.g. plant green-up, snow cover). Migration propensity varied across study areas. Red deer were, on average, three times more migratory than roe deer (56% versus 18%). This relationship was mainly driven by red deer males which were twice as migratory as females (82% versus 38%). The probability of roe deer migration was similar between sexes. Roe deer (both sexes) migrated earliest in spring. While territorial male roe deer migrated last in autumn, male and female red deer migrated around the same time in autumn, likely due to their polygynous mating system. Plant productivity determined the onset of spring migration in both species, but if plant productivity on winter ranges was sufficiently high, roe deer were less likely to leave. In autumn, migration coincided with reduced plant productivity for both species. This relationship was stronger for red deer. Our results confirm that ungulate migration is influenced by plant phenology, but in a novel way, that these effects appear to be modulated by species-specific traits, especially mating strategies.

International audience; Although individuals must generally trade off acquisition of high-quality resources against predation risk avoidance, removal of top predators by humans has resulted in many large herbivores experiencing novel conditions where their natural predators are absent. Antipredator behaviors should be attenuated or lost in such a context of relaxed predation pressure. To test this prediction, we analyzed daily and seasonal habitat selection and activity rhythm (both commonly linked to predation risk) of GPS-collared Sitka black-tailed deer (Odocoileus hemionus sitkensis Merriam, 1898) on predator-free islands (British Columbia, Canada). In marked contrast to the behavioral patterns commonly observed in populations subject to predation risk, we documented a very low day-night contrast in habitat selection. Moreover, we observed higher activity during daytime than nighttime, as expected for nonhunted populations. We also showed that resource selection was primarily driven by seasonal variations in resource availability. These results are consistent with the expected attenuation of antipredator behaviors in predation-free environments. However, we also observed marked crepuscular activity peaks, which are commonly interpreted as an antipredator response in ungulates. Our study indicates that large herbivores are able to adjust certain antipredator behaviors under relaxed selection, notably habitat selection and activity rhythm, while others persist despite the long-term absence of predators.; Les individus font face a` un compromis entre acquérir des ressources de qualité et éviter leurs prédateurs. De nombreuses populations de grands herbivores évoluent dans des environnements où leurs prédateurs naturels sont maintenant absents. Dans ces conditions, les comportements anti-prédateurs des individus devraient être amoindris, voire supprimés. Nous avons testé cette hypothèse en analysant les patrons journaliers et saisonniers de sélection des habitats et d’activité de cerfs Sitka a` queue-noire (Odocoileus hemionus sitkensis Merriam, 1898) vivant sur des îles exemptes de tout risque de prédation en Colombie Britannique (Canada). A la différence de ce qui est communément observé dans des populations soumises au risque de prédation, le contraste jour-nuit dans la sélection des habitats est peu marqué. L’activité est plus importante de jour que de nuit, comme attendu pour des populations non-chassées. De plus, la sélection des ressources est principalement influencée par les variations saisonnières et spatiales de leur disponibilité. Ces résultats sont cohérents avec une atténuation des comportements anti-prédateurs attendue dans des environnements exempts de risque de prédation. Cependant, des pics d’activité crépusculaires marqués persistent, qui sont pourtant communément interprétés comme des réponses anti-prédatrices. Notre étude montre donc qu’en l’absence de risque de prédation, certains comportements anti-prédateurs peuvent être ajustés, alors que d’autres sont maintenus.

Made available in DSpace on 2018-11-26T17:44:52Z (GMT). No. of bitstreams: 0 Previous issue date: 2018-01-26 Robert Bosch Foundation Animal movement is fundamental for ecosystem functioning and species survival, yet the effects of the anthropogenic footprint on animal movements have not been estimated across species. Using a unique GPS-tracking database of 803 individuals across 57 species, we found that movements of mammals in areas with a comparatively high human footprint were on average one-half to one-third the extent of their movements in areas with a low human footprint. We attribute this reduction to behavioral changes of individual animals and to the exclusion of species with long-range movements from areas with higher human impact. Global loss of vagility alters a key ecological trait of animals that affects not only population persistence but also ecosystem processes such as predator-prey interactions, nutrient cycling, and disease transmission. 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Fitotecn, P-7002554 Evora, Portugal Univ Evora, ICAAM, Inst Mediterranean Agr & Environm Sci, Evora, Portugal Univ Montpellier, Univ Paul Valery Montpellier, CNRS,EPHE, Ctr Ecol Fonctionnelle & Evolut,UMR 5175, F-34293 Montpellier 5, France Phillip Isl Nat Parks, Cowes, Vic, Australia Monash Univ, Sch Biol Sci, Melbourne, Vic, Australia Ohio State Univ, Dept Civil Environm & Geodet Engn, Columbus, OH 43210 USA Fiji Natl Univ, Dept Biol, POB 5529, Natabua, Lautoka, Fiji Univ Massachusetts, US Geol Survey, Massachusetts Cooperat Fish & Wildlife Res Unit, Amherst, MA 01003 USA Penn State Univ, US Geol Survey, Penn Cooperat Fish & Wildlife Res Unit, University Pk, PA 16802 USA Univ Oxford, Dept Zool, Oxford OX1 3PS, England Giraffe Conservat Fdn, POB 86099, Eros, Namibia German Primate Ctr, Behav Ecol & Sociobiol Unit, D-37077 Gottingen, Germany Tech Univ Munich, Dept Ecol & Ecosyst Management, Restorat Ecol, D-85354 Freising Weihenstephan, Germany Cary Inst Ecosyst Studies, Millbrook, NY 12545 USA Univ British Columbia, Irving K Barber Sch Arts & Sci, Unit 2, Biol, Okanagan Campus, Kelowna, BC V1V 1V7, Canada Univ Zurich, Dept Evolutionary Biol & Environm Studies, CH-8057 Zurich, Switzerland Univ Wyoming, Dept Zool & Physiol, Laramie, WY 82071 USA Univ Washington, Sch Environm & Forest Sci, Seattle, WA 98195 USA Univ Montana, Coll Forestry & Conservat, Dept Ecosyst & Conservat Sci, Wildlife Biol Program, Missoula, MT 59812 USA Bavarian Forest Natl Pk, Dept Conservat & Res, D-94481 Grafenau, Germany Albert Ludwigs Univ Freiburg, Wildlife Ecol & Management, D-79106 Freiburg, Germany Univ Toulouse, INRA, CEFS, Castanet Tolosan, France Univ Calif Davis, Anim Behav Grad Grp, Davis, CA 95616 USA Bionet Natr Onderzoek, NL-6171 EL Stein, Netherlands Univ Vet Med Vienna, Res Inst Wildlife Ecol, A-1160 Vienna, Austria Univ Coll Cork, Sch Biol Earth & Environm Sci, Cork, Ireland Univ Wyoming, US Geol Survey, Dept Zool & Physiol, Wyoming Cooperat Fish & Wildlife Res Unit, Laramie, WY USA North Carolina Museum Nat Sci, Raleigh, NC 27601 USA North Carolina State Univ, Dept Forestry & Environm Resources, Raleigh, NC 27695 USA Karatina Univ, Dept Nat Resource Management, POB 195710101, Karatina, Kenya Univ Lethbridge, Dept Psychol, Lethbridge, AB T1K 3M4, Canada Columbia Univ, Lamont Doherty Earth Observ, Palisades, NY 10964 USA Univ Valencia, Terr Vertebrates Grp, Cavanilles Inst Biodivers & Evolutionary Biol, E-46980 Valencia, Spain SUNY Stony Brook, Dept Anthropol, Stony Brook, NY 11794 USA TSG, SSC, IUCN, Rua Licuala 622,Damha 1,Campo Grande, BR-79046150 Mato Grosso Sul, Brazil IPE Inst Pesquisas Ecol Inst Ecol Res, Caixa Postal 47, BR-12960000 Sao Paulo, Brazil Univ Alicante, Dept Ciencias Ambient & Recursos Natur, Vertebrates Zool Res Grp, Alicante, Spain Embrapa Pantanal, BR-79320900 Corumba, MS, Brazil Chico Mendes Inst Conservat Biodivers, Natl Res Ctr Carnivores Conservat, BR-12952011 Atiba, SP, Brazil Univ Glasgow, Inst Biodivers Anim Hlth & Comparat Med, Glasgow, Lanark, Scotland NYU, Dept Biol, Ctr Genom & Syst Biol, New York, NY 10003 USA Univ Calif Davis, Dept Microbiol & Mol Genet, Davis, CA 95616 USA Univ Oslo, Dept Biosci, Ctr Ecol & Evolutionary Synth, NO-0316 Oslo, Norway Hebrew Univ Jerusalem, Alexander Silberman Inst Life Sci, Dept Ecol Evolut & Behav, Movement Ecol Lab, IL-91904 Jerusalem, Israel Norwegian Inst Nat Res, NO-0349 Oslo, Norway Norwegian Univ Sci & Technol NTNU, Dept Math Sci & Ctr Biodivers Dynam, N-7491 Trondheim, Norway Univ Fed Mato Grosso do Sul, Dept Ecol, BR-79070900 Campo Grande, MS, Brazil Field Museum Nat Hist, Integrat Res Ctr, Chicago, IL 60605 USA Consorzio Parco Nazl Stelvio, Bormio, Sondrio, Italy Univ Grenoble Alpes, Irstea, UR LESSEM, BP 76, F-38402 St Martin Dheres, France Univ Bayreuth, BayCEER, D-95447 Bayreuth, Germany Natl Park Schwarzwald, D-77889 Seebach, Germany Univ New South Wales, Evolut & Ecol Res Ctr, Sydney, NSW 2052, Australia Univ New South Wales, Sch Biol Earth & Environm Sci, Sydney, NSW 2052, Australia Penn Game Commiss, Harrisburg, PA 17110 USA Princeton Univ, Dept Ecol & Evolutionary Biol, Princeton, NJ 08544 USA Univ Konstanz, Dept Biol, D-78467 Constance, Germany Off Natl Chasse & Faune Sauvage, Studies & Expertise DRE, F-01330 Birieux, France Univ Haifa, Dept Evolutionary & Environm Biol, IL-3498838 Haifa, Israel Western Ecosyst Technol Inc, Laramie, WY 82070 USA Aarhus Univ, Arctic Res Ctr, DK-8000 Aarhus C, Denmark Polish Acad Sci, Inst Nat Conservat, PL-31120 Krakow, Poland Univ Porto, CIBIO InBIO Associate Lab, P-4485661 Vairao, Portugal Univ Lisbon, Inst Super Agron, InBIO Associate Lab, Ctr Appl Ecol Prof Baeta Neves, P-1349017 Lisbon, Portugal Univ Lisbon, Fac Ciencias, Ctr Ecol Evolut & Environm Changes, P-1749016 Lisbon, Portugal Univ Calif Davis, Dept Environm Sci & Policy, Davis, CA 95616 USA Jackson Hole Conservat, Jackson, WY 83001 USA Univ Vet Med Hannover, Inst Terr & Aquat Wildlife Res, D-30173 Hannover, Germany Univ Calif Santa Cruz, Environm Studies Dept, Ctr Integrated Spatial Res, Santa Cruz, CA 95060 USA Univ Pretoria, Dept Zool & Entomol, ZA-0028 Hatfield, South Africa Colorado State Univ, Dept Fish Wildlife & Conservat Biol, Ft Collins, CO 80523 USA Tatra Natl Pk, PL-34500 Zakopane, Poland Sao Paulo State Univ, Anim Biol Postgra Program, BR-15054000 Sao Jose Do Rio Preto, SP, Brazil

International audience; Routine movements of large herbivores, often considered as ecosystem engineers, impact key ecological processes. Functional landscape connectivity for such species influences the spatial distribution of associated ecological services and disservices. We studied how spatio-temporal variation in the risk-resource trade-off, generated by fluctuations in human activities and environmental conditions, influences the routine movements of roe deer across a heterogeneous landscape, generating shifts in functional connectivity at daily and seasonal time scales. We used GPS locations of 172 adult roe deer and step selection functions to infer landscape connectivity. In particular, we assessed the influence of six habitat features on fine scale movements across four biological seasons and three daily periods, based on variations in the risk-resource trade-off. The influence of habitat features on roe deer movements was strongly dependent on proximity to refuge habitat, i.e. woodlands. Roe deer confined their movements to safe habitats during daytime and during the hunting season, when human activity is high. However, they exploited exposed open habitats more freely during night-time. Consequently, we observed marked temporal shifts in landscape connectivity, which was highest at night in summer and lowest during daytime in autumn. In particular, the onset of the autumn hunting season induced an abrupt decrease in landscape connectivity. Human disturbance had a strong impact on roe deer movements, generating pronounced spatio-temporal variation in landscape connectivity. However, high connectivity at night across all seasons implies that Europe's most abundant and widespread large herbivore potentially plays a key role in transporting ticks, seeds and nutrients among habitats.

International audience; When they visit and revisit specific areas, animals may reveal what they need from their home range and how they acquire information. The temporal dimension of such movement recursions, that is, periodicity, is however rarely studied, yet potentially bears a species, population or individual‐specific signature.A recent method allows estimating the contribution of periodic patterns to the variance in a movement path. We applied it to 709 individuals from five ungulate species, looking for species signatures in the form of seasonal variation in the intensity of circadian patterns.Circadian patterns were commonplace in the movement tracks, but the amount of variance they explained was highly variable among individuals. It increased in intensity during spring and summer, when key resources were spatially segregated, and decreased during winter, when food availability was more uniformly low. Other periodicity‐inducing mechanisms supported by our comparison of species‐ and sex‐specific patterns involve young antipredator behaviour, territoriality and behavioural thermoregulation.Model‐based continuous‐time movement metrics represent a new avenue for researchers interested in finding individual‐, population‐ or species‐specific signatures in heterogeneous movement databases featuring various study designs and sampling resolutions. However, we observed large amounts of individual variation, so comparative analyses should ideally use both GPS and animal‐borne loggers to augment the discriminatory power and be based on large samples. We briefly outline potential uses of the intensity of circadian patterns as a metric for the study of animal personality and community ecology.

This study aimed to (1) identify the scale of environmental drivers of seasonal movements on the migration – residency behavior continuum in a large herbivore species and to (2) test the hypothesis that the same environmental drivers and spatio-temporal scaling should influence spatial processes in both migrants (long distance migration) and residents (short distance range shifts). We performed a comparative analysis of the influence of plant phenology and snow cover duration on seasonal movements of five partially migrating red deer populations with contrasting environmental conditions, at the seasonal range scale and at the study area scale. The five populations presented varying proportions of migrants, large gradients of migration distances and seasonal range shifts. The probability for a red deer to migrate was strongly influenced by large-scale environmental conditions, consistent with the resource heterogeneity hypothesis (high spatio-temporal scaling favors migration). Distances moved by both migrants and residents were strongly related to large-scale environmental conditions as well. We showed that similar proximal causes influenced these seasonal movements, reinforcing the idea of a continuum from migration to residency in response to seasonal environmental changes. Together, our findings suggest that global warming, by homogenizing large-scale environmental conditions, may thus decrease migratory tactics.

International audience; Non-consumptive effects of predators result from the cost of responses to perceived risk. Prey modulate risk exposure through flexible 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 first step towards a mechanistic understanding of non-consumptive effects. 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). There 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 first 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.

The seasonal activity of the Isabelline serotine bat Eptesicus isabellinus was studied in eight habitats of the Bou Hedma National Park, central Tunisia. From June 2010 to June 2011 two techniques, mist-netting and echolocation call recording, were implemented. This bat species was captured throughout the year but only at water bodies and particularly at Nouh basin. Captures peaked in late spring — early summer, and declined in winter. Pregnant females were captured from April to June, and lactating females in May and June. Echolocation calls were recorded throughout the year, but not every month in all habitats. Most activity was reported at water bodies and street lamps. Activity peaked in late spring and early autumn, and declined during winter period. Terminal buzzes were mainly detected at basins, peaking in late spring — early summer and early autumn. Our study provided further insights on the importance of water bodies which should draw particular attention in the management plan of the national park and in the whole country.

Partial migration, when only part of the population migrates seasonally while the other part remains resident on the shared range, is the most common form of migration in ungulates. Migration is often defined by spatial separation of seasonal ranges and consequently, classification of individuals as migrants or residents is usually only based on geographic criteria. However, the underlying mechanism for migration is hypothesized to be movement in response to spatiotemporal resource variability and thus, migrants are assumed to travel an “ecological distance” (ED) or shift their realized ecological niches. While ecological and geographic distances should be related, their relationship may depend on landscape heterogeneity. Here, we tested the utility of ecological niche theory to both classify migratory individuals and to understand the underlying ecological factors for migratory behavior. We developed an integrative approach combining measures in geographic and ecological niche space and used this to classify and explain migratory behavior of 71 annual roe deer (Capreolus capreolus) movement trajectories in five European study areas. First, to assess the utility of the ED concept for classifying migratory behavior, we tested whether roe deer sought the same ecological conditions year-round or moved to different ecological conditions by measuring the annual ED travelled and the seasonal niche overlap using multivariate statistics. Comparing methods to classify migrants and residents based on geographic and ecological niche space, we found that migratory roe deer switched between seasons both in geographic and in ecological dimensions. Second, we tested which seasonal ecological factors separated resident from migrant niches using discriminant analysis and which broad-scale determinants (e.g., spatiotemporal forage variation and population density) predicted migration probability using generalized linear models. Our results indicated that factors describing forage and topographic variability discriminated seasonal migrant from resident niches. Determinants for predicting migration probability included the temporal variation (seasonality) and also the spatial variability of forage patches. Last, we also found suggestive evidence for a positive relationship between population density and migration probability. By applying the ecological niche concept to the study of partial migration in ungulates, our work underlines that partial migration is a form of behavioral plasticity. Key words: behavioral plasticity; Capreolus capreolus; ecological distance; large herbivores; niche switching; Normalized Difference Vegetation Index; partial migration; realized niche; spatiotemporal variation; ungulates

International audience; Supplemental feeding for ungulates is a widespread practice in many human-dominated landscapes across Europe and North America, mainly intended to seasonally support populations. Surprisingly, little consideration was given so far to the effect of supplemental feeding on ungulate spatial ecology at a large scale, in management and conservation studies. Analyses of the main ecological drivers influencing the use of supplemental feeding sites by ungulates across a gradient of abiotic and biotic factors are currently lacking. We conducted a large-scale assessment of ecological and management drivers of use of feeding station sites in roe deer (Capreolus capreolus), a small cervid widely distributed across Europe that is particularly sensitive to winter severity. We tested four competing hypotheses by comparing the time spent at feeding station sites by 180 individual Global Positioning System-collared roe deer from nine populations spanning a wide latitudinal and altitudinal gradient. We found that roe deer used feeding station sites highly opportunistically in response to winter severity across its range. The harshest weather conditions at the northern range limit or the highest elevations provoked an intense use of feeding station sites, which typically peaked at the end of winter, in accordance with the adverse weather and nutritional condition hypotheses. Consistently, milder winters corresponded to a reduced and/or more homogeneous use of supplemental feeding. In general, intensively used feeding station sites heavily conditioned spatial behavior of roe deer. Importantly, biotic factors such as the presence of competitors decreased roe deer use of supplemental feeding station sites. Our results emphasize the importance of this human-induced alteration to resource distribution, especially in the context of the rapidly occurring climate change that is modifying resource availability for ungulate populations.

International audience; Parasite abundance has been shown to have major consequences for host fitness components such as survival and reproduction. However, although natal dispersal is a key life history trait, whether an individual's decision to disperse or not is influenced by the abundance of parasites it carries remains mostly unknown. Current and opposing hypotheses suggest that infected individuals should either be philopatric to avoid the energetic costs of dispersal (condition dependence) or disperse to escape from heavily parasitised habitats. From intensive monitoring of a roe deer population inhabiting a multi-use and spatially heterogeneous agricultural landscape, we evaluated the link between an individual's parasite abundance and its propensity to disperse, while accounting for confounding effects of body mass. Dispersal propensity generally decreased with both increasing nematode abundance and with decreasing body mass. Within the dispersing segment of the population, individuals with high nematode abundance left their natal home range later in the season than less parasitised deer. These results clearly show that parasite abundance is an important component of condition-dependent dispersal in large herbivores. However, unexpectedly, three individuals that were both heavily parasitised and of low body mass dispersed. We suggest that this 'leave it ' response to high parasite levels in the natal habitat could represent a last ditch attempt to improve reproductive prospects, constituting a form of emergency life history strategy.

International audience; Ungulate populations exhibiting partial migration present a unique opportunity to explore the causes of the general phenomenon of migration. The European roe deer Capreolus capreolus is particularly suited for such studies due to a wide distribution range and a high level of ecological plasticity. In this study we undertook a comparative analysis of roe deer GPS location data from a representative set of European ecosystems available within the EURODEER collaborative project. We aimed at evaluating the ecological factors affecting migration tactic (i.e. occurrence) and pattern (i.e. timing, residence time, number of migratory trips). Migration occurrence varied between and within populations and depended on winter severity and topographic variability. Spring migrations were highly synchronous, while the timing of autumn migrations varied widely between regions, individuals and sexes. Overall, roe deer were faithful to their summer ranges, especially males. In the absence of extreme and predictable winter conditions, roe deer seemed to migrate opportunistically, in response to a tradeoff between the costs of residence in spatially separated ranges and the costs of migratory movements. Animals performed numerous trips between winter and summer ranges which depended on factors influencing the costs of movement such as between-range distance, slope and habitat openness. Our results support the idea that migration encompasses a behavioural continuum, with one-trip migration and residence as its end points, while commuting and multi-trip migration with short residence times in seasonal ranges are intermediate tactics. We believe that a full understanding of the variation in tactics of temporal separation in habitat use will provide important insights on migration and the factors that influence its prevalence.

International audience; We studied the walking activity over the year of free-ranging adult red deer (Cervus elaphus) in a mountainous area with the aim of describing the dynamics of movement patterns at the individual level. We monitored the distance walked by two males and two females fitted with global positioning system collars to test the hypothesis that deer adopt behaviours to reduce costs of locomotion. We predicted that both sexes would travel less in winter when disadvantageous environmental conditions occurred. We also predicted that the males would (1) reduce their movement soon after the rut due to very high energy expenditure during the breeding season and (2) travel less than the females due to their larger body mass. As we expected, minimum walking activity occurred after the rut from November to February for the males and in late February for the females. The walking activity of males peaked during the rut whereas that of females decreased. But compared to males, females moved more both during winter and daylight hours. Although our study stems from just four individuals, these results and the methodology used can be inspirational for red deer research as well as for ungulate research in general.

International audience; Vole demographics are often modified close to wooded environments. Population monitoring in such settings reveals substantial edge effects, although these have seldom been quantified. This study compares the abundance of common vole Microtus arvalis (Pallas, 1778) populations at various distances from three types of wooded environment (hedgerow, copse and forest) and in two types of habitat (temporary and permanent grassland). In temporary grassland, substantial edge effects were detected along the boundaries of forests and hedgerows. At 100 m from the forest edge, voles were twice more abundant than at 50 m, four times more than at 25 m and eight times more than at 5 m. In permanent grassland, the edge effect was such that voles were practically absent up to 100 m from the forest edge. For both types of grassland, the edge effect was more pronounced for forest habitat than for hedgerows, that is, for elements with a large surface area of wooded environment. Our results suggest that landscape context may greatly affect the local abundance of voles. Given the considerable damage done by M. arvalis, the presence of wooded patches in grassland areas leads to lower vole densities in adjacent areas and this might be an alternative method to control M. arvalis as a pest species.

Accounting for spatial scale is essential for understanding habitat selection, but few studies have used spa- tial statistics to reveal the characteristic scale at which organisms respond to their environment. We studied habitat selection by GPS-tracked red deer (Cervus elaphus L., 1758) in the Pyrenees Mountains, France, by applying a geo- statistical model that compares autocorrelation of a resource between used and available sites to uncover the scale at which animals assess habitat. Using an artificial landscape, we demonstrated that the model can handle discrete habitat classes. Based on conventional hierarchical analysis, deer selected for open habitat, especially meadow, and avoided coniferous forest, more strongly at the coarse level of the home range than GPS locations. Home ranges ex- hibited generally lower autocorrelation in elevation and meadow habitat than random locations within the population range, indicative of preference for high habitat heterogeneity. Mean maximum discrepancy in autocorrelation, which was more pronounced at the level of the home range than GPS locations, occurred at 830 m for meadow habitat and at 1511 m for elevation, suggesting that red deer responded to their environment at this scale. Our study dem- onstrates how spatial statistics can serve as an instructive complement to conventional approaches to habitat selec- tion. Resume´ : L'etude de l'echelle spatiale est essentielle pour comprendre la selection de l'habitat, mais peu d'etudes ont utilisedes statistiques spatiales pour reveler l'echelle caracteristique alaquelle un organisme repond ason envi- ronnement. Nous avons etudiela selection de l'habitat par des cerfs rouges (Cervus elaphus L., 1758) suivis par GPS dans les montagnes pyreneennes, en France, en appliquant un modele geostatistique, qui compare l'autocorre´- lation d'une ressource entre les sites utilises et disponibles pour decouvrir l'echelle a laquelle les animaux evaluent leur habitat. Apartir d'un paysage artificiel, nous demontrons que ce modele peut etre generalisepour traiter des classes discretes d'habitat. D'apres une analyse hierarchique conventionnelle, les cerfs selectionnent un habitat ouvert, plus particulierement les prairies, et evitent les forets de coniferes, de facon plus marquee au niveau plus grossier du domaine vital que celui des localisations GPS. Les domaines revelent generalement une plus faible autocorrela- tion pour l'altitude et l'habitat prairie que les localisations aleatoires dans le domaine populationnel, revelateur d'une preference pour une forte heterogeneited'habitats. L'ecart moyen maximum d'autocorrelation, plus prononcea` l'echelle du domaine individuel que des localisations GPS, est de 830 m pour l'habitat prairie et de 1511 m pour l'altitude, ce qui laisse croire que le cerf reagit ason environnement a cette echelle. Notre etude demontre comment les statistiques spatiales peuvent servir de complement instructif aux approches conventionnelles de selection de l'habitat.

International audience; In this study, we sought to understand how landscape structure affects roe deer movements within their home-range in a heterogeneous and fragmented agricultural system of south-western France. We analysed the movements of 20 roe deer fitted with GPS collars which recorded their locations every 2–6 h over several months (mean = 9 months). Based on empirical observations and previous studies of roe deer habitat use, we hypothesised that roe deer should avoid buildings and roads, move preferentially along valley bottoms and through the more wooded areas of the landscape. To test these hypotheses we paired each observed movement step with 10 random ones. Using conditional logistic regression, we modelled a step selection function, which represents the probability of selecting a given step as a function of these landscape variables. The selectedmodel indicated thatmovements were influenced by all the tested landscape features, but not always in the predicted direction: our results suggested that roe deer tend to avoid buildings, roads, valley bottoms and possibly the more wooded areas (although the latter result should be interpreted with caution, as it may be influenced by a bias in the rate of GPS fix acquisition in woods). The distances to buildings and to roads were the most influential variables in the model, suggesting that the avoidance of potential sources of disturbancemay be a key factor in determining ranging behaviour of roe deer in human dominated landscapes.

International audience; To determine the spatial resolution of Global Positioning System (GPS) receiver data, rigorous testing is essential. We tested performance of the Lotek 3300 GPS collar for medium-sized mammals (Lotek Engineering, Inc., Newmarket, ON, Canada). To mimic real wildlife monitoring situations, we performed both static (stationary receiver) and mobile tests, placing the receiver collar on a dog. We compared fix locations of the mobile receiver with the actual trajectory described by a portable Trimble high-precision GPS. We determined performance in relation to habitat type and leaf cover. Location error was habitat-dependent, with the best results in open habitat and much poorer ones in forest, particularly coniferous-dominated forest. For both static and mobile tests, location accuracy was higher when the number of satellites contacted was high and when the residual positional dilution of precision (PDOP) value was low. However, location error was highly variable, even for a given PDOP value and a given number of satellites contacted. Finally, mobile collars performed less well than their static counterparts, presumably because of frequent changes of GPS position and orientation.

1. High-density populations of large herbivores are now widespread. Wildlife managers commonly attempt to control large herbivores through hunting to meet specific management objectives, considering population density as the minimal key source of information. Here, we review the problems of censusing populations of large herbivores and describe an alternative approach, employing indicators of ecological change. 2. Estimating density of large herbivores with high precision and accuracy is difficult, especially over large areas, and requires considerable investment of time, people and money. Management decisions are often made on an annual basis, informed by population changes over the previous year. However, estimating year-to-year changes in density is not a realistic goal for most large herbivores. Furthermore, population density per se provides no information on the relationship between the population and its habitat. 3. For successful management of large herbivores, we need to consider not only the fate of the population, but rather changes in both population and habitat features, as well as their interaction. Managers require information on trends in both the animal population and habitat quality in order to interpret changes in the interaction between these two compartments. 4. We propose that a set of indicators of animal performance, population abundance, habitat quality and/or herbivore habitat impact provides relevant information on the population– habitat system. Monitoring temporal changes in these indicators provides a new basis for setting hunting quotas to achieve specific management objectives. This sort of adaptive management is employed widely in France for managing roe deer Capreolus capreolus . 5. Synthesis and applications . The management of large herbivores would be improved by investing fewer resources in trying to estimate the absolute abundance of ungulates, and more resources in collecting additional data to inform understanding of the ecological status of the ungulate–habitat system being managed. This paper presents a set of indicators of ecological change for monitoring the interaction between a population and its habitat as a basis for adaptive management to attain explicit goals and to improve knowledge of the system. This approach could improve management for a variety of large herbivores, by harmonizing actions at wide spatial scales.

International audience; 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.

International audience; Despite frequent exposure to disturbances or threats, some species seem to be successful in human-dominated landscapes. In order to better understand how these species respond to disturbance linked to human activities, we studied the immediate response of roe deer to playback experiments conducted in an agricultural landscape where human activities are omnipresent and hunting is frequent. During two periods of contrasted hunting risk, we performed playback experiments (with dog barks or tractor engine noises) with experimenters that were either stationary or walked towards the deer. We modelled the probability of fleeing immediately after the start of the experiment. The probability of fleeing was around 0.3 when experimenters were located 100 m away from the deer and did not move but was greater when the experimenters were initially closer to the animal or approached closer to the animal during the experiment. Surprisingly, the responses of roe deer did not vary between periods of contrasting risk or in relation to the type of sound played. This study provides experimental evidence of a high level of reactivity of this sedentary species when confronted with disturbance. We argue that this may partly explain its success in human-dominated environments, allowing it to exploit the rich feeding resources provided by agricultural landscapes in a relatively safe way.

Summary 1 A crucial question in wildlife management concerns the definition of ecologically meaningful management units. Management action needs to be co-ordinated at the appropriate spatial scale. There are few practical tools for delimiting pertinent management units for large mammals such as ungulates. 2 Previous work has favoured a molecular approach to determine genetically distinct units, but this may be too costly for routine management. In addition, recent studies have revealed fine-scale heterogeneity in population dynamics of free-ranging ungulates in relation to environmental variability. 3 We combined spatial analysis of environmental heterogeneity (vegetation cover, topography and climate), animal morphology (jaw length) and genetic structure (microsatellites) to define biologically meaningful population units for roe deer Capreolus capreolus in the Belluno province, north-eastern Italy. Jaw length is a sensitive measure of density-dependence and a good proxy for spatial and temporal variation in roe deer population growth. 4 Spatial analysis of environmental variables suggested that the study area should be divided into two, or possibly four, biogeographical regions in relation to variation in altitude and habitat type. There was significant spatial variability in jaw length across the province that clustered into two main regions (with shorter jaws in the north compared with the south), which matched the previously defined biogeographical regions. This spatial structuring was also supported by microsatellite analysis, which revealed two genetically distinct populations, one in the north and one in the south. 5 Synthesis and applications. A multiparameter approach, combining environmental information with data on indices of density-dependence such as jaw length, could be extremely useful for defining ecologically meaningful management units. Indeed, monitoring spatial and temporal variation in jaw length could provide deer managers with a simple way to index population structure and fluctuations in time and space.

International audience; Dispersal is frequently more prevalent in one sex compared to the other. Greenwood proposed that patterns of sex-biased dispersal among birds and mammals are linked to their mating strategies. For species where males defend resources rather than females, he predicted female-biased dispersal, because males should remain at their birth site where they are familiar with the distribution of the resources that they must defend. Greenwood's hypothesis has been extensively supported among birds, where most species exhibit a resource-defence mating strategy. However, almost no equivalent information is available for mammals as males generally defend mates in this group. An exception is the European roe deer, a resource-defence mating ungulate. We thus tested Greenwood's hypothesis on this atypical mammalian model, looking for female-biased dispersal using sex-specific inter-individual genetic distances. We conclusively show that gene flow is not higher among females compared to males in the studied roe deer population, and hence that dispersal is not female-biased, suggesting that male mating strategy is not the primary selective force driving the evolution of dispersal in roe deer. We discuss the role of female mate choice and intra-sexual competition as possible alternative selective pressures involved

International audience; Breeding dispersal, defined as the net movement between successive breeding sites, remains a poorly understood and seldom reported phenomenon in mammals, despite its importance for population dynamics and genetics. In large herbivores, females may be more mobile during the breeding season, undertaking short-term trips (excursions) outside their normal home range. If fertilisation occurs, leading to gene flow of the male genome, this behaviour could be considered a form of breeding dispersal from a genetic point of view. Here, we investigated ranging behaviour of 235 adult roe deer using intensive GPS monitoring in six populations across Europe within the EURODEER initiative. We show that excursions are common from June to August among females, with 41.8 % (vs. 18.1 % of males) making at least one excursion. Most individuals performed only one excursion per season and departure dates for females were concentrated in time, centred on the rutting period, suggesting a link with reproduction. The distance females travelled during excursions was significantly greater than the site-specific average diameter of a male home range, while travel speed decreased once they progressed beyond this diameter, indicating search behaviour or interaction with other male(s) outside the resident male's territory. Because adults are normally highly sedentary, the potential for mating with relatives is substantial; hence, we conclude that rut excursions could be an alternative tactic enabling females to avoid mating with a closely related male. To understand better the ultimate drivers at play, it will be crucial to explore the genetic causes and consequences of this behaviour.

When individuals disperse, they modify the physical and social composition of their reproductive environment, potentially impacting their fitness. The choice an individual makes between dispersal and philopatry is thus critical, hence a better understanding of the mechanisms involved in the decision to leave the natal area is crucial. We explored how combinations of behavioural (exploration, mobility, activity and stress response) and morphological (body mass) traits measured prior to dispersal were linked to the subsequent dispersal decision in 77 roe deer Capreolus capreolus fawns. Using an unusually detailed multi-trait approach, we identified two independent behavioural continuums related to dispersal. First, a continuum of energetic expenditure contrasted individuals of low mobility, low variability in head activity and low body temperature with those that displayed opposite traits. Second, a continuum of neophobia contrasted individuals that explored more prior to dispersal and were more tolerant of capture with those that displayed opposite traits. While accounting for possible confounding effects of condition-dependence (body mass), we showed that future dispersers were less neophobic and had higher energetic budgets than future philopatric individuals, providing strong support for a dispersal syndrome in this species.

International audience; 1. Because many large mammal species have wide geographical ranges, spatially distant populations may be confronted with different sets of environmental conditions. Investigating how home range (HR) size varies across environmental gradients should yield a better understanding of the factors affecting large mammal ecology. 2. We evaluated how HR size of a large herbivore, the roe deer (Capreolus capreolus), varies in relation to seasonality, latitude (climate), weather, plant productivity and landscape features across its geographical range in Western Europe. As roe deer are income breeders, expected to adjust HR size continuously to temporal variation in food resources and energetic requirements, our baseline prediction was for HR size to decrease with proxies of resource availability. 3. We used GPS locations of roe deer collected from seven study sites (EURODEER collaborative project) to estimate fixed-kernel HR size at weekly and monthly temporal scales. We performed an unusually comprehensive analysis of variation in HR size among and within populations over time across the geographical range of a single species using generalized additive mixed models and linear mixed models, respectively. 4. Among populations, HR size decreased with increasing values for proxies of forage abundance, but increased with increases in seasonality, stochastic variation of temperature, latitude and snow cover. Within populations, roe deer HR size varied over time in relation to seasonality and proxies of forage abundance in a consistent way across the seven populations. Thus, our findings were broadly consistent across the distributional range of this species, demonstrating a strong and ubiquitous link between the amplitude and timing of environmental seasonality and HR size at the continental scale. 5. Overall, the variability in average HR size of roe deer across Europe reflects the interaction among local weather, climate and seasonality, providing valuable insight into the limiting factors affecting this large herbivore under contrasting conditions. The complexity of the relationships suggests that predicting ranging behaviour of large herbivores in relation to current and future climate change will require detailed knowledge not only about predicted increases in temperature, but also how this interacts with factors such as day length and climate predictability.

International audience; Natal dispersal (i.e. movements between the natal range of an individual and its first breeding site) is a complex process which can have profound impacts on population dynamics. In most species, only a proportion of juveniles actually disperse, but few empirical data are available on the factors that drive the decision to disperse. To understand the behavioural ontogeny of the decision to disperse, we investigated the ranging behaviour of 66 juvenile roe deer, Capreolus capreolus, monitored with GPS collars prior to and during the dispersal phase. We compared the number and features (distance and duration) of exploratory movements during the period prior to dispersal between philopatric individuals and dispersers. Overall, 76% of the juveniles that dispersed during their first spring performed exploratory movements prior to the dispersal period. Indeed, exploration prior to the dispersal phase was much more common among future dispersers than among future philopatric individuals, suggesting that dispersal of juveniles is facilitated to some degree by predispersal exploration forays. Furthermore, the direction of the dispersal event was correlated with the direction of previous exploration, although there was no relationship between exploration distance and subsequent dispersal distance. Thus, it appears that individuals that subsequently disperse first explore their environment, prior to definitively leaving their natal range, probably to obtain information on potential suitable adult ranges. Exploratory movements prior to dispersal can thus provide information that influences an individual's decision on whether or not to disperse and, if so, in which direction. (C) 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

International audience; Most forests in Europe are patchily distributed within the agricultural landscape. Therefore, forest biogeochemistry in Europe cannot be understood without considering the connectivity of nutrient cycles between forest patches and fertilized cropland. In this paper, we quantified the role of roe deer, the most widespread wild ungulate in Europe, as a vector of nitrogen and phosphorus from fertilized fields to forest patches, in a typical agricultural landscape of southwestern Europe. We derived a model of nutrient transfer from a data set on deer density, landscape-use by individual deer, and nutrient content in feces. The model shows that the magnitude of nutrient transfer is highly sensitive to the proportion of forest patches within the landscape, and to the way deer use the landscape to feed and defecate. Hence, the magnitude of nutrient transfer varies substantially across the landscape. Locally, deer may significantly fertilize the forest, transferring the equivalent of almost 20% of the atmospheric deposition of nitrogen, and the equivalent of 0.13% of the total stock of phosphorus from cropland to forest patches each year. These inputs may markedly influence the biogeochemistry of forests in the long run, and the nitrogen to phosphorus ratio available to trees and forest plants. These results highlight the significant, but, heterogeneous, role of wild ungulates in forest biogeochemistry across Europe.

International audience; 1. Natal dispersal is defined as the movement between the natal range and the site of first breeding and is one of the most important processes in population dynamics. The choice an individual makes between dispersal and philopatry may be condition dependent, influenced by either phenotypic attributes and/or environmental factors. 2. Interindividual variability in dispersal tactics has profound consequences for population dynamics, particularly with respect to metapopulation maintenance. A better understanding of the mechanisms underlying this variability is thus of primary interest. 3. We investigated the ranging behaviour of 60 juvenile European roe deer, Capreolus capreolus, monitored with GPS collars for 1 year prior to their first reproduction, from 2003 to 2010 in South-West France. Dispersal occurs across a spatial continuum so that dividing individuals into two categories (dispersers vs. philopatric) may lead to information loss. Therefore, to investigate condition-dependent dispersal more accurately, we developed an individual-based measure of dispersal distance, which took into account interindividual variation in ranging behaviour. We assessed the influence of body mass, the degree of habitat heterogeneity and sex on dispersal initiation date, dispersal propensity and distance. 4. The overall population dispersal rate was 0.34, with a mean +/- SD linear distance between natal and post-dispersal home ranges of 12.3 +/- 10.5 km. Dispersal distances followed a classical leptokurtic distribution. We found no sex bias in either dispersal rate or distance. Forest animals dispersed less than those living in more heterogeneous habitats. Heavier individuals dispersed with a higher probability, earlier and further than lighter individuals. Our individual-based standardised dispersal distance increased linearly with body mass, with some suggestion of a body mass threshold of 14 kg under which no individual dispersed. 5. Natal dispersal in roe deer was thus dependent on both phenotypic attributes and environmental context. Our results suggest that population connectivity can be altered by a change in average body condition and is likely higher in the rich and heterogeneous habitats typical of modern day agricultural landscapes.

International audience; Forest fragmentation may benefit generalist herbivores by increasing access to various substitutable food resources, with potential consequences for their population dynamics. We studied a European roe deer (Capreolus capreolus) population living in an agricultural mosaic of forest, woodlots, meadows and cultivated crops. We tested whether diet composition and quality varied spatially across the landscape using botanical analyses of rumen contents and chemical analyses of the plants consumed in relation to landscape metrics. In summer and non-mast winters, roe deer ate more cultivated seeds and less native forest browse with increasing availability of crops in the local landscape. This spatial variation resulted in contrasting diet quality, with more cell content and lower lignin and hemicellulose content (high quality) for individuals living in more open habitats. The pattern was less marked in the other seasons when diet composition, but not diet quality, was only weakly related to landscape structure. In mast autumns and winters, the consumption of acorns across the entire landscape resulted in a low level of differentiation in diet composition and quality. Our results reflect the ability of generalist species, such as roe deer, to adapt to the fragmentation of their forest habitat by exhibiting a plastic feeding behavior, enabling them to use supplementary resources available in the agricultural matrix. This flexibility confers nutritional advantages to individuals with access to cultivated fields when their native food resources are depleted or decline in quality (e.g. during non-mast years) and may explain local heterogeneities in individual phenotypic quality.

International audience; We investigated space-use patterns of the West African savanna buffalo (Syncerus caffer brachyceros), a little-studied subspecies occurring at the northern limit of the African buffalo's geographical range. This buffalo generally ranges in small herds (about 45 individuals) and has a low body mass (approximately 400 kg) relative to the Cape buffalo (S. c. caffer). We monitored the movements of 7 breeding herds in W Regional Park (Burkina Faso, Benin, Niger) using global positioning system collars and activity data loggers. Habitat selection was analyzed at both large (interseasonal) and small (intraseasonal) scales in a context where resources are segregated spatially at some times of year. Both biotic (primary production and vegetation types) and abiotic (timing of rainfall and surface water) covariates, and the extent to which neighboring herds shared space, were considered. In the dry season buffalo herds ranged close (within 5.3 +/- 2.0 km, mean +/- SD) to segments of permanent rivers. At the onset of the monsoon all herds but 1 (which had year-round access to suitable resources) performed a large (35 +/- 10 km) directional movement in response to a large-scale gradient of primary production. Spatiotemporal dynamics of forage and water resources thus jointly stimulated interseasonal directional movements and shaped large (335 +/- 167 km(2)) annual home ranges. Furthermore, the establishment of home ranges in the wet season appears to be conditioned by a threshold (about 10%) in the availability of perennial grasses. Habitat-selection analysis at intraseasonal scale also underlines the key role played by perennial grasses for buffaloes. The spatial arrangements of home ranges of neighboring herds also suggest that interherd behavioral avoidance is a high-level constraint on foraging processes. The ability of the African buffalo to cope with contrasting environmental conditions throughout most sub-Saharan ecosystems highlights the high behavioral plasticity of this species.

International audience

We investigated age-specific variation in male yearly breeding success (YBS) using genetic estimates obtained from 2 populations of a territorial ungulate, the European roe deer (Capreolus capreolus). YBS in both populations was markedly age-structured, with 3 distinct stages, supporting the dome-shaped pattern of variation commonly reported for age-dependent variation in life-history traits of ungulates. YBS was low at 2 years of age, peaked at 3–8 years of age, and tended to decline afterwards (senescence). Most males successfully reproduced for the 1st time at 3 years of age, which is well after their physiological maturity. The few successful young males (i.e., 2 year olds) were likely fast-growing individuals that could successfully hold a territory. The high variance in YBS and antler size for old males at Bogesund, Sweden, suggests that only some males of this age class are able to maintain large antlers and, hence, retain their territories.

International audience; 1- Territoriality is commonly associated with resource defence polygyny, where males are expected to gain access to females by anticipating how resources will influence female distribution and competing for resource-rich sites to establish their zone of dominance. 2-We tested this hypothesis in European roe deer (Capreolus capreolus) by simultaneously assessing the influence of resources on female distribution and the influence of female distribution on male distribution and breeding success using paternity analyses. 3-Females did not fully distribute themselves among male territories in relation to resources. As a result, relative female abundance in a male's territory depended on territory size, but not on its habitat quality. In turn, relative female abundance in a male's territory determined, at least partially, his breeding success. 4-Interestingly, male territory size, and hence access to females, was partly determined by male body mass (all males) and by residual antler size (subadults only). The latter result suggests that large antlers may be important to young males for establishing their first territory, which is then usually retained for all subsequent reproductive seasons. 5-To conclude, although territoriality of male roe deer has certainly evolved as a tactic for ensuring access to mates, our results suggest that it does not really conform to a conventional resource defence polygyny strategy, as males seem to gain no obvious benefit from defending a territory in an area of high habitat quality in terms of enhanced access to mates. 6-This may explain the stability of male territories between years, suggesting that male territoriality conforms to an 'always stay' and 'low risk–low gain' mating strategy in roe deer.

Locating and monitoring animals using tracking devices is a method commonly used for many taxa to study characteristics such as home-range size, habitat selection, movement patterns and other aspects of ranging behaviour. Fitting such devices requires the capture and handling of the study organism and researchers must then assume that a monitored animal behaves in a 'normal' way. We investigated whether the capture and handling of roe deer Capreolus capreolus induced behavioural alterations. In particular, we expected that the roe deer would exhibit a 'seeking a refuge and waiting before returning' strategy immediately after release, taking shelter far from the capture scene, in closed habitat, and exhibiting a reduced activity level. We evaluated the effect of capture and handling on 112 roe deer equipped with GPS collars, during a period of 50 days after release. We compared the first 10 days after release with the subsequent days for the following behavioural parameters: distance to the barycentre of their GPS fixes, presence in forest habitat, distance to the nearest forest patch, distance to a source of human disturbance, and activity level. We found pronounced differences in terms of spatial behaviour, habitat use and overall activity level between the two periods in GPS monitored roe deer. We also found differences in terms of spatial displacement between the sexes, with females responding less than males, and among age classes, with yearlings responding most and fawns least, to the capture and handling event. Finally, spatial displacement of roe deer increased with openness of the habitat due, in part, to the scarcity of available shelter in open areas. We conclude that the roe deer exhibited a strategy consisting of seeking a refuge and waiting before returning after capture, handling and fitting of a collar, with displacement towards a refuge habitat, in or near woodland, avoidance of sources of human disturbance and reduced activity levels. From a practical point of view, we recommend removing data during the first days of monitoring as behavioural alterations due to capture and handling may be pronounced.

International audience; The mortality risk from hunting/predation should increase animals' vigilance and modify their selection of feeding sites. This risk may thus be costly if vigilance interferes with feeding and/or if animals select poorer but safer feeding sites. We observed the vigilance behaviour of roe deer, Capreolus capreolus, feeding in a fragmented landscape during and outside the hunting season and compared food availability and local landscape features at these feeding sites with random paired sites. Roe deer spent more time vigilant during the hunting season than outside it. During the hunting season, vigilance decreased as the woodland extent within an 800 m radius increased, but this was not the case outside the hunting season. Vigilance decreased with increasing distance to houses, both during and outside the hunting season. When food is abundant, interference with feeding may be low because animals can simultaneously process food (chewing) and be vigilant. During the hunting season, the total time spent vigilant while chewing increased with increasing food abundance to a lesser extent than outside the hunting season, suggesting a higher level of costly exclusive vigilance during the hunting season. Outside the hunting season animals selected feeding sites that provided more food, but during the hunting season, as risk (proximity to houses) was positively correlated with food availability, animals no longer selected feeding sites on the basis of food availability. Taken together, our results indicate that roe deer trade off risk avoidance for food availability in hunted populations.

From a literature review of five wildlife ecology journals since 1937, we document how using indices to monitor ungulate body condition is common practice, with the kidney fat index (KFI = weight of fat around the kidneys/weight of kidneys without fat × 100) as the favoured tool (82% of studies). In this context, we highlight the problems of using indices when underlying statistical assumptions are not met (isometry, parallel slopes between treatments). We show, with real and simulated data for two cervids with contrasting fat storage strategies, how results from analysis of variance of KFI values differ from analysis of covariance (ANCOVA) of raw data. We conclude that the KFI is affected by the restrictions typically associated with derived index values, and as a consequence, statistical analysis of the KFI could generate spurious results leading to erroneous interpretations concerning variation in body condition of ungulate populations. Thus, we recommend analysing fat weight as an untransformed variable in ANCOVA (kidney weight as covariate) to describe body condition variation in ungulates.

Radiotracking of two roe deer Capreolus capreolus (1 female, 1 male) in a Mediterranean habitat (Matorrals) with a detailed analysis of the vegetation showed that within their annual home range the deers selected particular habitat types, according to the season. The northern slope was preferred by both animals. Comparatively to female, male had a larger annual home range (150 ha vs. 130 ha), composed of more widely scattered sites of preferred use in more seasonal vs variable habitat types. Habitat selection in different seasons was affected by several factors: i) in all seasons, abundance of Rubus ulmifolius and Rosa ssp., ii) strong winds, especially in winter, that caused the male to move to southern slope, iii) disturbance by hunting dogs in autumn that forced deers into one or two refuge zones, iv) aspect of deer biology, such as territory marking by the male and giving birth by the female, which modified not only behaviour but also use of space. Anecdotal evidence has suggested that, during the rutting period, female roe deer may undertake short excursions, outside of their normal home range, possibly to mate with a reproductive partner. To address this question, we analysed the ranging behaviour of 27 female roe deer Capreolus capreolus, equipped with GPS collars, inhabiting a fragmented landscape in France. We compared female movements during the rutting period with a non-rutting period over two summers using a recently published approach. Search intensity and home range size were significantly greater during the rutting period. The difference in home range size between the two periods was significantly greater in 2006 compared to 2005 and in open compared to closed habitat. We were not able to identify any influence of body mass on the difference in ranging behaviour between the two periods. Visual analysis of movement trajectories for 11 females revealed that 5 (45%) performed an excursion for a duration of a few hours to several days. We speculatively suggest that female rut excursions provide an opportunity for active mate choice in roe deer, where males are territorial, although we cannot rule out the alternative explanation that these movements are a means to avoid male harassment.