Search

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.

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.

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.

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.

Millions of animals are killed by vehicle collisions each year. As mitigation measures, wildlife warning reflectors have become increasingly popular, although clear evidence for their effectiveness is lacking. A reason for inconclusive results in the literature may be that most previous studies on the effectiveness of wildlife warning reflectors compare animal-vehicle collision rates with and without reflectors, a setting characterised by low event rates and weak experimental control. Animal behaviour can be expected to provide a more direct evidence for a possible effect of reflectors. In this study, we analyse roe deer behaviour in the presence of a blue semicircle reflector, one of the most frequently applied wildlife warning reflectors in Germany and other parts of Europe. Behavioural response to these reflectors (classified as no reaction, vigilance, short-distance flight and long-distance flight) was recorded both under controlled experimental conditions with captive roe deer and for free-ranging roe deer at road sections with traffic occurrence. We used generalised linear mixed models (GLMMs) to test if reflector presence induced threat-related behaviour (vigilance, flight) and movement away from the reflectors. We found no significant evidence that the light stimulus emitted by reflectors was perceived as a threat or induced evasive movement. We conclude that our study provides no evidence that blue semicircle reflectors induce behaviour in roe deer that seems suitable to reduce roe deer-vehicle collisions.

Human induced land use changes negatively impact the viability of many wildlife species through habitat modifications and mortality, while some species seem to benefit from it. Roe deer (Capreolus capreolus), a wide spread ungulate increased both its abundance and range throughout Europe. This pattern is also reflected in the increasing hunting bags over the last 40 years. Such a development raises questions about the relationship between human hunting and population dynamics and, in particular, about the potential of human hunting to control related populations. We analysed and reconstructed annual hunting bags of roe deer for three federal states of northern Germany, Brandenburg, Lower Saxony and Mecklenburg West Pomerania for the years 1972 to 2011. Since 1992 the hunting bags from these three states are significantly higher than those reported for the years 1972–1991. Our reconstruction takes into consideration effects of climate variability, expressed by inter-annual changes in the North Atlantic Oscillation and impacts from rapeseed and wheat cultivation. We found that severe winters, which are indicated by negative values of the North Atlantic Oscillation during the months December–March, directly, or with a time lag of two years affect the number of deer shot. In contrast, an increase in the area used for rapeseed cultivation coincides with higher numbers of roe deer shot, with respect to the overall mean value. Consequently, we recommend that wildlife management addresses changes in large scale processes including land use pattern and climate variability.

In most studies that estimate abundance of foxes from faeces counts, scanning is done along trails and roads or along linear features such as hedges, because it is supposed that foxes defecate mainly along these structures. As a consequence, only part (i.e. trails or linear features) of the total habitat is searched and results are possibly biased if usage by foxes of these searched features is subject to spatial or temporal variation. We therefore investigated three methods for counting red fox Vulpes vulpes faeces, that differ in the shape of the sampling units: trails and two alternatives; i.e. transects and squares. We searched for faeces using these three methods in two study areas (the Upper Rhine Valley and the Black Forest valleys) at 61 study plots and found a total of 257 fox faeces. Methods for estimating abundance should ideally have high accuracy and high precision. As actual fox densities in the areas were unknown, we were unable to assess the accuracy of our sampling methods and thus focused on method precision. We fit separate negative binomial regression models for each method with the number of faeces found as the dependent variable and a set of landscape variables as possible explanatory variables. The transect method detected significant differences in the number of faeces found between the study areas and was most precise. Even though we did find more faeces with the trail method, the precision of this method was lower than that of the transect method. For the methods trail and square, variance in the number of faeces found was large in comparison to their mean. Bias caused by methods that only sample part of the habitat is not limited to faecal counts and red fox studies, but can also occur with other species and methods.