La dispersion chez le chevreuil européen, Capreolus capreolus, dans un paysage hétérogène

La dispersion, définie comme l'ensemble des mouvements qui éloigne les organismes de leur lieu de naissance de façon permanente, est un processus biologique fondamental de par son rôle déterminant dans la dynamique des populations. Dans la plupart des populations, tous les individus ne dispersent pas, et les individus qui dispersent ne sont pas un échantillon aléatoire de la population. Le but de cette thèse est d'appréhender les facteurs qui peuvent être une source de variabilité entre les individus dans leur comportement de dispersion au cours des différentes phases de la dispersion natale chez une espèce de grand herbivore, le Chevreuil. Dans une population de chevreuils évoluant dans un milieu hétérogène, plus de 100 individus juvéniles ont été capturés et équipés de collier GPS entre 2003 et 2012, permettant un suivi régulier de leurs déplacements pendant plus d'une dizaine de mois. Il a ainsi été mis en évidence l'importance de facteurs internes (tels que la masse corporelle des individus, leurs traits de comportement avant dispersion ou leur sexe) et externes aux individus (tel que le degré d'ouverture du paysage) au cours des différentes phases de la dispersion (émigration, trajet, immigration). Une variabilité inter-individuelle du comportement de dispersion s'observe tout au long du processus de dispersion, de la phase préparatoire à l'installation dans un domaine post-dispersion. De plus, plusieurs particularités du comportement de dispersion semblent participer à la réduction des coûts directs et indirects qui lui sont associés. Cette variabilité pouvant avoir des répercutions importantes sur le fonctionnement de la population, révèle ainsi l'importance de sa prise en compte dans les études sur la dispersion.Dispersal, defined as the movements that take individuals away from their birth site permanently, is a fundamental biological process that impact population dynamic and genetic. In most populations, not all individual disperse, and dispersers are not a random subset of the source population. The general aim of this thesis is to better understand the factors generating inter-individual variability across the three phases of the dispersal behaviour in a large herbivore specie, the European roe deer. In a heterogeneous landscape, more than 100 juveniles from a natural population were captured and equipped with a GPS collar between 2003 and 2012 and then intensively monitored during approximately ten month. This study highlights the role of internal factors (such as individual body mass, behavioural traits before dispersal or sex) and external factors (such as degree of landscape openness) across the different phases of the dispersal process (emigration, transience, immigration). A high inter-individual variability of the dispersal behaviour was observed across the whole process, from the preliminary phase to the settlement in the post-dispersal home range. Moreover, some behavioural characteristics of dispersal should reduce the direct and indirect costs associated with dispersal. Due to the important consequences on population fonctionning this inter-individual variability on dispersal behaviour may have, it seems essential to take into account this factor when studying dispersal

Implications of the forage maturation hypothesis for activity of partially migratory male and female deer

Partial migration is common in a large variety of taxa in seasonally variable environments. Understanding the mechanisms underlying migration is important, as migration affects individual fi tness. Migratory herbivores bene fi t from delayed forage maturation and hence higher food quality during migra- tion and at their summer range, termed the forage maturation hypothesis (FMH). The link between diet quality and rumination time allows migrants eating a higher quality diet to spend less time on rumination, and they can thus allocate more time to additional feeding. However, such an argument implicitly assumes that deer are energy maximizers, while studies have reported also time minimization strategies under risk of predation. Male and female distributions are limited by different factors linked to both body size differ- ences and reproductive strategies, but there is no study investigating differences in activity pattern accord- ing to the individual migratory patterns for male and female deer. We here unify the FMH with the hypotheses predicting sex-speci fi c time allocation strategies. To test predictions of sex-speci fi c activity of resident and migratory red deer ( Cervus elaphus ), we analyzed activity data of 286 individuals that were fi tted with GPS collars from a population in western Norway. While migrants were more active during the migration itself, we found no differences in activity pattern between migrant and resident deer during the main growth season, neither in terms of proportion of daily time active nor in terms of daily mean movement speed, thus rejecting that deer were energy maximizers. Overall, we found that females were more active during the main growth season even after controlling for body size differences. These patterns are consistent with patterns predicted from sexual segregation theory linked to the reproductive strategy hypothesis. Our study highlights how the understanding of migration can be advanced by considering it in the context of different reproductive strategies of males and females.publishedVersio

Island tameness and the repeatability of flight initiation distance in a large herbivore

This is the author accepted manuscript. The final version is available from NRC Research Press via the DOI in this record.Antipredator behaviours can be lost relatively quickly in populations that are relieved of predation, as is known for several species inhabiting islands. Flight initiation distance (FID) is often studied in the context of island tameness; however, little is known about the factors that influence and maintain FID variation in predation-free populations. Here, we studied FID in foals of an isolated predator-free population of feral horses (Equus caballus L., 1758) on Sable Island, Canada, to determine if FID could be used for research on consistent individual differences in risk aversion and island tameness. In addition to testing for temporal, spatial, and sex effects on FID, we compared repeatability estimates at two temporal scales (within and among days). Similar FID for measurements obtained on the same day and for males and females indicated an absence of short-term desensitization and sex effects. In contrast, FID decreased for measurements made on subsequent days and from east to west, which could reflect habituation to human presence and (or) other temporal and spatial processes. Repeatability was high (0.42 ± 0.06), but tended to decrease with increasing time intervals. This study highlights the potential of FID for individual-based research on the ecology and evolutionary dynamics of risk aversion in predation-free populations.Funding was provided by the Natural Sciences and Engineering Research Council of Canada (NSERC Discovery Grant No. 371535-2009 to P.D.M.), the Canada Foundation for Innovation (Leaders Opportunity Grant No. 25046 to P.D.M.), and a Royal Society International Exchange grant (J.P. and P.D.M.). D.A. was supported by an NSERC Ph.D. scholarship. D.C. received support from the University of Exeter M.Sc. program in Evolutionary and Behavioural Ecology. J.P. was supported by a Leverhulme Trust Early Career Research Fellowship

Sex differences in condition dependence of natal dispersal in a large herbivore: dispersal propensity and distance are decoupled

International audienceEvolution 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

Flexibility but no coordination of visits in provisioning riflemen

Parental care strategies occupy a continuum from fixed investments that are consistent across contexts to flexible behaviour that largely depends on external social and environmental cues. Determining the flexibility of care behaviour is important, as it influences the outcome of investment games between multiple individuals caring for the same brood. We investigated the repeatability of provisioning behaviour and the potential for turn taking among breeders and helpers in a cooperatively breeding bird, the rifleman, Acanthisitta chloris. First, we examined whether nest visit rate is an accurate measure of investment by assessing whether carers consistently bring the same size of food, and whether food size is related to nest visit rate. Our results support the use of visit rate as a valid indicator of parental investment. Next, we calculated the repeatability of visit rate and food size to determine whether these behaviours are fixed individual traits or flexible responses to particular contexts. We found that riflemen were flexible in visit rate, supporting responsive models of care over ‘sealed bids’. Finally, we used runs tests to assess whether individual riflemen alternated visits with other carers, indicative of turn taking. We found little evidence of any such coordination of parental provisioning. We conclude that individual flexibility in parental care appears to arise through factors such as breeding status and brood demand, rather than as a real-time response to social partners

Wherever I may roam-Human activity alters movements of red deer (Cervus elaphus) and elk (Cervus canadensis) across two continents

Human activity and associated landscape modifications alter the movements of animals with consequences for populations and ecosystems worldwide. Species performing long-distance movements are thought to be particularly sensitive to human impact. Despite the increasing anthropogenic pressure, it remains challenging to understand and predict animals' responses to human activity. Here we address this knowledge gap using 1206 Global Positioning System movement trajectories of 815 individuals from 14 red deer (Cervus elaphus) and 14 elk (Cervus canadensis) populations spanning wide environmental gradients, namely the latitudinal range from the Alps to Scandinavia in Europe, and the Greater Yellowstone Ecosystem in North America. We measured individual-level movements relative to the environmental context, or movement expression, using the standardized metric Intensity of Use, reflecting both the directionality and extent of movements. We expected movement expression to be affected by resource (Normalized Difference Vegetation Index, NDVI) predictability and topography, but those factors to be superseded by human impact. Red deer and elk movement expression varied along a continuum, from highly segmented trajectories over relatively small areas (high intensity of use), to directed transitions through restricted corridors (low intensity of use). Human activity (Human Footprint Index, HFI) was the strongest driver of movement expression, with a steep increase in Intensity of Use as HFI increased, but only until a threshold was reached. After exceeding this level of impact, the Intensity of Use remained unchanged. These results indicate the overall sensitivity of Cervus movement expression to human activity and suggest a limitation of plastic responses under high human pressure, despite the species also occurring in human-dominated landscapes. Our work represents the first comparison of metric-based movement expression across widely distributed populations of a deer genus, contributing to the understanding and prediction of animals' responses to human activit

Wherever I may roam—Human activity alters movements of red deer (Cervus elaphus) and elk (Cervus canadensis) across two continents

Human activity and associated landscape modifications alter the movements of ani-mals with consequences for populations and ecosystems worldwide. Species perform-ing long-distance movements are thought to be particularly sensitive to human impact. Despite the increasing anthropogenic pressure, it remains challenging to understand and predict animals' responses to human activity. Here we address this knowledge gap using 1206 Global Positioning System movement trajectories of 815 individuals from 14 red deer (Cervus elaphus) and 14 elk (Cervus canadensis) populations spanning wide environmental gradients, namely the latitudinal range from the Alps to Scandinavia in Europe, and the Greater Yellowstone Ecosystem in North America. We measured individual-level movements relative to the environmental context, or movement ex-pression, using the standardized metric Intensity of Use, reflecting both the directional-ity and extent of movements. We expected movement expression to be affected by resource (Normalized Difference Vegetation Index, NDVI) predictability and topogra-phy, but those factors to be superseded by human impact. Red deer and elk movement expression varied along a continuum, from highly segmented trajectories over relatively small areas (high intensity of use), to directed transitions through restricted corridors (low intensity of use). Human activity (Human Footprint Index, HFI) was the strong-est driver of movement expression, with a steep increase in Intensity of Use as HFI increased, but only until a threshold was reached. After exceeding this level of impact, the Intensity of Use remained unchanged. These results indicate the overall sensitivity of Cervus movement expression to human activity and suggest a limitation of plastic responses under high human pressure, despite the species also occurring in human-dominated landscapes. Our work represents the first comparison of metric- based movement expression across widely distributed populations of a deer genus, contribut-ing to the understanding and prediction of animals' responses to human activity.publishedVersio

Beyond climate envelope projections : roe deer survival and environmental change.

Research on climate change impacts has focused on projecting changes in the geographic ranges of species, with less emphasis on the vital rates giving rise to species distributions. Managing ungulate populations under future climate change will require an understanding of how their vital rates are affected by direct climatic effects and the indirect climatic and non-climatic effects that are often overlooked by climate impact studies. We used generalized linear models and capture–mark–recapture models to assess the influence of a variety of direct climatic, indirect climatic, and non-climatic predictors on the survival of roe deer (Capreolus capreolus) at 2 sites in Sweden. The models indicated that although direct climatic effects (e.g., precipitation) explained some variation in survival, indirect climatic effects (e.g., an index of vegetation production), and non-climatic effects (hunting by lynx [Lynx lynx] and humans) had greater explanatory power. Climate change is likely to increase vegetation productivity in northern Europe, and, coupled with the positive effects of vegetation productivity on roe deer survival, might lead to population increases in the future. Survival was negatively affected by lynx presence where these predators occur and by human harvest in the site that lacked predators. In the future, managers might find that a combination of increased harvest and predation by recovering carnivore populations may be necessary to mitigate climate-induced increases in roe deer survival. Considering vegetation availability and predation effects is likely to improve predictions of ungulate population responses to variation in climate and, therefore, inform management under future climate change

Effects of rearing conditions on natal dispersal processes in a long-lived predator bird

Natal or prebreeding dispersal is a key driver of the functioning, dynamics, and evolution of populations. Conditions experienced by individuals during development, that is, rearing conditions, may have serious consequences for the multiple components that shape natal dispersal processes. Rearing conditions vary as a result of differences in parental and environmental quality, and it has been shown that favorable rearing conditions are beneficial for individuals throughout their lives. However, the long‐term consequences of rearing conditions on natal dispersal are still not fully understood in long‐lived birds. In this study, we aim to test the following hypotheses to address the relationship between rearing conditions and certain components of the natal dispersal process in Bonelli's eagle (Aquila fasciata): (1) The body condition of nestlings depends on the quality of the territory and/or breeders; and (2) the survival until recruitment, (3) the age of recruitment, and (4) the natal dispersal distance (NDD) all depend on rearing conditions. As expected, nestlings reared in territories with high past productivity of chicks had better body condition, which indicates that both body condition and past productivity reflect the rearing conditions under which chicks are raised. In addition, chicks raised in territories with high past productivity and with good body condition had greater chances of surviving until recruitment. Furthermore, birds that have better condition recruit earlier, and males recruit at a younger age than females. At last, although females in good body condition exhibited higher NDD when they recruited at younger ages, this pattern was not observed in either older females or males. Overall, this study provides evidence that rearing conditions have important long‐term consequences in long‐lived birds. On the basis of our results, we advocate that conservation managers work actively in the promotion of actions aimed at improving the rearing conditions under which individuals develop in threatened populations

Habitat quality, configuration and context effects on roe deer fecundity across a forested landscape mosaic

Effective landscape-scale management of source-sink deer populations will be strengthened by understanding whether local variation in habitat quality drives heterogeneity in productivity. We related female roe deer Capreolus capreolus fecundity and body mass to habitat composition and landscape context, separately for adults and yearlings, using multi-model inference (MMI) applied to a large sample of individuals (yearlings: fecundity=202, body mass=395; adults: fecundity=908, body mass=1669) culled during 2002-2015 from an extensive (195 km2) heterogeneous forest landscape. Adults were heavier (inter-quartile, IQ, effect size=+0.5kg) when culled in buffers comprising more arable lands while contrary to our prediction no effects on body mass of grassland, young forest or access to vegetation on calcareous soil were found. Heavier adults were more fertile (IQ effect size, +12% probability of having two embryos instead of one or zero). Counter-intuitively, adults with greater access to arable lands were less fecund (IQ effect of arable: -7% probability of having two embryos, instead of one or zero), and even accounting for greater body mass of adults with access to arable, their modelled fecundity was similar to or lower than that of adults in the forest interior. In contrast, effects of grassland, young forest and calcareous soil did not receive support. Yearling body mass had an effect on fecundity twice that found in adults (+23% probability of having one additional embryo), but yearling body mass and fecundity were not affected by any candidate habitat or landscape variables. Effect of arable lands on body mass and fecundity were small, with little variance explained (Coefficient of Variation of predicted fecundity across forest sub-regions=0.03 for adults). More variance in fecundity was attributed to other differences between forest management sub-regions (modelled as random effects), suggesting other factors might be important. When analysing source-sink population dynamics to support management, an average value of fecundity can be appropriate across a heterogeneous forest landscape