Nicolas Morellet

Landscape connectivity influences gene flow in a roe deer population inhabiting a fragmented landscape: an individual-based approach

Changes in agricultural practices and forest fragmentation can have a dramatic effect on landscape connectivity and the dispersal of animals, potentially reducing gene flow within populations. In this study, we assessed the influence of woodland connectivity on gene flow in a traditionally forest‐dwelling species — the European roe deer — in a fragmented landscape. From a sample of 648 roe deer spatially referenced within a study area of 55 × 40 km, interindividual genetic distances were calculated from genotypes at 12 polymorphic microsatellite loci. We calculated two geographical distances between each pair of individuals: the Euclidean distance (straight line) and the ‘least cost distance’ (the trajectory that maximizes the use of wooded corridors). We tested the correlation between genetic pairwise distances and the two types of geographical pairwise distance using Mantel tests. The correlation was better using the least cost distance, which takes into account the distribution of wooded patches, especially for females (the correlation was stronger but not significant for males). These results suggest that in a fragmented woodland area roe deer dispersal is strongly linked to wooded structures and hence that gene flow within the roe deer population is influenced by the connectivity of the landscape.

Landscape composition influences roe deer habitat selection at both home range and landscape scales

Understanding how patterns of habitat selection vary in relation to landscape structure is essential to predict ecological responses of species to global change and inform management. We investigated behavioural plasticity in habitat selection of roe deer (Capreolus capreolus) in relation to variable habitat availability across a heterogeneous agricultural landscape at the home range and landscape scales. As expected, woodland was heavily selected, but we found no functional response for this habitat, i.e. no shift in habitat selection with changing habitat availability, possibly due to the presence of hedgerows which were increasingly selected as woodlands were less abundant. Hedgerows may thus function as a substitutable habitat for woodlands by providing roe deer with similar resources. We observed a functional response in the use of hedgerows, implying some degree of landscape complementation between hedgerows and open habitats, which may in part compensate for lower woodland availability. We also expected selection for woodland to be highest at the wider spatial scale, especially when this habitat was limiting. However, our results did not support this hypothesis, but rather indicated a marked influence of habitat composition, as both the availability and distribution of resources conditioned habitat selection. There was no marked between-sex difference in the pattern of habitat selection at either scale or between seasons at the landscape scale, however, within the home range, selection did differ between seasons. We conclude that landscape structure has a marked impact on roe deer habitat selection in agricultural landscapes through processes such as landscape complementation and supplementation.

Testing Global Positioning System Performance for Wildlife Monitoring Using Mobile Collars and Known Reference Points

Abstract 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.

Habitat use under predation risk: hunting, roads and human dwellings influence the spatial behaviour of roe deer

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

Seasonality, weather and climate affect home range size in roe deer across a wide latitudinal gradient within Europe

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.

The Effect of Capture on Ranging Behaviour and Activity of the European Roe Deer Capreolus capreolus

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.

Landscape fragmentation generates spatial variation of diet composition and quality in a generalist herbivore

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.

Condition‐dependent natal dispersal in a large herbivore: heavier animals show a greater propensity to disperse and travel further

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. 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. 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. 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. 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.

Selection for nutrients by red deer hinds feeding on a mixed forest edge

We studied the nutritional behaviour of hinds foraging on a mixed-forest edge by direct observation of their choices at each season and by measuring nutrient concentration in the plants. We compared nutrient concentrations in the observed diets with those in the total available vegetation, and with those of 1,000 randomly simulated diets in which we included only those plants that were actually eaten by the animal. Whether the available or the consumed feeds were used as the basis for comparisons had important consequences due to the presence of conifers and ferns, which were high in soluble sugars but were never eaten by the animals (potentially due to their toxicity). The selected diets were lower in sugars than the total available vegetation in summer, but were actually higher in sugars than the random diets generated from consumed forage species only. Hind diets contained more soluble sugars but not more protein than simulated diets in all seasons. Contrary to our prediction, anti-nutritional compounds (ADL and tannins) were avoided only in winter. Compared to simulated diets, hinds consumed more tannins in spring and summer and more ADL in summer and autumn. We suggest that this was a consequence of selection for soluble sugars, because the preferred plant species, which had high soluble sugar concentrations, also contained a large proportion of the anti-nutritional compounds eaten. In winter, the grass-dominated diets contained more fibre (NDF) and less ADL than the simulated diets, indicating that hinds orient their feeding towards digestible fibres. The switch from a browser to a grazer diet was related to a change in the availability of the nutrients, mainly soluble sugars. In our study, grasses contained more soluble sugars and proteins than deciduous browse during winter. This calls into question the dichotomy usually assumed in the literature between grass and browse in terms of nutrient content.

Spatiotemporal dynamics of forage and water resources shape space use of West African savanna buffaloes

Abstract 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 buffalos 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 km2) 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.