Use of space and habitat selection by roe deer Capreolus capreolus in a Mediterranean coastal area: how does woods landscape affect home range?

The home-range dynamics and habitat selection of nine roe deer were studied from March 1994 to August 1994 in the Maremma Natural Park along the Tyrrhenian coast of Italy. The habitat was highly fragmented, with open agricultural fields prevailing in the study area (57%); the climate was Mediterranean. Data on spatial behaviour were collected by radio-tracking techniques. Habitat selection and structure were investigated by compositional and landscape analysis, both within the study area and within the home ranges. Animals of our sample showed spatial-use patterns varying from stationary to roaming. Stationary individuals used small home ranges while roaming ones moved, especially during the reproductive period in July and August. The percentage and structure of woodlands influenced the size of home ranges and the behaviour of males: stationary males used large amounts of woodlands within their home ranges and showed a territorial behaviour whereas males that used a high percentage of fields showed wider home ranges even during the territorial period. Females seemed to be less influenced by the presence and patch-structure of woodland within their home range. Landscape structure and habitat composition seemed to be important factors influencing the spatial behaviour of this roe deer population.     

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.     

Life at the edge: Roe deer occurrence at the opposite ends of their geographical distribution,Norway and Portugal

In the face of climate change and habitat fragmentation there is an increasingly urgent need to learn more about factors that influence species distribution patterns and levels of environmental tolerance. Particular insights can be obtained by looking at the edges of a species range, especially from species with wide distributions. The European roe deer was chosen as a model species due to its widespread distribution. By using pellet group counts, we studied summer and winter habitat use of this herbivore at two of the extreme edges of its distribution – southwest of Portugal, and northeast of Norway – in relation to a range of fine-scale environmental factors including forest structure, vegetation characteristics and human disturbance. Our first prediction that roe deer would respond differently to human activity in both counties was supported. While in Norway roe deer are always close to houses, in Portugal they are either far (in summer) or indifferent (winter). However, everywhere and in every season, roe deer are far from roads. Our second prediction that roe deer better tolerate anthropogenic disturbances in the area where the importance of limiting factors is higher (Norway) was validated. However, our third prediction that anthropogenic disturbance would be less tolerated by roe deer outside the limiting seasons in each country was not supported. Our results suggest that roe deer perceive human activities differently in the two countries and that roe deer better tolerate anthropogenic disturbances in Norway.     

Do male roe deer clump together during the rut?

Movement s an d habitat use of 7 male and 7 female roe deerCapreolus capreolus (Linnaeus, 1758) were studied by radioteleme try from March 1999 to February 2001. Annual and bimonthly home ranges of males were small (ca 10 ha, 95% kernel), with large overlap among individuals throughout the year. Exclusive core areas (ca 0.4 ha, 25% kernel) were concentrated in the forest, a limited and sought-after resource in the study area. The difference in overlap between male exclusive core areas and female home ranges in the pre-rut and rut periods suggested that females made excursions to search for territorial males during the rut Our results support the mating strategy hypothesis of territorial behaviour Different space use patterns occurred between the sexes, with females apparently playing an active role in mate choice by visiting males at clumps of core areas in the forest.     

There is only one winner: The negative impact of red deer density on roe deer numbers and distribution in the Słowiński National Park and its vicinity

Red and roe deer are the most numerous cervids in Europe, and they occur in sympatry in most regions. Roe deer were considered to be an inferior competitor in studies in which they co‐occurred with fallow deer or muntjac. Despite the remarkable overlap of their ranges, there are few studies on the competition between the red and roe deer. Since interspecific interactions among ungulates are often related to their mutual densities, the current study focused on the effects of high red deer density on the roe deer numbers and spatial distribution in the unhunted Słowiński National Park (SNP) in northern Poland and forest districts open to hunting bordering the park. Using fecal pellet group counts, it was found that in the forest districts (where red deer densities were 2–3 times lower than in the SNP), roe deer densities were significantly higher than in the park. The red‐to‐roe deer density ratio was 10.8 and 2.7, in the SNP and the surrounding forest districts, respectively. Moreover, in the SNP, the roe deer distribution was negatively affected by the red deer habitat use, while in the hunting areas, such an effect was not recorded. The negative influence of the red deer on the roe deer population in the park was most probably due to the red deer impact on food availability. The biomass of the plant groups forming the staple food of the roe deer (Rubus spp., forbs, dwarf shrubs) was significantly higher in the fenced plots than in the unfenced ones. Lack of hunting in the protected areas may benefit only some species in ungulate assemblages which, in turn, may contradict one of their objectives—to maintain viable and ecologically functional populations.     

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 x 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-scale habitat characteristics and neonatal white-tailed deer survival

Landscape-level habitat characteristics affect neonatal white-tailed deer (Odocoileus virginianus) survival. Little is known, however, about how changes in maternal habitat use after parturition affect neonate survival. We quantified survival rates and determined if neonate survival to 8 weeks was affected by weekly maternal habitat use in the agricultural Glaciated Plains (GP) and forest-grassland Ozark (OZ) eco-regions of Missouri, USA. We captured 127 pregnant female deer during 2015–2017, and fitted each with a global positioning system (GPS) radio-collar and vaginal implant transmitter (VIT). We captured 226 neonatal deer during 2015–2017, fitted each with an expandable radio-collar, and monitored survival status daily. We estimated weekly maternal home ranges and calculated habitat metrics within these home ranges. We used the Kaplan-Meier estimator to calculate 8-week survival estimates and Cox proportional hazards models to investigate the influence of habitat metrics on neonate survival. The 8-week survival estimates were 0.43 (95% CI = 0.35–0.54) and 0.47 (95% CI = 0.38–0.57) in the GP and OZ, respectively. Both of these survival estimates were lower than expected but particularly so in the GP because it is dominated by agricultural fields, a land cover type typically associated with high survival. Neonate survival in the GP was negatively correlated with the amount of edge and forest patch size within maternal home ranges. In the OZ, female neonate survival was positively correlated with birth mass, male neonate survival was not affected by birth mass, and survival of both sexes was negatively correlated with grassland patch density. We suspect these habitat metrics were related to predator searching efficiency and abundance. In the highly fragmented GP, predators might be able to easily search the largest cover habitat patches, whereas in the more contiguous OZ landscape, where cover habitat patch sizes were > 10 times the size of patches in the GP, large patches might be difficult for predators to search efficiently. Therefore, we recommend managers consider the larger landscape context when making habitat management decisions to increase white-tailed deer population productivity. © 2019 The Wildlife Society.     

Home range dynamics of male roe deerCapreolus capreolus in a mountainous habitat

Male spacing behaviour of roe deer Capreolus capreolus (Linnaeus, 1758) was studied in a wooded mountainous habitat in the Casentinesi Forest National Park, Italy. Data were collected using radio-tracking techniques from March 1997 to February 1998. Annual, seasonal, and bimonthly home ranges were analysed. Different factors may influence male spacing behaviour throughout the year. Winter home range sizes may be dependent on environmental conditions, while social factors could determine a high level of individual variability during the territorial and reproductive period. Prime age males showed great spatial stability, in contrast to the ranging movements of yearlings.     

What shapes intra‐specific variation in home range size? A case study of female roe deer

Spatial distribution in mammals, and thereby home range size, is influenced by many different factors including body size, sex, age, reproductive status, season, availability of forage, availability of water, fragmentation of landscape, trophic level and intra- and inter-specific competition. Using linear mixed models, we looked for factors shaping the variation in size of spring-summer and winter home ranges for 51 radio-collared adult female roe deer at Trois Fontaines forest, Champagne–Ardenne, France (1996–2005). Home range size of females was larger in winter than in spring–summer, decreased with age, and decreased with increasing quality. Females in low quality areas adjusted the size of their home range to include more patches of habitat so that all female deer obtained similar amounts of food resources (total biomass of 6.73±2.34 tons (mean±SE) for each home range). Such adjustments of home range size in response to patchiness of resources led to marked between-female variation in home range size. Our results demonstrate that roe deer females have different tactics of habitat use according to spatial variations in habitat quality so that females get similar food resources in highly productive environments such as the Trois Fontaines forest.     

Multi-Scale Foraging Habitat Use and Interactions by Sympatric Cervids in Northeastern China

Abstract: Moose (Alces alces) and roe deer (Capreolus pygargus) are sympatric in the forest region of northeastern China. Using univariate analyses of feeding sign data, we found the 2 species were positively associated, but there were distinctions between their use of forage resources across landscape, patch, and microhabitat scales. We used resource selection function models to predict the influence of environmental covariates on moose and roe deer foraging; we detected covariate effects at the landscape and microhabitat scales but not at the patch scale. Forage resources used by the 2 species were similar, but moose used wetter areas and more low-visibility habitats than did roe deer, which strongly avoided areas with sparse vegetation. Both species were influenced by forage abundance and distribution at the microhabitat scale but exhibited differences in intensity of use of plant species and microhabitats. Moose used areas with deeper snow and avoided hiding cover; roe deer avoided areas with higher total basal areas of tree stems and preferred areas with high plant species richness. For moose, there was a trade-off in the use of concealment cover between the landscape and microhabitat scales. We detected avoidance by moose of roads where roe deer occurred. Roe deer exhibited more capacity for coping with human disturbance and interspecific interaction. In areas similar to our study area, road closures and suppression of roe deer near roads within 3–5 years postlogging may benefit moose. Furthermore, a mosaic of areas with different logging intervals may contribute to spatial separation of moose and roe deer and promote their coexistence.     

Predation processes: behavioural interactions between red fox and roe deer during the fawning season

Predation by red fox (Vulpes vulpes) is the most important mortality cause for neonatal roe deer (Capreolus capreolus) in Scandinavia. With the objective of investigating how the fox finds fawns and how antipredatory behaviour of roe deer females influences choice of hunting method, I analysed observations of interactions between red fox and roe deer females. The observations were collected over 14 years in a mixed forest/agricultural landscape in Sweden. Of 49 fox–doe encounters, the doe attacked the fox in 59%. In 90% of these attacks the fox was successfully deterred. In two observations a doe saved a fawn attacked by a fox. Two hunting methods used by the fox were discerned. In 28 cases foxes searched the ground, and in 18 cases they surveyed open areas, often from a forest edge. The latter behaviour seemed more directed at fawns and was seen leading to a capture attempt. Searching seemed less efficient and also difficult to conduct due to the aggressiveness of does. A surveying sit-and-wait type of hunting method thus appeared as the most successful. The possibility to use this method could explain why roe deer fawns are more vulnerable to fox predation in open habitats.     

Occurrence patterns of crop‐foraging sika deer distribution in an agriculture–forest landscape revealed by nitrogen stable isotopes

Conflicts arising from the consumption of anthropogenic foods by wildlife are increasing worldwide. Conventional tools for evaluating the spatial distribution pattern of large terrestrial mammals that consume anthropogenic foods have various limitations, despite their importance in management to mitigate conflicts. In this study, we examined the spatial distribution pattern of crop‐foraging sika deer by performing nitrogen stable isotope analyses of bone collagen. We evaluated whether crop‐foraging deer lived closer to agricultural crop fields during the winter and spring, when crop production decreases. We found that female deer in proximity to agricultural crop fields during the winter and spring were more likely to be crop‐foraging individuals. Furthermore, the likelihood of crop consumption by females decreased by half as the distance to agricultural crop fields increased to 5–10 km. We did not detect a significant trend in the spatial distribution of crop‐foraging male deer. The findings of spatial distribution patterns of crop‐foraging female deer will be useful for the establishment of management areas, such as zonation, for efficient removal of them.     

Spatial patterns and density dependence in the dynamics of a roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>) population in central Italy

The demography of roe deer living in a mountain area of central Italy was studied from 1995 to 1999 with 104 radio-tagged animals, including fawns and adults of both sexes. From spring surveys we estimated population density by mark-resighting (average: 53.8dž.8 individuals km^-2) and found an average fawn/doe ratio of 0.75ǂ.4. The fawn/doe ratio was negatively correlated to density suggesting density-dependent regulation in this population. Using culled and net-trapped individuals we evaluated the dressed body weights of adult males (23.1ǃ.0 kg) and females (22.0ǃ.0 kg), which indicated a low level of sexual dimorphism. The potential litter size (1.44ǂ.1 embryos) depended on female body weight and a threshold of 20.9ǃ.4 kg separated adult females carrying one or two embryos. Both fawn (0.38ǂ.07) and adult survival (0.90ǂ.07) were evaluated from radiotagged individuals and no gender effect was observed in either age class. During the study period we recorded a population decline in one part of the study area and an increasing fawn mortality, which was attributed to the spreading of an enteropathogenic desease. The study revealed an unexpected spatial structure in population dynamics at a scale of few square kilometres. In the two studied subareas, which are very close and ecologically similar, we documented significant differences in several demographic parameters: females in the subarea with the highest deer density produced smaller litters and allocated their reproductive effort preferentially to males, which is consistent with the hypothesis that local resource competition determines sex allocation in roe deer. The importance of spatial variability with respect to roe deer demography was overlooked in previous studies and our results raise new interesting research questions relative to the study of population equilibria which are also relevant for the management of this important game species.     

Tick burden on European roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>)

In our study we assessed the tick burden on roe deer (Capreolus capreolus L.) in relation to age, physical condition, sex, deer density and season. The main objective was to find predictive parameters for tick burden. In September 2007, May, July, and September 2008, and in May and July 2009 we collected ticks on 142 culled roe deer from nine forest departments in Southern Hesse, Germany. To correlate tick burden and deer density we estimated deer density using line transect sampling that accounts for different detectability in March 2008 and 2009, respectively. We collected more than 8,600 ticks from roe deer heads and necks, 92.6% of which were Ixodes spp., 7.4% Dermacentor spp. Among Ixodes, 3.3% were larvae, 50.5% nymphs, 34.8% females and 11.4% males, with significant seasonal deviation. Total tick infestation was high, with considerable individual variation (from 0 to 270 ticks/deer). Adult tick burden was positively correlated with roe deer body indices (body mass, age, hind foot length). Significantly more nymphs were found on deer from forest departments with high roe deer density indices, indicating a positive correlation with deer abundance. Overall, tick burden was highly variable. Seasonality and large scale spatial characteristics appeared to be the most important factors affecting tick burden on roe deer.     

Maternal and individual effects in selection of bed sites and their consequences for fawn survival at different spatial scales

We examined the relationship between survival of roe deer (Capreolus capreolus) fawns at Trois Fontaines, Champagne-Ardennes, France, and factors related to bed-site selection (predator avoidance and thermoregulation) and maternal food resources (forage availability in the maternal home range). Previous studies have demonstrated that at small scales, the young of large herbivores select bed sites independently from their mothers, although this selection takes place within the limits of their mother’s home range. Fawn survival was influenced largely by the availability of good bed sites within the maternal home range, not by the fawn’s selection of bed sites; however, selection for thermal cover when selecting bed sites positively influenced survival of young fawns. Typical features of a good home range included close proximity to habitat edges, which is related to forage accessibility for roe deer. The availability of bed sites changed as fawns aged, probably due to an increased mobility of the fawn or a different use of the home range by the mother; sites offering high concealment and thermal protection became less available in favor of areas with higher forage accessibility. Despite the minor influence of bed-site selection on survival, roe deer fawns strongly selected their bed sites according to several environmental factors linked to predator avoidance and thermoregulation. Fawns selected for sites providing concealment, light penetration, and avoided signs of wild boar (Sus scrofa) activity. Avoidance of sites with high light penetration by young fawns positively affected their survival, confirming a negative effect on thermoregulation due to reduced thermal cover. Selection for light penetration by older fawns was less clear. We discuss these results in the context of cross-generational effects in habitat selection across multiple scales, and the potential influence of the ‘ghost of predation past’.     

Distribution of an increasing roe deer population in a fragmented Mediterranean landscape

This paper describes roe deer Capreolus capreolus distribution in central Spain, where the species has spread from the mountains into peripheral agricultural areas In this region, it is more abundant in forest fragments near the mountains than in those further away, in pine and oak woodlands than in sclerophyllous forests, and in forests with open water than in those lacking this resource The paper also analyzes whether habitat quality and geographic location of forests with respect to roe deer source areas are the two basic causes of its distribution, as predicted by some models of the species distribution in fragmented landscapes The results corroborate this hypothesis revealing that roe deer abundance is linked negatively to sclerophyllous forest cover and distance from mountains We conclude that this type of farmland seems to be sub-optimal for roe deer in comparison with forested, moist mountains     

Ranging behaviour of roe deer in an experimental high-density population: Are females territorial?

We studied the ranging behaviour and spatial relationships between seven roe deer during more than 4 years in a partly wooded 14.2-ha enclosure. The animals (three young males, four adult females) were monitored with GPS telemetry collars. As expected, the surface area and overlap of the males’ bimonthly ranges decreased, and the distance between their arithmetic centres increased, as they became adult and, for two of them, territorial. Unexpectedly, females also tended to space out, the surface area and overlap of their bimonthly ranges being minimal in May to June, i.e. during the birth period. The distance between their arithmetic centres reached its maximum at the same time. Overlap between females’ ranges was consistently lower than those between males and females’ ranges, or between 1-year old males’ ranges. Our results raise the questions of female seasonal territoriality and of independence of the spacing systems of the two sexes in roe deer.     

Ranging Movements of Female Roe Deer: Do Home-loving Does Roam to Mate?

Bimonthly changes in home range size of seven mature roe deer Capreolus capreolus L. females were analysed over one year. Great stability of home range size and high site fidelity was common to all females, which remained bimonthly within areas of 6.1–20.8 ha. Two groups of females could be identified on the basis of bimonthly variations in their home range sizes: females with larger areas (LHR females), who decreased their ranges in the birth season only, and females with smaller home ranges (SHR females), who increased significantly their home ranges during the rut. The most likely explanation is that the areas inhabited by SHR females included high-quality food resources allowing survival even in small home ranges. In the rut, SHR females may have expanded their ranges to increase mating opportunities.     

Siberian Roe Deer (<Emphasis Type="Italic">Capreolus pygargus</Emphasis> Pallas, 1771) in Ukraine: Analysis of the Mitochondrial and Nuclear DNA

A molecular-genetic analysis of the nucleotide sequences of the cytochrome b gene (1140 base pairs) of the mitochondrial DNA and 17 microsatellite loci of eight samples of roe deer from the Samara forest of Dnipropetrovsk oblast (Ukraine) was carried out. For comparison, 212 corresponding mtDNA sequences of the Siberian and European roe deer and data on the variability of microsatellite markers in 49 representatives of these species were included in the study. It was noted that all the analyzed mitochondrial sequences of individuals from the Samara forest are characteristic of the Siberian roe Capreolus pygargus Pallas, 1771. Four haplotypes were described, all of which belonged to the haplogroup typical for the western part of the range of C. pygargus. A fragment analysis of the microsatellite loci of nuclear DNA confirmed the identification of the investigated group with the Siberian species.