Density-dependent responses of fawn cohort body mass in two contrasting roe deer populations

We investigated the influence of population density on juvenile body mass in two contrasting roe deer populations, in Sweden (Bogesund) and France (Chizé), in which density was monitored for ≥15 years. We investigated the effect of population density and climatic conditions on cohort performance. We predicted that: (1) body mass of growing fawns should be sensitive to environmental changes, showing marked between-year variation (i.e., cohort effects), (2) fawns in the less productive (weakly seasonal, weakly predictable summer weather) habitat of Chizé should show stronger density-dependent responses due to more severe food competition during summer than fawns in the more productive (markedly seasonal, moderately predictable summer weather) habitat of Bogesund, and (3) fawns at Bogesund should be heavier both in absolute terms and relative to their size than their conspecifics in Chizé due to a higher degree of fat accumulation in northern environments. In both study sites we found marked cohort variation and clear effects of density, with body mass varying by as much as 29% over years. While neither summer nor winter climate influenced fawn body mass at Bogesund, fawns tended to be lighter after summers with high temperatures at Chizé. In addition, fawns were heavier after acorn mast years experienced in utero at Bogesund. As expected, the strength of the density-dependent response of fawn body mass was greater at Chizé than at Bogesund. For a given density, male fawns were consistently heavier than females in both sites. Lastly, both sexes at Bogesund had higher absolute body mass and were larger for a given body size than in Chizé. Our results clearly demonstrate that absolute density is a poor predictor of roe deer performance and supports the view that habitat quality has an overwhelming importance for determining fawn body mass in roe deer populations.     

Early survival in roe deer: causes and consequences of cohort variation in two contrasted populations

Time- and sex-specific summer survival of roe deer fawns was estimated using capture-mark-recapture methods in two enclosed populations living in contrasting conditions. The population of Trois Fontaines (eastern France) was roughly constant in size throughout the study period, while in Chizé (western France), the population experienced frequent summer droughts and numbers decreased continuously during the study. Early survival of fawns was low and highly variable over the years at both Chizé and Trois Fontaines, and demonstrated marked variations between cohorts that need to be taken into account when modelling roe deer population dynamics. In Trois Fontaines, fawn survival was positively correlated with early body growth and total rainfall in May and June. In Chizé, fawn survival decreased with increasing density and tended to increase with increasing rainfall in May and June and adult female body mass. These factors explained more than 75% of the variability in early survival observed in both populations. Variation between cohorts had different consequences for the two populations. At Trois Fontaines, cohort variation was limited to a numerical effect on early survival. However at Chizé, cohort variation was long-lasting and affected the phenotypic quality of survivors at later ages, and thereby future survival and breeding abilities (both numerical and quality effects). Male and female fawns had similar survival over their first summer in both populations. This result contrasts with the lower survival of young males often observed in ungulates. Two ultimate causes can be proposed to account for the low and variable survival of roe deer fawns over the first summer: the high energy expenditures incurred by does during each breeding attempt and/or the low absolute body size of newborn roe deer fawns. Received: 28 April 1997 / Accepted: 14 July 1997     

High red deer density depresses body mass of roe deer fawns

Many previous studies have pointed out that, when resources are limited, the potential for competition should be high among sympatric species that display overlaps in habitat and nutritional niches. However, reliable evidence of competition between red deer (Cervus elaphus) and roe deer (Capreolus capreolus) has not been yet reported for life history traits directly measuring performance such as body mass, reproduction, or survival. From long-term monitoring of deer populations in the reserve of La Petite Pierre (France), we measured the sex-specific responses of roe deer fawn body mass to changes in red deer density after accounting for possible confounding effects of date of shooting, climatic conditions, and roe deer density. As expected under the hypothesis of competition, red deer density in a given year had a marked negative influence on body mass of roe deer fawns born the same year and the following year. Fawn mass of roe deer males and females responded in similar ways to changes in red deer density. Our study provides the first evidence of a negative response of roe deer performance to high red deer density.     

Reproductive constraints,not environmental conditions,shape the ontogeny of sex‐specific mass–size allometry in roe deer

In polygynous mammals, sex‐specific patterns of body growth are linked to divergent selection pressures on male and female body size, resulting in sexual dimorphism (SD). For males, reproductive success is generally linked to body size, hence, males should prioritise early growth. For females, reproductive success is linked to resource availability, so they may adopt a more conservative growth tactic. Using longitudinal monitoring of known‐age animals in two contrasting populations and an allometric approach to disentangle the relative contribution of structural size and physiological condition to SD, we addressed these issues in the weakly polygynous roe deer. Despite very different environmental conditions, we found remarkably similar patterns in the two populations in the mass–size allometric relationship at each life history stage, suggesting that relative allocation to structural size and physiological condition is highly constrained. SD in structural size (indexed by hind foot length) involved sex‐specific growth trajectories governed by a single mass–size allometric relationship during the juvenile stage, such that males were both bigger and heavier than females. In contrast, SD in physiological condition (indexed by the allometric relationship between body mass and hind foot length, expressed as body mass for a given body size) developed markedly during the sub‐adult stage in relation to sex differences in the timing of first reproduction. Among adults, males were heavier for a given size than females, suggesting that, relative to females, males express a capital breeder tactic, accumulating fat reserves to offset reproductive costs. By the senescent stage, SD in physiological condition had disappeared, with both sexes governed by a single allometric relationship, suggesting more rapid senescence in males than females. Individuals born into poor cohorts were generally lighter for a given size, indicating growth priority for skeletal size over physiological condition in both sexes. However, sex differences in cohort effects among sub‐adults resulted in lower size‐specific SD in poor cohorts, indicating that body condition of sub‐adult females is buffered against environmental harshness. We conclude that sex‐differences in reproductive tactics impose constraints on the ontogeny of SD in roe deer, leading to sex‐specific trajectories in structural size and physiological condition.     

Spatial variation in springtime food resources influences the winter body mass of roe deer fawns

It is well established that the dynamics of mammalian populations vary in time, in relation to density and weather, and often in interaction with phenotypic differences (sex, age and social status). Habitat quality has recently been identified as another significant source of individual variability in vital rates of deer, including roe deer where spatial variations in fawn body mass were found to be only about a tenth of temporal variations. The approach used was to classify the habitat into blocks a priori, and to analyse variation in animal performance among the predefined areas. In a fine-grained approach, here we use data collected over 24 years on 1,235 roe deer fawns captured at known locations and the plant species composition sampled in 2001 at 578 sites in the Chizé forest to determine the spatial structure at a fine scale of both vegetation and winter body mass of fawns, and then to determine links between the two. Space and time played a nearly equal role in determining fawn body masses of both sexes, each accounting for about 20% of variance and without any interaction between them. The spatial distribution of fawn body mass was perennial over the 24 years considered and predicted values showed a 2 kg range according to location in the reserve, which is much greater than suggested in previous work and is enough to have strong effects on fawn survival. The spatial distribution and the range of predicted body masses were closely similar in males and females. The result of this study is therefore consistent with the view that the life history traits of roe deer are only weakly influenced by sexual selection. The occurrence of three plant species that are known to be important food items in spring/summer roe deer diets, hornbeam (Carpinus betulus), bluebell (Hyacinthoides sp.) and Star of Bethlehem (Ornithogalum sp.) was positively related to winter fawn body mass. The occurrence of species known to be avoided in spring/summer roe deer diets [e.g. butcher's broom (Ruscus aculeatus) and beech (Fagus sylvatica)], was negatively related to fawn body mass. We conclude that the spatial variation in the body mass of fawns in winter in this forest is as important as the temporal variation, and that the distribution of plant species that are actively selected during spring and summer is an important determinant of spatial variation in winter fawn body mass. The availability of these plants is therefore likely to be a key factor in the dynamics of roe deer populations.     

How does environmental variation influence body mass,body size,and body condition? Roe deer as a case study

We tested the influence of population density and of drought intensity (measured as the Gaussen Index in spring and summer of the year of birth) on winter body mass, hind foot length, and body condition of roe deer fawns. Body mass decreased with increasing density and increased with increasing Gaussen Index in summer, in a similar way for both males and females. Hind foot length of males showed the same response. On the other hand, hind foot length of females decreased with increasing density only after dry summers, hence when environmental conditions were very harsh. Body condition was affected neither by density nor by drought intensity. Our results indicate that body mass and size are much better indicators of phenotypic quality than body condition in roe deer. The sex-specific responses of body size to environmental conditions could correspond to a differential allocation in favour of daughters by heavier than average roe deer mothers.     

Long‐lived and heavier females give birth earlier in roe deer

In seasonal environments, parturition of most vertebrates generally occurs within a short time‐window each year. This synchrony is generally interpreted as being adaptive, as early born young survive better over the critical season than late born young. Among large herbivores, the factors involved in driving among‐ and within‐individual variation in parturition date are poorly understood. We explored this question by analyzing the relative importance of attributes linked to female quality (longevity, median adult body mass and cohort), time‐dependent attributes linked to female condition (reproductive success the previous year, relative annual body mass and offspring cohort (year)), and age in shaping observed variation in parturition date of roe deer. A measure of quality combining the effects of female longevity and median adult body mass accounted for 11% of the observed among‐individual variation in parturition date. Females of 2 yr old give birth 5 d later than older females. Our study demonstrates that high quality (heavy and long‐lived) females give birth earlier than low quality females. Temporally variable attributes linked to female condition, such as reproductive success in the previous year and relative annual body mass, had no detectable influence on parturition date. We conclude that parturition date, a crucial determinant of reproductive success, is shaped by attributes linked to female quality rather than by time‐dependent attributes linked to female condition in income breeders (individuals that rely on current resource intake rather than on accumulated body reserves to offset the increased energy requirements due to reproduction) such as roe deer.     

Landscape fragmentation influences winter body mass of roe deer

Body size of large herbivores is a crucial life history variable influencing individual fitness‐related traits. While the importance of this parameter in determining temporal trends in population dynamics is well established, much less information is available on spatial variation in body size at a local infra‐population scale. The relatively recent increase in landscape fragmentation over the last century has lead to substantial spatial heterogeneity in habitat quality across much of the modern agricultural landscape. In this paper, we analyse variation in body mass and size of roe deer inhabiting a heterogeneous agricultural landscape characterised by a variable degree of woodland fragmentation. We predicted that body mass should vary in relation to the degree of access to cultivated meadows and crops providing high quality diet supplements. In support of our prediction, roe deer body mass increased along a gradient of habitat fragmentation, with the heaviest deer occurring in the most open sectors and the lightest in the strict forest environment. These spatial differences were particularly pronounced for juveniles, reaching >3 kg (ca 20% of total body mass) between the two extremes of this gradient, and likely have a marked impact on individual fates. We also found that levels of both nitrogen and phosphorous were higher in deer faecal samples in the more open sectors compared to the forest environment, suggesting that the spatial patterns in body mass could be linked to the availability of high quality feeding habitat provided by the cultivated agricultural plain. Finally, we found that adults in the forest sector were ca 1 kg lighter for a given body size than their counterparts in the more open sectors, suggesting that access to nutrient rich foods allowed deer to accumulate substantial fat reserves, which is unusual for roe deer, with likely knock‐on effects for demographic traits and, hence, population dynamics.     

Assessing the intensity of sexual selection on male body mass and antler length in roe deer Capreolus capreolus: is bigger better in a weakly dimorphic species?

Little is known about traits under sexual selection in territorial mammals with low sexual size dimorphism. We examined the potential for sexual selection on male body mass and antler length in the European roe deer Capreolus capreolus, a territorial ungulate in which males are less than 10% heavier than females. Independently, both body mass and antler length (irrespective of age) had a positive effect on male yearly breeding success. However, when corrected for body mass, antler length at a given mass only had a slight effect on male breeding success. This suggests that: (1) ‘bigger is better’ and (2) sexual selection is responsible for at least part of the observed variation in body mass and antler length in roe deer. High body mass and large antlers may be advantageous to males for two reasons: (1) they enhance fighting ability and dominance, so allowing males to defend better their territory and hence access mates, and (2) they attract females because they are honest signals of male phenotypic quality. This suggests that, even in weakly dimorphic ungulate species, sexual selection may lead to a marked influence of body mass on male breeding success, as long as body mass is also strongly selected in females.     

Bigger teeth for longer life? Longevity and molar height in two roe deer populations

The role of tooth wear as a proximate cause of senescence in ruminants has recently been highlighted. There are two competing hypotheses to explain variation in tooth height and wear; the diet-quality hypothesis predicting increased wear in low-quality habitats, and the life-history hypothesis predicting molar height to be related to expected longevity. We compared tooth height and wear from roe deer of known age from two contrasting populations of roe deer (Capreolus capreolus) in France: Trois Fontaines (TF) with good habitat and shorter animal life expectancy and Chizé (CH) with poor habitat and longer animal life expectancy. There was no population difference in tooth wear, leading to rejection of the diet-quality hypothesis. However, despite their smaller body size, initial molar height for animals from CH was larger than for animals from TF. This provides the first evidence that variation in longevity between populations can lead to differences in molar height within a 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.     

Demographic crash associated with high parasite load in an experimental roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>) population

In March 2002, ten roe deer (Capreolus capreolus) were released in a partly wooded 14.2-ha enclosure to investigate the effect of high population density on vegetation cover and demographic parameters. However, in mid 2003, five animals died after rapid emaciation. Two other deer were killed to carry out further post-mortem examination. In addition to a general loss of body mass and injuries in males caused by the healthiest of them, we found a high level of parasitism. The remaining animals received anti-parasite treatment, and four other treated roe deer were added into the enclosure. All the deer were then captured and treated against parasites twice a year, the fawns being removed when first captured. Four years after the crash, the population seemed healthy: deaths no longer occurred; mean adult body mass reached 27.6 kg in March; females produced on average 1.6 fawns per year, with a mean body mass of 13.8 kg in October and 16.7 kg in March. Furthermore, undergrowth cover changed little since March 2004. These results suggest that outbursts of parasites may cause population crashes in high-density situations before the onset of nutritive stress and may explain certain cases of ‘abnormal’ (massive) mortality recorded in roe deer.     

Effects of population density and female body mass on litter size in European roe deer at a continental scale

Roe deer (Capreolus capreolus) has a wide distributional range in Europe and inhabits a broad range of habitats and environmental conditions. Thus, populations of roe deer show substantial variation in demographic parameters. We aimed to determine whether body mass and population density—which influence the reproductive potential of roe deer at a local scale—affect their reproductive potential at a biogeographical scale as well. We reviewed the literature (covering years 1948–2015) on in utero litter size in roe deer from 59 locations in 14 countries in Europe. Across study sites, mean litter size varied from 1.0 to 2.4 embryo or corpora lutea per female, and population density ranged from 4.5 to 73.5 individuals/km². Mean body mass varied from 11.2 to 20.8 kg in subadult females and from 12.1 to 22.4 kg in adult females. Between 46° and 56° N, body mass of females did not show a significant trend of increase, whereas between 56° and 63° N, it increased with latitude (Bergmann’s rule). We used linear mixed-effects models (LMMs) to analyse the influence of body mass and population density (analysed separately) on litter size. Females with larger body mass produced larger litters, and this pattern was pronounced at both higher and lower latitudes. Higher population densities negatively affected litter size in a sample of all females. This macroecological analysis showed that factors influencing the reproductive potential of roe deer females at local scales produce similar effects at the biogeographical scale.     

Variation in cohort mandible size as an index of roe deer (Capredus capreolus) densities and population trends

This paper examines the effects of increasing population density on the skeletal development of roe deer as indexed by variation in the size of the mandible. We tested for density-dependence in total jaw length and minimum diastema height among cohorts in a single, intensively studied population in France. A strong relationship of average adult jaw length of both males and females with deer density in the year of birth of a given cohort was observed, and presumably reflects inadequate nutrition of juveniles at high population density. No such relationship was observed for the minimum diastema height which remained constant over the range of densities studied. This relationship was exploited to investigate population trends of roe deer in Britain. In nine populations, where equivalent data were available for between 16 and 28 separate cohorts, a consistent pattern of variation in cohort jaw size was observed, with a decrease in average total length, but not diastema height, towards the present. This trend is discussed in relation to changes in habitat structure in the coniferous forests of Britain. The utility of cohort jaw length as a management tool to index population trends of roe deer is highlighted.     

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.     

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     

Abundance estimation of <Emphasis Type="Italic">Ixodes</Emphasis> ticks (Acari: Ixodidae) on roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>)

Despite the importance of roe deer as a host for Ixodes ticks in central Europe, estimates of total tick burden on roe deer are not available to date. We aimed at providing (1) estimates of life stage and sex specific (larvae, nymphs, males and females, hereafter referred to as tick life stages) total Ixodes burden and (2) equations which can be used to predict the total life stage burden by counting the life stage on a selected body area. Within a period of 1½ years, we conducted whole body counts of ticks from 80 hunter-killed roe deer originating from a beech dominated forest area in central Germany. Averaged over the entire study period (winter 2007–summer 2009), the mean tick burden per roe deer was 64.5 (SE ± 10.6). Nymphs were the most numerous tick life stage per roe deer (23.9 ± 3.2), followed by females (21.4 ± 3.5), larvae (10.8 ± 4.2) and males (8.4 ± 1.5). The individual tick burden was highly aggregated (k = 0.46); levels of aggregation were highest in larvae (k = 0.08), followed by males (k = 0.40), females (k = 0.49) and nymphs (k = 0.71). To predict total life stage specific burdens based on counts on selected body parts, we provide linear equations. For estimating larvae abundance on the entire roe deer, counts can be restricted to the front legs. Tick counts restricted to the head are sufficient to estimate total nymph burden and counts on the neck are appropriate for estimating adult ticks (females and males). In order to estimate the combined tick burden, tick counts on the head can be used for extrapolation. The presented linear models are highly significant and explain 84.1, 77.3, 90.5, 91.3, and 65.3% (adjusted R ²) of the observed variance, respectively. Thus, these models offer a robust basis for rapid tick abundance assessment. This can be useful for studies aiming at estimating effects of abiotic and biotic factors on tick abundance, modelling tick population dynamics, modelling tick-borne pathogen transmission dynamics or assessing the efficacy of acaricides.     

The modular organization of roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>) body during ontogeny: the effects of sex and habitat

Background

As a small artiodactyl, the roe deer (Capreolus capreolus L.) is characterized by biological plasticity and great adaptability demonstrated by their survival under a wide variety of environmental conditions. In order to depict patterns of phenotypic variation of roe deer body this study aims to quantify variation during ontogenetic development and determine how sex-specific reproductive investment and non-uniform habitat differences relate to phenotypic variation and do these differential investments mold the patterns of phenotypic variation through modular organisation.

Results

Patterns of phenotypic correlation among body traits change during the ontogeny of roe deer, with differential influence of sex and habitat type. Modularity was found to be a feature of closed habitats with trunk+forelimbs+hindlimbs as the best supported integration/modularity hypothesis for both sexes. The indices of integration and evolvability vary with habitat type, age and sex where increased integration is followed by decreased evolvability.

Conclusion

This is the first study that quantifies patterns of correlation in the roe deer body and finds pronounced changes in correlation structure during ontogeny affected by sex and habitat type. The correlation structure of the roe deer body is developmentally written over the course of ontogeny but we do not exclude the influence of function on ontogenetic changes. Modularity arises with the onset of reproduction (subadults not being modular) and is differentially expressed in males and females from different habitats. Both adult males and females show modularity in primordial, closed habitats. Overall, all these findings are important as they provide support to the idea that modularity can evolve at the population level and change fast within a species.

     

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.     

Access to mates in a territorial ungulate is determined by the size of a male's territory, but not by its habitat quality

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.