Patterns of Lynx Predation at the Interface between Protected Areas and Multi-Use Landscapes in Central Europe

In Central Europe, protected areas are too small to ensure survival of populations of large carnivores. In the surrounding areas, these species are often persecuted due to competition with game hunters. Therefore, understanding how predation intensity varies spatio-temporally across areas with different levels of protection is fundamental. We investigated the predation patterns of Eurasian lynx (Lynx lynx) on roe deer (Capreolus capreolus) and red deer (Cervus elaphus) in both protected areas and multi-use landscapes of the Bohemian Forest Ecosystem. Based on 359 roe and red deer killed by 10 GPS-collared lynx, we calculated the species-specific annual kill rates and tested for effects of season and lynx age, sex and reproductive status. Because roe and red deer in the study area concentrate in unprotected lowlands during winter, we modeled spatial distribution of kills separately for summer and winter and calculated-the probability of a deer killed by lynx and-the expected number of kills for areas with different levels of protection. Significantly more roe deer (46.05–74.71/year/individual lynx) were killed than red deer (1.57–9.63/year/individual lynx), more deer were killed in winter than in summer, and lynx family groups had higher annual kill rates than adult male, single adult female and subadult female lynx. In winter the probability of a deer killed and the expected number of kills were higher outside the most protected part of the study area than inside; in summer, this probability did not differ between areas, and the expected number of kills was slightly larger inside than outside the most protected part of the study area. This indicates that the intensity of lynx predation in the unprotected part of the Bohemian Forest Ecosystem increases in winter, thus mitigation of conflicts in these areas should be included as a priority in the lynx conservation strategy.     

Country,Cover or Protection: What Shapes the Distribution of Red Deer and Roe Deer in the Bohemian Forest Ecosystem?

The Bohemian Forest Ecosystem encompasses various wildlife management systems. Two large, contiguous national parks (one in Germany and one in the Czech Republic) form the centre of the area, are surrounded by private hunting grounds, and hunting regulations in each country differ. Here we aimed at unravelling the influence of management-related and environmental factors on the distribution of red deer (Cervus elaphus) and roe deer (Capreolus capreolus) in this ecosystem. We used the standing crop method based on counts of pellet groups, with point counts every 100 m along 218 randomly distributed transects. Our analysis, which accounted for overdispersion as well as zero inflation and spatial autocorrelation, corroborated the view that both human management and the physical and biological environment drive ungulate distribution in mountainous areas in Central Europe. In contrast to our expectations, protection by national parks was the least important variable for red deer and the third important out of four variables for roe deer; protection negatively influenced roe deer distribution in both parks and positively influenced red deer distribution in Germany. Country was the most influential variable for both red and roe deer, with higher counts of pellet groups in the Czech Republic than in Germany. Elevation, which indicates increasing environmental harshness, was the second most important variable for both species. Forest cover was the least important variable for roe deer and the third important variable for red deer; the relationship for roe deer was positive and linear, and optimal forest cover for red deer was about 70% within a 500 m radius. Our results have direct implications for the future conservation management of deer in protected areas in Central Europe and show in particular that large non-intervention zones may not cause agglomerations of deer that could lead to conflicts along the border of protected, mountainous areas.     

Wolf (Canis lupus) feeding habits during the first eight years of its occurrence in Germany

Due to the fact that the feeding habits of large carnivores are the main contentious point when they start resettling regions they were absent from for several decades, the diet composition of the wolves in Germany was analysed from the beginning of this process. Wolves in Germany primarily feed on wild ungulates, which make up more than 96% of their diet. The dominating prey species is the roe deer (55.3%), followed by red deer (20.8%) and wild boar (17.7%). The second important food category are the leporids (2.9% of Biomass), whereas livestock makes up only 0.6% of all biomass consumed. Wolves clearly prefer hunting on juvenile to adult red deer; roe deer are not selected after their age. We found seasonal differences in the diet composition with a higher amount of wild boar in spring and winter, when a high amount of juveniles and weakened animals, respectively, are available. In the first years of the study the percentage of red deer was much higher, and the percentage of roe deer therefore was lower than the following years. The amount of wild boar in the wolf diet fluctuated most in the first three years. Diet composition remained constant during the last five years. Wolves needed less than two generations for adapting to the new conditions in the cultivated landscape of eastern Germany.     

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.     

Behavioural and spatial adaptation of the Eurasian lynx to a decline in prey availability

The distribution and abundance of food resources is a major factor influencing animal populations. I studied the effect of a roe and red deer population decline on diet composition, home range size and foraging pattern in the Eurasian lynxLynx lynx (Linnaeus, 1758) in the Bia?owie?a Primeval Forest (BPF), eastern Poland. The population of cervids in BPF experienced a nearly two-fold reduction in size from 1991 through 2006 due to severe hunting pressure between 1991 and 1996. Comparison of published data on lynx diet during the high abundance of ungulates with new data obtained for the low abundance period showed that despite a significant decline in their availability, cervids (roe and red deer) continued to form the majority of the diet of lynx, with roe deer being most preferred in both periods. Home range sizes of lynx showed a tendency to increase with declining prey densities, as indicated by relative percentage increases in average yearly home range sizes amongst different sex/age groups. In response to lower availability of their main prey, lynx increased their daily straight-line movement distances by 44% and doubled the ranges covered in 5-day periods. This illustrated that, with declining prey abundance, the lynx increased their hunting efforts by either spending more time actively searching for prey or continuing foraging even after a successful hunt. Spatial analysis of the distribution of ungulates and lynx indicated that deer were evenly distributed throughout lynx ranges in BPF and spatial proximity of the predator to prey sites did not play an important role in the efficiency of hunting. Lynx may adapt to changing prey availability by increasing search effort, but this was not sufficient to prevent the negative influences of the prey decline on the lynx population. Prey depletion has an immediate effect on lynx spatial organization and, in consequence, on their density. This information has to be considered in prioritizing lynx conservation measures and management of ungulates.     

Foraging of lynxes in a managed boreal-alpine environment

Foraging of Eurasian lynxes Lynx lynx was studied with telemetry and snow tracking in central Norway. In all habitats and at all seasons, medium-sized ungulates (roe deer Capreolus capreolus , reindeer Rangifer tarandus and domestic sheep Ovis aries) dominated the diet (81% of ingested biomass estimated from faeces). Mountain hares Lepus timidus and galliform birds comprised the remainder of the diet (15% and 3%, respectively). Lynxes with different life history status did not differ in prey choice, but adult males utilised carcasses of ungulate prey considerably less (16% of the edible parts) than did females with offspring (80%) and subadults (58%.). Forest habitats in lowlands and adjacent to cultivated fields were the most favourable foraging habitats (indexed as the prey encounter rate per km lynx track) primarily owing to the presence of roe deer. Two family groups tracked in winter killed 0.2 ungulate per day. The importance of agricultural land as a foraging habitat and the dominance of livestock in the diet in remoter areas indicate that the lynx has responded to agriculture and reindeer husbandry during the past century by switching from smaff game to ungulates.     

The consequences for deer of ingesting oilseed rape (Brassica napus): feeding experiments with roe deer (Capreolus capreolus) and red deer (Cervus elaphus)

In two experiments, in the growing season March–May, freshly harvested double-low oilseed rape ( Brassica napus ) plants were fed ad libitum to penned roe deer ( Capreolus capreolus ) and red deer ( Cervus elaphus ). In Experiment 1, oilseed rape was fed to five roe and 12 red deer as 60% of their daily dry matter (DM) intake for four weeks, after a two-week dietary change-over period. The rest of the diet comprised cattle-rearing pellets (33%) and freshly cut heather ( Calluna vulgaris ) plants (7%). In Experiment 2, oilseed rape was fed to seven roe and eight red deer as 100% of the diet for up to six weeks, after a two-week change-over period.
In both experiments, Heinz bodies were found in roe deer blood 2–3 weeks after the start of the change-over period, with a fall in mean packed cell volume (PCV) and blood glutathione (GSH) concentration. In Experiment 2, after 5–8 days of feeding on 100% oilseed rape, four roe deer showed signs of inappetance and were changed to a diet of 80% oilseed rape. At the same time, the other three roe deer had very low PCV (< 60% of normal values) and oilseed rape feeding was discontinued for those animals. The red deer showed no symptoms of haemolytic damage nor any change in blood GSH concentration in either experiment, but showed a gradual fall in PCV in Experiment 2. No other ill-effects were observed in any of the animals.
Intakes of oilseed rape (gDM/kg^0–75/day) were lower for the roe deer than the red deer in both experiments. The severity of haemolytic anaemia in the roe deer was related to the proportion of oilseed rape in the diet, rather than the amount ingested. It was concluded that the health of roe deer ingesting oilseed rape may be affected if other foods are not available.     

完达山林区森林采伐和非采伐区马鹿、狍子对冬季生境因子选择的比较

2003—2005年的冬季,在黑龙江省完达山地区的五泡林场,通过样线调查,利用逐步判别分析的方法,对冬季马鹿和狍子在森林采伐区和非采伐区生境选择的差异进行了比较研究,其结果表明,在该地区虽然两物种在生境选择上发生部分重叠,但在生境利用方式上均存在显著差异在采伐区马鹿主要利用中坡位或上坡位、半阴半阳或阳坡、人为干扰距离大于1000m、海拔相对较高、食物丰富度较高、乔木胸径较小、灌木密度较高的灌丛或杂木林生境;狍子主要利用中坡位或下坡位、阳坡、海拔相对较低、食物丰富度适中、乔木胸径较大、灌木密度较低、人为干扰距离要求不严格的杂木林或柞树林生境。在非采伐区,马鹿主要选择距人为干扰大于1000m、海拔较高、食物丰富度较高、乔木胸径较小的杂木林生境;狍子主要选择中坡位或下坡位、人为干扰距离要求不严格、海拔较低、食物丰富度较适中和乔木胸径较大的杂木林或杨-桦混交林生境。     

Effects of red deer on tree regeneration and growth in Aorangi Forest, Wairarapa

New Zealand forests have been substantially modified by introduced red deer over the past century. New Zealand’s indigenous forest managers need to know if regeneration of palatable tree species can be restored following control or eradication of browsing ungulates. Aorangi Forest, Wairarapa, suffered dramatic changes in forest understorey composition by the 1950s after more than seven decades of colonisation by red deer (Cervus elaphus), feral goats (Capra hircus) and pigs (Sus scrofa). Since then feral goats have been eradicated, but red deer and pigs persist throughout Aorangi Forest despite ongoing recreational hunting. This study uses data from paired fenced and unfenced plots established at seven sites in Aorangi Forest between 1981 and 1987, and re-measured in 2004, to show the effects of ungulates on tree (≥ 2 cm diameter at breast height) regeneration and growth. Our results show that browsing by red deer has prevented regeneration of kanono (Coprosma grandifolia), a highly palatable, fast-growing sub-canopy hardwood tree. Deer reduced the growth of mahoe (Melicytus ramiflorus) trees, probably by directly browsing epicormic shoots. The regeneration of other less palatable sub-canopy trees (e.g. porokaiwhiri, Hedycarya arborea), and slower-growing canopy species (e.g. hinau, Elaeocarpus dentatus and rewarewa, Knightia excelsa) appears to have been unaffected by deer browsing. Accordingly, tree species composition does not appear to have been greatly affected by browsing from deer populations in Aorangi Forest over the past two decades.     

In vitro digestibility of seasonal forages used by wapiti and roe deer in northeastern China

Seasonal changes in in vitro dry matter digestibility (IVDMD) of five forage classes (browses, sedges, forbs. ferns and the bark of Ulmus spp.) used by wapiti and roe deer were determined in relation to chemical composition, using the two-stage technique of in vitro trial with rumen inocula from one wapiti and one roe deer captured in winter. Wapiti and roe deer showed remarkable fluctuation in IVDMD among seasons and forage classes. IVDMD decreased from 43.8-64.3% in spring to 29.4-40.5% in winter for wapiti, and from 52.4-77.4% in spring to 25.9-36.5% in winter for roe deer. There was significant variation in IVDMD among species of browses for wapiti and roe deer. IVDMD ranged from 28.5% to 58.7% in twigs and from 39.8% to 58.2% in leaves for wapiti, and from 24.1% to 67.7% in twigs and from 40.0% to 71.7% in leaves for roe deer. Roe deer had higher IVDMD (40.0-71.7%) in both leaves and twigs of browses than wapiti (35.9-58.7%) during snow-free seasons, but IVDMD of twigs was lower for roc deer (25.9-36.5%) than wapiti (29.4-38.4%) in winter. Significant multiple correlation was found between NDF and ADF, and IVDMD in sedges, forbs and ferns for both wapiti and roe deer, whereas only roe deer showed the significant multiple correlation for leaves of browses.     

Relationship between prey availability and population dynamics of the Eurasian lynx and its diet in northern Belarus

The diet and population dynamics of the Eurasian lynxLynx lynx Linnaeus, 1758 as well as an index of its main prey abundance were studied in transitional mixed forests of northern Belarus in 1985-2004. Monitoring of the lynx population and its main potential prey (the mountain hareLepus timidus, and the roe deerCapreolus capreolus) was done by snow-tracking. Also, abundance of tetraonids (Tetraonidae) was monitored by sight count. Hare numbers were fairly stable during the study period, whereas density of the roe deer population markedly increased, and tetraonids decreased. Composition of the lynx diet was stable seasonally. Lynx fed mostly on hares, roe deer and birds (usually tetraonids) year-round. However, the share of roe deer in lynx diet increased significantly during the period of its higher abundance and the share of tetraonids decreased with their decreasing numbers. There was also a remarkable increase of lynx population, which followed that of the roe deer, despite the pronounced decline of tetraonids. The results of the study emphasised the importance of roe deer as a prey of the Eurasian lynx.     

Feeding patterns by roe deer and rabbits on pine, willow and birch in relation to spatial arrangement

The location of a plant and its association with the surrounding vegetation may be a strong determinant of herbivore foraging decisions. The attractiveness of a food plant may be reduced if associated with a less preferred one (repellent–plant hypothesis) or, conversely, it may escape herbivory via association with a more preferred species (attractant–decoy hypothesis). In this study we tested the hypothesis that selection of the same food plants by two herbivore species with different body mass, i.e. roe deer and rabbits, is affected by the spatial disposition of preferred plants in relation to less preferred ones. We planted willows, birches and pines, representing food species of higher, intermediate and lower preference, respectively, in different spatial arrangements, to manipulate patch quality and accessibility to herbivores. Contrasting patches were constructed by planting willows or pines in the midst of birches and single-species arrays of all three species, in a blocked design, replicated six times in an area occupied by both roe deer and rabbits. Preference patterns were studied by recording browsing on current years' shoots and older plant parts. Across all trees in the experiment, we confirmed that roe deer clearly preferred willow over birch, pine was not browsed at all. There was greater herbivory by roe deer on birches occurring peripherally in an array, when the associated central species was willow or pine, as compared to birches in a single-species birch patch. Therefore food choices by roe deer appear to be determined by both patch and individual level selection. The rabbits only showed preference at a species level and did not respond to spatial arrangement of trees. The results from this study indicate that the location and association of trees can strongly affect foraging patterns of roe deer, but not in a manner wholly consistent with the repellent–plant or the attractant–decoy hypotheses.     

Diet quality and immunocompetence influence parasite load of roe deer in a fragmented landscape

The influence of landscape structure and host diet on parasite load of wildlife is still largely unknown. We studied a roe deer (Capreolus capreolus) population in a fragmented agricultural landscape in southern France to explore the relationship of gastrointestinal nematode load with spleen mass (to index immunocompetence), faecal nitrogen (to index diet quality), landscape structure and age of 33 hunt-harvested roe deer. Gastrointestinal worm counts were negatively related to faecal nitrogen and spleen mass, explaining respectively 24.7% and 9.2% of the observed variability in parasite load. Landscape structure did not appear to have a direct influence on gastrointestinal worm counts, but since animals from more open areas have a diet that is richer in nitrogen, its influence may be indirect. In conclusion, in the study area, the colonisation of the agricultural landscape does not seem to have increased the risk of gastrointestinal nematode parasitism for roe deer, possibly because access to high-quality food enhances immunocompetence.

     

How does oak mast seeding affect the feeding behavior of sympatric red and roe deer?

Oak reproduction is characterized by mast seeding with high inter-annual fluctuations in fruit production. Such resource pulses can greatly affect ecosystem functioning and may cause seed consumers to alter their mobility, demography, or diet. Consequences of mast seeding for seed consumers remain poorly understood as their long timescale makes them difficult to study. We investigated impacts of oak mast seeding on the feeding behavior of two sympatric European deer species: red deer (Cervus elaphus) and roe deer (Capreolus capreolus). We analyzed their rumen content over a 31-year period in tandem with 10 years of data on oak fructification (i.e. 8 years of field monitoring and two modelled years). Acorn production is strongly correlated with consumption by both deer species. In years of high fructification, acorns represent more than 50% and 35% of red and roe deer diet, respectively, confirming assumptions that deer favor acorns when these are available. Red deer eat more acorns than roe deer both between and within years. High acorn production in mast years appears to saturate the capacity of deer to consume acorns. As the proportion of acorns increase in their diet, red deer eat more grasses and less conifer browse. No dietary shift was found for roe deer. By inducing dietary shifts in consumers, oak mast seeding can have cascading effects on ecosystem processes, notably on the damages on conifers caused by red deer and the consequences for forest dynamics.     

Spatial variability in wolf diet and prey selection in Galicia (NW Spain)

We studied wolf (Canis lupus) diet for three different landscapes in the north-western Iberian Peninsula, differing in land uses and availability of food for wolves. We examined 2740 scats, collected over a period of 4 years, in order to describe wolf diet, its geographic variation, and trophic preferences. The most consumed species were wild pony, roe deer and cattle. We observed differences in wolf diet among the three study sites, related to the availability and accessibility of food resources in each habitat. For the two study sites in northern and central Galicia, wolves showed similar diet, with high occurrence of wild pony (37 vs. 34%) and cattle (20 vs. 23%), but differing in the consumption of wild ungulates (16 vs. 8%) and carrion (7 vs. 14%). Wolf diet in eastern Galicia’s mountain ranges was entirely different, due to the higher consumption of wild ungulates (70%). Wolves showed clear prey selection patterns. Between wild ponies and livestock, wolves positively selected ponies. Among wild ungulates, wolves positively selected roe deer. Wild pony and roe deer are key species for wolf feeding in Galicia. In the Galician wild pony range, ponies are the main food for wolves. Given that the availability of wild ponies may contribute to the decrease in wolf predation on cattle, it is essential to develop innovative administrative decisions in such areas to preserve this traditional equid population. In the same way, the population of roe deer should be strengthened in the livestock areas outside the range of wild pony.     

Evaluation of the importance of roe deer fawns in the spring–summer diet of red foxes in southeastern Norway

Red fox Vulpes vulpes predation on roe deer Capreolus capreolus fawns has the potential to strongly affect prey population dynamics, but it is unclear whether this relationship is symmetrical or not. We analysed the spring–summer diet of adult foxes and of their cubs in a fragmented agricultural area of southeastern Norway, where a parallel study showed that the predator kills annually 25% of the radio-monitored roe deer fawns. The overall diet was highly varied and was dominated by small mammals (33% volume), especially Microtus agrestis, and medium-large mammals (25%), largely represented by fawns. The frequency of occurrence (FO) of fawns in the diet of adult foxes was highest in early spring, thus, supporting previous studies showing that the predator started actively hunting for fawns from the very beginning of the birth season. During the summer, the FO of both fawns and small mammals markedly declined, while that of berries and invertebrates increased. As expected for central-place foragers, cubs consumed a higher proportion of large prey items compared to adults. In particular, 25% of scats from cubs—versus 9% from adults—contained roe deer remains, suggesting a high profitability of fawns for vixens raising offspring. However, considering the wide food spectrum and the availability of several large prey items in our study area, it seems unlikely that the importance of fawns to the diet and population dynamics of red foxes could be as great as the impact of the predator on roe deer populations. This asymmetrical relationship implies that there are unlikely to be any stabilising feedback mechanisms in the predator–prey relationship.     

Diet of Eurasian lynx, Lynx lynx, in the boreal forest of southeastern Norway: the relative importance of livestock and hares at low roe deer density

The year-round food habits of lynx were studied using radio-telemetry and snow-tracking in the boreal forest of southeastern Norway. The main objectives of the study were to clarify the importance of domestic sheep and small prey species in the diet of lynx in an area with a very low density of roe deer. During the period 1995–1999, we found 193 scats and 358 kills made by lynx. Our results indicate that roe deer were the most common prey species (contributing to 83 and 34% of the biomass consumed in winter and summer, respectively), although a wide range of other species were also found, including mountain hares, tetranoids, red foxes, domestic sheep, wild reindeer, and even moose. Most of the diet was obtained by predation, although we did document several cases of scavenging. Roe deer were more important in the diet in winter than in summer, perhaps because they were easier to locate in winter as they clustered around feeding sites. In summer, domestic sheep and small prey increased in importance. Despite the very low density of roe deer in this study area, lynx seemed to still specialise on them, although domestic sheep did constitute a significant amount to their diet, especially for males and yearlings. However, the contribution of sheep to summer diet was far from that expected if their relative density was considered.     

Antibodies to granulocytic Ehrlichia in moose, red deer, and roe deer in Norway

Serum samples from 104 moose (Alces alces), 124 red deer (Cervus elaphus) and 114 roe deer (Capreolus capreolus), collected from different counties in southern Norway from 1994 to 2000, were analysed by an indirect immunofluorescent antibody staining method for antibodies to Ehrlichia equi. The overall seroprevalences for granulocytic Ehrlichia spp. in moose, red deer, and roe deer from Ixodes ricinus infested counties were 43%, 55%, and 96%, respectively. Antibody prevalence was significantly higher in roe deer than in moose and red deer (P < 0.001). Mean antibody titers (log10 +/- SD) to E. equi in sera from moose, red deer, and roe deer were 1:1,497 (3.17 +/- 0.646), 1:234 (2.37 +/- 0.424) and 1:676 (2.83 +/- 0.404), respectively. The present work indicates that all these wild ruminant species are exposed to granulocytic Ehrlichia in Norway.     

Intolerance amongst deer species at feeding: roe deer are uneasy banqueters

The function of inter-specific interactions in large herbivore assemblies is under debate. We have studied inter-specific interactions between roe and fallow deer on pastures, to assess whether competition between these Cervidae occurs through behavioural interference. Roe were displaced by fallow deer (i.e. the former moved away from the latter at a distance of >50 m) in 83% of cases (N = 127). Ninety-four percentage (N = 83) of displacement events occurred while roe deer were feeding; in 50% of these cases (N = 78), roe stopped grazing and left the feeding ground. Even when fallow deer did not show any sign of direct aggression to roe, these moved away from fallow in 72% of cases (N = 127). Vigilance rate was significantly greater in roe than in fallow deer, irrespectively of the presence of the other species. When roe and fallow deer grazed within 50 m from each other, vigilance rate increased significantly in roe, but not in fallow deer. Roe deer, in a group, were significantly more tolerant of the presence of fallow deer (even in group), than when solitary. Fallow deer seemed to be able to exclude roe deer from feeding sites through behavioural intolerance.     

Food competition between a large ruminant and a small hindgut fermentor: the case of the roe deer and mountain hare

In this study, we demonstrate that the mountain hare and roe deer compete with each other. This was determined using "natural experiments" of populations found in sympatry and allopatry on the islands along the west coast of Norway. We demonstrate that both species occupy the same habitats, share the same food resources and that food availability is limited. Two browsing species as different in size as roe deer and mountain hare might be expected to partition the available vegetation (e.g. woody scrub) in terms of height above ground level. However, from the evidence collected, the feeding-height-separation hypothesis must be rejected as an explanation for ecological separation between roe deer and mountain hares because there was extensive height overlap in resource utilisation by both species and neither species changed its feeding height in response to the presence of the other. Total browse utilisation did not increase when both species were together; rather, species-specific browse utilisation declined when the other species was present. However, the foraging behaviour of each herbivore varied significantly between the allopatric and sympatric sites. When both herbivores were present, the clip diameter of shoots browsed by mountain hares declined to match those selected by roe deer, while roe deer switched from a browse-dominated diet to a diet dominated by winter-green gramineae. The change to smaller-diameter shoots likely resulted in the hare increasing its intake of digestion-inhibiting or toxic secondary metabolites, while the alternative choice of digging through the snow like roe deer to reach the winter-green gramineae is a practice considered energetically too costly for hares. On this basis, we conclude that the enforced switch to a nutritionally inferior diet by mountain hares at the sympatric sites may result in changes to growth rate and body size which consequently impact on mortality and may explain the competitive superiority of the roe deer.