Scattered woody vegetation promotes European brown hare population

European brown hare populations have declined during the last decades. Agricultural intensification has been identified as a relevant driver of this process and agri-environment schemes have been implemented to foster biodiversity in agricultural landscapes. Because species-specific outcomes of measures strongly depend on tailored design of the policy framework and the local management, while changing climate may pose additional challenges, policy and management need science-based information of which landscape composition should be promoted to achieve set biodiversity goals.Here, we used direct observations of European brown hares over 20 years for evaluating the effects of landscape composition and weather conditions on European brown hare density. For the first time, our analysis compared the estimates of machine learning (gradient boosting machine) and linear mixed models in terms of importance of a wide range of explanatory variables for European brown hare densities and effect trends.Scattered woody vegetation, as represented by the two variables transitional woodland-shrub and small woody features, was on top rankings among the predictors and greater proportions of these elements were accompanied by sharp increases of European brown hare density. Also warmer winter temperature had a positive effect.We conclude that promoting scattered woody vegetation in agricultural landscapes is a powerful tool for improving European brown hare habitat quality. Particularly with the increasing dynamic in agriculture due to climate change, incentives and regulations that create a long-lasting heterogeneity in the landscape composition through near-natural elements can support the population of this popular mammal.     

Factors related to the occurrence of hybrids between brown hares Lepus europaeus and mountain hares L. timidus in Sweden

Hybridization occurs among many species, and may have implications for conservation as well as for evolution. Interspecific gene flow between brown hares Lepus europaeus and mountain hares L. timidus has been documented in Sweden and in continental Europe, and has probably to some extent occurred throughout history in sympatric areas. What local factors or ecological relationships that correlate with or trigger hybridization between these species has however been unclear. We studied spatial distribution of hybrids between brown hares and mountain hares in Sweden in relation to characteristics of the sampled localities (hunting grounds). In a sample of 70 brown hares collected from 39 populations in south‐central Sweden during 2003–2005, 11 (16%) showed introgressed mtDNA from mountain hares. Among the brown hares from their northern range, i.e. in general the most recent establishments, the corresponding figure was 75% (9/12). The frequency of samples with hybrid ancestry increased significantly with latitude, altitude and hilliness, and were higher (p<0.1) in recently established populations and/or where the proportion of arable land was low. Several site‐specific parameters were correlated, e.g. latitude as expected to hilliness, and no parameter explained the occurrence of hybrids exclusively. Instead, the appearance of mountain hare mtDNA among brown hares was associated with a conglomerate of parameters reflecting landscapes atypical for the brown hare, e.g. forest dominated and steep areas where the species quite recently was established. We suggest that these abiotic factors mirror the main aspect influencing hybridization frequency, namely the density or relative frequency of the two species. In atypical brown hare landscapes with recent establishment, mountain hares are probably relatively more common. When one species dominate in numbers, or when both species display low densities, increased frequency of hybridization is expected due to low availability of conspecific partners, a phenomenon referred to as Hubbs’ principle.     

The consequences of predator control for brown hares (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) on UK farmland

The brown hare Lepus europaeus is a valued game species but also a species of conservation concern owing to its severe decline in abundance on farmland throughout Europe during the twentieth century. Changes in the farmland habitat and predation have both been cited as causative factors. Their relative roles have been unclear, but most conservation action has focused on improving habitat. We analyse data from a sequence of three unique studies (one experiment and two demonstrations) covering the period 1985–2006 in which control of several common predator species was undertaken to increase densities of wild game on farmland in England. Across the three studies, regression modelling of the proportional change in hare numbers between successive years showed that—after site, year differences and harvesting were accounted for—predator control was a significant determinant of hare population change. Where habitat improvement also took place, hares reached autumn densities that were exceptional for the UK and which could sustain substantial harvests. When predation control was stopped, hare densities fell, even where habitat improvements remained in place. This analysis demonstrates that even where farmland habitat is greatly improved, uncontrolled predation prevents hares making full use of its carrying capacity. This helps explain the mixed—and at best modest—success of agri-environment schemes in the UK and elsewhere in Europe to increase hare densities. Game-shooting estates, on which effective predator control takes place, probably have a special significance within the landscape as source areas for brown hares.     

A quantitative analysis of the abundance and demography of European hares Lepus europaeus in relation to habitat type, intensity of agriculture and climate

1. European hares Lepus europaeus have declined throughout Europe since the 1960s. Possible reasons for this include agricultural intensification and changes in climate and predator numbers, but no clear consensus has been reached as to the relative importance of each of these. We aimed to identify factors associated with high and low hare numbers throughout Europe, to determine which could have caused population declines. 2. Results of 77 research papers from 12 European countries were summarized. Relationships between hare density and demographics and habitat, climate, hunting and predator variables were examined and quantified where possible. Temporal changes in factors identified as being associated with high or low numbers of hares were then examined to see if they could explain population declines. 3. Data from pastural habitats were limited, but densities of hares were low. Arable habitats had higher densities than mixed areas in spring, unless farming was intensive in which case densities were similar. In autumn the two habitats had similar densities. Field size, temperature, precipitation and hunting had no effect on density throughout Europe. Fecundity was affected by climate. 4. Arable land, various crops, fallow habitat and temperature were positively associated, and monoculture, precipitation and predators negatively associated with hare abundance. The relationship of field size, pasture and woodland with abundance depended on spatial scale. 5. Habitat changes caused by agricultural intensification are the ultimate cause of hare population declines. Effects of changes in climate or predator numbers are magnified by the loss of high-quality year-round forage and cover. Further research is required on how habitat changes affect fecundity and survival, and to identify which parameters have the greatest impact on population numbers. Farmland management policies that target the re-establishment of some of the habitat diversity lost within fields, farms and landscapes will help to reverse the decline of the European hare.     

The effect of landscape structure on the distribution of brown hare Lepus europaeus in farmlands of Germany and Poland

Habitat management should be an important part of the brown hare (Lepus europaeus) conservation, but the habitat requirements of this species are not fully recognised. The aim of our research was to estimate these requirements by analysing the effect of various agricultural landscape structure features on the distribution of hares in five agricultural areas in Germany and Poland. The local density of hares was assessed in the spring and autumn of 2006 by using the method of spotlight–strip counts on 9–15 subareas in each research region. The structure of agricultural landscape has been described for each subarea: the share of grain, other crops and grasses as well as the density of crop edges and uncultivated places with wild vegetation. The density of hares was considerably higher in Germany than in Poland (18.8–48.4 vs. 4.1–9.5 indiv./km²). The hare density was positively correlated with non-grain crops in an area, with crop edges in two areas and with wild vegetation without trees in two areas, and negatively correlated with grassfields in two areas. The occurrence of wild vegetation without trees affected the hare density only in the study areas, where this habitat was relatively rare (<3 km/km²). It was suggested that proper projects aimed at habitat management for brown hares should be elastic, i.e. the projects should be modified depending on the structure of local landscapes. Moreover, the protection and creation of structures with wild vegetation among cropland seem to be considerable methods of brown hare or generally wildlife conservation; therefore, such measures should be an important part of agro-environmental packages.     

Spring and autumn habitat preferences of active European hares (Lepus europaeus) in an agricultural area with low hare density

The number of European brown hares (Lepus europaeus) has been declining throughout much of Europe since the 1960s. Consequently, many studies have focused on analysing habitat selection of European hares in order to improve the suitability of the habitat for this species. Habitat preferences of European hares are known to be affected by hare density, but most studies have been conducted in agricultural areas where hare densities were medium to high. Finding habitat preferences at high densities is difficult as most available habitats are occupied. In addition, in agricultural areas, field size might influence the hares’ habitat selection because it affects the distribution and availability of certain habitat types. However, most studies relate to areas with large field sizes. In this study, we analysed the habitat preferences of European hares in spring and autumn during the activity period, in the early hours of the night, in an agricultural area with low hare density and small average field size using Chesson’s electivity index. Moreover, we focused on the question whether two different habitat classifications varying in their specificity might cause contradictory results regarding European hares’ habitat preferences. Our results show that in this agricultural area with low hare density, European hares avoided several habitat types which were preferred in other study areas with higher hare densities. Therefore, we assume that hare density has an influence on the species’ habitat selection. In contrast, the small average field size of our study area seemed not to have an effect on hare habitat preference. Furthermore, by pooling habitat types into broader groups, substantial information was lost in some categories. Hence, for some categories, e.g. grassland or agricultural crop land, more detail might be needed than for others, such as urban areas, when analysing hares’ habitat selection. In conclusion, our results imply that studies on habitat preferences have to be conducted in areas with low hare density to be able to gain knowledge on the species’ habitat requirement and hereinafter improve the suitability of the habitat for this species.     

Vegetation quality and habitat selection by European haresLepus europaeus in a pastural landscape

European haresLepus europaeus Pallas, 1778 have lower population densities and body condition in pastural landscapes than in arable landscapes, but reasons for this are not understood. The aim of this study was to determine whether forage quality is low in pastural landscapes during certain seasons. We carried out chemical analysis of the nutritional quality of 5 habitat types to determine whether hares select high quality habitats, and whether nutritional quality explains seasonal differences in range sizes of hares in pastural landscapes. Hares did not tend to select habitats of high nutritional quality (protein, fat or energy) over those of lower quality. Hares did not increase active range size as the overall energy content of forage at the study site decreased; seasonal differences in active range size were not explained by nutritional quality. Differences may be explained by behavioural changes related to breeding. Pastural habitat is fairly stable in terms of nutritional quality through the year, and results suggest that poor forage quality is unlikely to be responsible for the poor body condition of hares in pastural landscapes. Hares in these landscapes are more likely to be limited by habitat quality in terms of cover than by forage.     

The occurrence of mountain hare mitochondrial DNA in wild brown hares

If interspecific hybrids are fertile and backcross to either parental species, transmission of mitochondrial DNA over the species barrier can occur. To investigate if such transmission has occurred between the brown hare Lepus europeus Pall and the mountain hare L. timidus L. in Scandinavia, an analysis of genetic variation in mitochondrial DNA from 36 hares, collected from 15 localities, was performed. Sequence divergence of mtDNA between species was estimated at 8 ± 1% (SD). Intraspecific mtDNA sequence divergence varied between 0.09 and 0.38% in brown hares and 0.10 and 1.44% in mountain hares. In six out of 18 brown hares examined, two different haplotypes of mountain hare origin were detected, demonstrating a transmission of mtDNA haplotypes from mountain hares to brown hares. The results indicate that interspecific hybridization between the two species occurs in wild populations.     

Bioenergy crops and farmland biodiversity: benefits and limitations are scale-dependant for a declining mammal,the brown hare

Biomass energy crops are prompting major land-use changes in agricultural and marginal land in an effort to reduce dependency on fossil fuels. Miscanthus × giganteus, a perennial giant grass, is one of the main such crops in Europe but few studies exist of its interaction with farmland wildlife, particularly mammals. Understanding ecological impacts of bioenergy planting schemes is vital for mitigating potential negative effects on already declining farmland biodiversity and for maximising any benefits from these low-management, structurally diverse crops. We assessed in a mixed farming area in the UK the impact of Miscanthus crops on the brown hare (Lepus europaeus), a widespread but declining farmland species of conservation concern. We intensively radio-tracked hares in Miscanthus blocks of contrasting size and analysed hare diet for evidence of the consumption of Miscanthus. Home ranges differed starkly averaging 10.5 versus 49.6 ha in the small and the large Miscanthus blocks, respectively. Despite entirely avoiding the crop as food, hares appeared able to exist and even thrive in areas planted with Miscanthus though their populations may be significantly limited by reduced food availability and increased energy use where dense Miscanthus is planted over a wide area. As a component of a mixed farming landscape, Miscanthus may provide biodiversity benefits by increasing spatial heterogeneity and refuge areas for declining farmland species like brown hares but any effect is likely to be strongly scale-dependant.     

Modelling changes in species’ abundance in response to projected climate change

Aim Existing climate envelope models give an indication of broad scale shifts in distribution, but do not specifically provide information on likely future population changes useful for conservation prioritization and planning. We demonstrate how these techniques can be developed to model likely future changes in absolute density and population size as a result of climate change. Location Great Britain. Methods Generalized linear models were used to model breeding densities of two northerly‐ and two southerly‐distributed bird species as a function of climate and land use. Models were built using count data from extensive national bird monitoring data and incorporated detectability to estimate absolute abundance. Projections of likely future changes in the distribution and abundance of these species were made by applying these models to projections of future climate change under two emissions scenarios. Results Models described current spatial variation in abundance for three of the four species and produced modelled current estimates of national populations that were similar to previously published estimates for all species. Climate change was projected to result in national population declines in the two northerly‐distributed species, with declines for Eurasian curlew Numenius arquata projected to be particularly severe. Conversely, the abundances of the two southerly distributed species were projected to increase nationally. Projected maps of future abundance may be used to identify priority areas for the future conservation of each species. Main conclusions The analytical methods provide a framework to make projections of impacts of climate change on species abundance, rather than simply projected range changes. Outputs may be summarized at any spatial scale, providing information to inform future conservation planning at national, regional and local scales. Results suggest that as a consequence of climate change, northerly distributed bird species in Great Britain are likely to become an increasingly high conservation priority within the UK.     

The recent expansion of the brown hare (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) in Sweden with possible implications to the mountain hare (<Emphasis Type="Italic">L. timidus</Emphasis>)

The brown hare (Lepus europaeus) expanded its Swedish distribution since the 1980s northwards and locally to new areas within its former range. Of 115 brown hare populations within the former range reported in a hunter enquiry, those established after 1980 were situated higher above the sea level than older ones and higher than neighbouring (<50 km) older populations. Reports on increased use of forest habitats by brown hares were equally frequent among recent and older populations, suggesting a process promoted solely by less harsh winters. Supposed hare hybrids were more often reported from hunting grounds with recent brown hare establishment, i.e. where the species expands in time and in space. In a 27-year dataset on brown hare observations, the recent increased use of forest habitats was supported in that maximum distances to agricultural land for brown hare sightings were higher in mild winters, whereas the proportions of the annual observations made during winter were lower. In 40-year bag records from two Swedish counties, the dynamics of the mountain hare (Lepus timidus) responded positively to snow parameters, whereas brown hares responded negatively. We suggest that the state of mountain hare populations primarily depends on winter conditions and predation pressure, whereas possible effects of hybridization are unclear. If winter conditions remain as in the last 15 years, mountain hare numbers are not likely to increase in southern Sweden, whereas the brown hare may expand even further. In either case, hybrids will occur in sympatric areas in frequencies probably related to the density of the respective true species.     

Methodological problems related to spotlight count as a census technique for Lepus europaeus in an alpine environment

Population estimations are necessary for effective conservation management. In Italy, brown hare populations are commonly censused by spotlighting, but this method does not seem to suit an alpine environment due to its vegetational and orographical complexity. The aim of this study is to evaluate the critical aspects related to spotlight census method in an alpine environment. Spotlight was carried out along two transects of a typical alpine environment. Observed animals were used to define density (number of animal seen/100 ha) and method precision (coefficient of variation (CV) applied to monthly repeated observations). Animal detectability was evaluated using half-normal function with cosine expansion (Distance 5.0®). Animal observability was evaluated by analyzing density estimates related to habitat conformation (unseen areas or full visibility). The exact surface surveyed by spotlighting was evaluated, defining the observation spotlight beam range (OTA) and the land useful sighting (LUS). In the end, LUS was classified in three patches according to hare presence: no hares, occasional hare presence, and constant hare presence. To evaluate habitat influence onto CV, we used a bootstrap simulation. The results show that spotlighting alone is not the most suitable method to apply in the alpine environment because habitat structure highly influences census results. Recommendations to improve spotlight surveys for monitoring European brown hare populations are given.     

The diet of the mountain hare (Lepus timidus hibernicus) on coastal grassland

Across most of their range in Europe, mountain hares are usually restricted to upland areas with poor food quality. In these areas they generally feed on browse species such as heather or twigs and barks of trees. On lowland areas in Europe, with better food quality, the mountain hare is replaced by the brown hare ( Lepus europaeus ) which feeds predominantly on greasses. This khas led some authors to conclude that mountain hares are primarily adapted for browsing. In the absence of brown hares in Ireland, mountain hares are found on a wide variety of habitats including grassland. On grassland, their diet consists almost exclusively of grasses, up to 94% of their annual diet, which is more than has been reported for brown hares on similar habitat. Based on this evidence, and other work, it is proposed that the mountain hare in primarily a grazing animal and competitive exclusion by brown hares may underlie much of their present distribution in Europe.     

Relationship Between Fecal Pellet Counts and Snowshoe Hare Density in Western Wyoming

ABSTRACT Snowshoe hares (Lepus americanus) are an important prey species for Canada lynx (Lynx canadensis) and are considered critical for lynx population persistence. Determination of snowshoe hare distribution and abundance is needed by land management agencies for lynx conservation. An accepted approach for estimating snowshoe hare abundance is the use of fecal-pellet plot counts. Locally derived regression equations are preferred for accurate calibration of pellet counts to snowshoe hare density due to local differences in pellet deposition and decomposition. We used linear regression to examine correlations between snowshoe hare density, as determined by mark–recapture estimates, and pellet plot counts on both uncleared plots and annually cleared plots on the Bridger-Teton National Forest, western Wyoming, USA. We found significant correlations between snowshoe hare density estimates and fecal pellet counts for both uncleared and annually cleared pellet counts; however, the relationship was stronger (higher r) when using pellet counts from annually cleared plots. In addition, we found that adjusting the buffer size by omitting hard habitat edges (not used by hares) around trapping grids improved correlations between snowshoe hare density and fecal pellet counts for both uncleared plots and annually cleared plots. Though precision is sacrificed when using uncleared plots, they may be useful as a coarse index of habitat use by snowshoe hares. Our derived regression equations may be useful to identify important foraging habitat for Canada lynx in western Wyoming. Land managers responsible for conserving snowshoe hare habitat in western Wyoming may use these equations to monitor changes in hare populations among habitats and during prescribed management actions.     

European hare (Lepus europaeus) invasion ecology: implication for the conservation of the endemic Irish hare (Lepus timidus hibernicus)

European hare Lepus europaeus populations have undergone recent declines but the species has successfully naturalised in many countries outside its native range. It was introduced to Ireland during the mid-late nineteenth century for field sport and is now well established in Northern Ireland. The native Irish hare Lepus timidus hibernicus is an endemic subspecies of mountain hare L. timidus and has attracted major conservation concern following a long-term population decline during the twentieth century and is one of the highest priority species for conservation action in Ireland. Little is known about the European hare in Ireland or whether it poses a significant threat to the native mountain hare subspecies by compromising its ecological security or genetic integrity. We review the invasion ecology of the European hare and examine evidence for interspecific competition with the mountain hare for habitat space and food resources, interspecific hybridisation, disease and parasite transmission and possible impacts of climate change. We also examine the impact that introduced hares can have on native non-lagomorph species. We conclude that the European hare is an emerging and significant threat to the conservation status of the native Irish hare. Invasive mammal species have been successfully eradicated from Ireland before and immediate action is often the only opportunity for cost-effective eradication. An urgent call is issued for further research whilst the need for a European hare invasive Species Action Plan (iSAP) and Eradication strategy are discussed.     

High mtDNA haplotype diversity among introduced Swedish brown haresLepus europaeus

The brown hareLepus europaeus Pallas, 1778 occurs naturally in central Eurasia, but has been introduced to parts of northern Europe, South- and North America, Australia and New Zealand. Brown hares were introduced to Sweden from central Europe for hunting purposes during the 19th century. We investigated how the human--mediated brown hare colonisation of Sweden is reflected in the amount of genetic variation present by assessing variation and composition of mitochondrial DNA (mtDNA) lineages among Swedish brown hares. MtDNA from a total of 40 brown hare specimens from 15 localities were analysed for Restriction Fragment Length Polymorphisms. The haplotype diversity is surprisingly high (0.893 ± 0.002) when compared to the mtDNA diversity among brown hares on the European continent as well as to other mammalian species. Admixture of haplotypes from different source populations combined with a reduced effect of random genetic drift and a relaxed selection pressure due to rapid population growth after introduction are mechanisms that are likely to account for the observed high mtDNA haplotype diversity.     

Assessment of Canada Lynx Research and Conservation Needs in the Southern Range: Another Kick at the Cat

Abstract The ecology of Canada lynx (Lynx canadensis) and their main prey, snowshoe hares (Lepus americanus), is poorly understood in southern Canada and the contiguous United States compared to the boreal forest of Canada and Alaska, USA, where both species are well studied. However, given recent listing of lynx under the Endangered Species Act, accurate understanding of lynx and snowshoe hare ecology and conservation requirements in the United States is a high priority. We critically examined unchallenged perceptions and important research needs related to lynx and hare ecology and conservation at the southern extent of their range. Contrary to popular dogma, lynx do not require old-growth forest for denning, but further research on lynx and hare use of fragmented landscapes at lower latitudes is required. The contention that southern lynx are subject to higher interference or exploitative competition compared to their northern counterparts remains without strong empirical support. Lynx rely more on red squirrels (Tamiasciurus hudsonicus) and possibly other alternate prey at lower latitudes, but hares are the predominant food type for lynx across their range. Southern lynx and hare populations do not exhibit periodic cyclicity, but harvest statistics suggest that lynx abundance in the southern range is highly variable, implying that numerical fluctuations likely are fueled by immigration from Canada. Southern lynx population viability in the absence of ingress is suspect and thus maintaining connectivity with northern areas of occupancy should be a priority. Successful conservation of lynx populations in the contiguous United States will require 1) improved understanding of lynx population and habitat ecology at lower latitudes, 2) protection and management of large tracts of lynx and snowshoe hare habitat, and 3) ensured connectivity between lynx populations at the core and periphery of the species' range. However, in light of the numerous challenges facing conservation of populations of many species at their southern distributional limit, the long-term prognosis for lynx in the southern range currently is uncertain.     

A review of the status and decline in abundance of the Irish Hare (Lepus timidus hibernicus) in Northern Ireland

The Northern Ireland Hare Survey documented the distribution of the Irish Hare (Lepus timidus hibernicus). Historical game bag records and other, more contemporary, records of hare distribution were examined. These data indicate how numbers of L. t. hibernicus may have changed over the last 140 years. The results of the Northern Ireland Hare Survey suggested that L. t. hibernicus was widespread throughout Northern Ireland. Current average densities are no more than 0.65 hares/km². Game bag records indicate that hare densities may have been much higher in the past, with a maximum of 138 hares/km² recorded on Crom Estate, Co. Fermanagh, in 1864. Evidence from hare distribution recorded during the Northern Ireland Rabbit Survey indicates that hare numbers declined between 1984 and 1994. Evidence from all sources suggests that L. t. hibernicus has declined in abundance substantially, with present total population estimates for Northern Ireland ranging from 8250 to 21 000 individuals. Flushing data indicate that rushes and hedgerows are important diurnal resting areas for hares. While the principal reason for the decline in numbers of L. t. hibernicus in Northern Ireland is not clear, more species‐rich pasture and provision of areas of cover, such as rushes, may arrest further declines, or indeed promote numbers of hares, particularly in lowland areas.     

NIGHT COUNTS OF HARES AND OTHER ANIMALS IN EAST AFRICA

Night counts of hares and other animals were made on a disused airstrip in the Queen Elizabeth National Park, Uganda and on the road between Magadi and Nairobi in Kenya. Lepus crawshayi is the only species of hare which occurs in the Queen Elizabeth Park but L. capensis is also found in Kenya. No regular seasonal fluctuations were found in the number of hares recorded due probably to the continuous breeding of these equatorial species. The number of waterbuck seen in Uganda on moonlit nights was significantly greater than that counted on dark nights. This is not thought to be due to better viewing conditions under a full moon since the phenomenon was not found with other species. The number of hares recorded varied inversely with the number of other mammals present but bore no relationship with the number of birds. It is concluded that night counts can be useful in revealing population trends but not for assessing total numbers or for comparing densities in different areas.