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.     

The effect of experimental removal of red foxesVulpes vulpes on spring density of brown haresLepus europaeus in western Poland

Red foxesVulpes vulpes (Linnaeus, 1758) were experimentally removed in two nearby areas located in western Poland to verify the hypothesis about the limiting impact of their predation on the low-density population of brown haresLepus europaeus (Pallas, 1778) (4.4-10.6 ind./km² in late autumn). In 1996/1997–2001/2002 foxes were culled (mainly in autumn and winter) in the reduction area (32 km²), whereas in the control area (34 km²) intensive culling was carried out only in 2000/2001–2001/2002. Indices of fox and hare spring densities were estimated using spotlight counts, as mean numbers of individuals observed per 10 km of the counting route. Annual changes in the fox density indices were negatively correlated with the bag of foxes, and annual changes in the hare density indices were negatively related to the annual changes in fox density indices. The fox density indices were significantly lower in the reduction area than in the control one only in 2000–2001 (2.8 times, on average), and in the same years, the hare population responded with higher density (1.7 times, on average). The hare responses took place without time delay, which suggests that the changes in fox abundance affected the situation of hares primarily in the autumn-winter season.     

Integrating field sports,hare population management and conservation

Conflicts between field sports, animal welfare and species conservation are frequently contentious. In Ireland, the Irish Coursing Club (ICC) competitively tests the speed and agility of two greyhounds by using a live hare as a lure. Each coursing club is associated with a number of discrete localities, known as preserves, which are managed favourably for hares including predator control, prohibition of other forms of hunting such as shooting and poaching and the maintenance and enhancement of suitable hare habitat. We indirectly tested the efficacy of such management by comparing hare abundance within preserves to that in the wider countryside. In real terms, mean hare density was 18 times higher, and after controlling for variance in habitat remained 3 times higher, within ICC preserves than the wider countryside. Whilst we cannot rule out the role of habitat, our results suggest that hare numbers are maintained at high levels in ICC preserves either because clubs select areas of high hare density and subsequently have a negligible effect on numbers or that active population management positively increases hare abundance. The Irish hare Lepus timidus hibernicus Bell, 1837 is one of the highest priority species for conservation action in Ireland and without concessions for its role in conservation, any change in the legal status of hare coursing under animal welfare grounds, may necessitate an increase in Government subsidies for conservation on private land together with a strengthened capacity for legislation enforcement.     

The German wildlife information system: population densities and development of European Hare (<Emphasis Type="Italic">Lepus europaeus</Emphasis> PALLAS) during 2002–2005 in Germany

The German Wildlife Information System, founded in 2001, is a long-term monitoring program documenting occurrence, number, and development of game populations throughout Germany. Population numbers are recorded by standardized counting methods in so-called reference areas. The population densities of the European hare are calculated by spotlight strip censuses in the reference areas each spring and autumn all across Germany. From 2002 to 2005, the censuses were carried out by local hunters in 510 to 676 reference areas each year. During these years, the calculated spring densities increased significantly from 11.0 (2002) to 14.5 hares/km² (2005) nationwide. The overall increase in spring densities was primarily caused by the population rise from spring 2003 to 2004, which correlates with the high net growth rate in 2003. In 2005, the number of counted hares varied between less than 1 and more than 107 hares/km² in spring and between 0 and more than 170 hares/km² in autumn. Because of differing landscapes in Germany, three regions were differentiated. In spring 2005, the average population densities (median) in East Germany (5.4 hares/km²) and Southwest Germany (14.6 hares/km²) were significantly lower than in Northwest Germany (23.9 hares/km²). These regional differences had been similarly distinct in former years.     

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.     

Numerical responses by populations of red fox and mountain hare during an outbreak of sarcoptic mange

Summary During a severe outbreak of sarcoptic mange (Sarcoptes scabiei vulpes) starting among red foxes (Vulpes vulpes) in Sweden in the 1970s, we studied: 1) the establishment and spread of the disease in northernmost Sweden (by inquiries), and 2) the 1970–84 bag records for foxes and mountain hares (Lepus timidus) (an alternative prey to the fox's main prey, voles). Since the first case of sarcoptic mange in 1975 the disease spread rapidly, with >50% of the hunting organizations having reported the disease in 1981 and >75% in 1983. Also the disease became more abundant within the areas affected. In areas with a low mange infection rate (index) the number of foxes killed in the 1980s did not deviate markedly from the average level in the 1970s. However, there was a slight tendency towards a decline in areas with a medium index and numbers declined markedly where the index was high. Hare harvests initially were low (after a tularemia epidemic) in the 1970s. In that decade harvests increased dramatically and stabilized, increased gradually or changed little, respectively, where mange infection rates were low, medium or high in the early 1980s. In areas with a low mange index hare harvests remained cyclical and at the same level in the 1980s as in most of the 1970s. However, in areas with a medium index harvests increased and seemed to begin to lose their cyclicity, and where the index was high the low and relatively stable hare harvests increased annually. A predator-prey hypothesis, assuming predators to synchronize alternative prey declines to those of the cyclic main prey, predicts that a predator reduction would cause a gradual disappearance of the cyclicity and increasing numbers among alternative prey. Our hare data are partially consistent with this prediction.     

Niche overlap of mountain hare subspecies and the vulnerability of their ranges to invasion by the European hare; the (bad) luck of the Irish

Niche conservatism is the tendency of related species to retain ancestral tolerances after geographic separation. We used Ecological Niche Modelling and Principal Components Analysis of bioclimatic and habitat variables to describe the extent of the species niche, and degrees of bioclimatic–habitat niche conservatism within the mountain hare (L. timidus) clade. Mountain hare niche space was contrasted with that of the European hare (L. europaeus), to shed light on species interactions in contact zones throughout Europe. All five subspecies of mountain hare had quantifiably distinct niches. Fennoscandian (L.t. sylvaticus, L.t. timidus) and highland (L.t. scoticus, L.t. varronis) subspecies, however, were most similar, exhibiting greatest apparent niche conservatism. They inhabit tundra, boreal forest and uplands, and, hence are presumed most similar to the ancestral form. The Irish hare was distinct, being consistently distinguished from other mountain hares in both 2D and nth dimensional (4D) niche space. The ecological distinctiveness of the Irish hare provides further evidence that it is an Evolutionarily Significant Unit, particularly vulnerable to displacement by introduced European hares with which it competes and hybridises. Projections under global climate change suggest that, by 2070, bioclimatic space for invasive European hares in Ireland will expand (by 79%) but contract for endemic Irish hares (by 75%), further facilitating their replacement. The near complete species replacement of the heath hare (L.t. sylvaticus) in southern Sweden, where the European hare has also been introduced, may suggest a similar fate may be in store for the Irish 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.     

Long-term changes in the feeding pattern of red foxes Vulpes vulpes and their predation on brown hares Lepus europaeus in western Poland

The aim of this study was to estimate long-term changes in the winter feeding pattern of red foxes Vulpes vulpes and in their predation on brown hares Lepus europaeus in relation to the decreasing abundance of hares in western Poland in 1965/1966–2006/2007. The frequencies of occurrence in the stomachs of culled foxes (N?=?726) were used as indices of prey capture rates. The average autumn density of brown hares in the study area decreased from 48 individuals/km² at the turn of the 1960s and 1970s to seven individuals/km² in 1999–2006. Hares and small rodents were the main food classes of foxes in western Poland at the turn of the 1960s and 1970s; however, the occurrence of hares in the fox diet subsequently decreased, and they were replaced by livestock carrion. The relationship between the occurrence frequency of hares in the fox diet and the hare density was best described by sigmoid equation. It indicates that the red fox showed a type III functional response to long-term changes in hare abundance. When predation rate index was estimated on the basis of functional response, the potential fox predation was density-dependent at low to intermediate hare densities (<25 individuals/km²). This finding suggests that the increase in the number of low-density hare populations may require intensive management measures, e.g. simultaneous use of fox control and habitat improvement.     

Distribution of mountain hares Lepus timidus in Scotland: results from a questionnaire

A questionnaire survey of land owners, managers and gamekeepers was conducted in order to assess the distribution of mountain hares in Scotland, assess their current management, collate numbers harvested in 2006–07 and estimate distribution change by comparing with similar data collected in 1995–96. The land area covered by returned questionnaires was 71098km² (90% of Scotland). Mountain hares were reported as present on 34359km² (48%) and absent from 36739km² (52%). Mountain hare presence was strongly associated with heather moorland managed for red grouse shooting. Moorland managed for driven grouse shooting had the highest percentage area of mountain hare presence (median 64%) followed by moorland managed for walked‐up grouse shooting (median 9%) and moorland with no grouse shooting (median 0%). Approximately 25000 mountain hares were harvested in 2006–07. Based on the estimated UK population in 1995 of 350000 (range ±50%), this represents around 7% of the population (range 5–14%). Reasons given by respondents for harvesting hares were tick control (50%), sport (40%) and forestry or crop protection (10%). Comparison of the estates surveyed in both 2006–07 and 1995–96 (a total area of 20462km²) indicated no net gain or loss in hare distribution. Furthermore, there was no evidence that levels of harvest had reduced the range of mountain hares in this area. It is not possible to comment on any distribution change outside this area (58737km²). Similarly, as no data were collected on abundance, it is not possible to draw conclusions on changes in density. Regular monitoring of mountain hare distribution within Scotland is required to identify any distribution changes. Measures of abundance throughout the range are necessary to estimate the population size, investigate the relationship between harvest intensity and changes in abundance and further assess the conservation status of this UK Biodiversity Action Plan species.     

Mountain hare populations on islands: effects of predation by red fox

Summary On islands off the west coast of Sweden the density of mountain hares (Lepus timidus L.) is very high. One of the main predators on hares, the red fox (Vulpes vulpes L.), is only present during short periods. Data on hare density and predation by red fox and eagle owl (Bubo bubo (L.)) has been analyzed from five islands over several years. Winter mortality in years with low predation pressure was independent of hare density. But when red fox or eagle owl were present on islands (i.e., high predation pressure) winter mortality became density dependent. Thus, at low density, winter mortality did not increase through red fox predation. But at densities up to two hares/ha, predation pressure was increasing and could be limiting for these populations. At still higher hare density predation pressure became less intensive. The functional response for foxes preying on hares showed a type II or a sigmoid type III response pattern. In normal summers, the population increase due to reproduction was at least two-fold. When a fox was present there was instead a sharp decrease in hare numbers. Fox predation had a stronger effect in summer than in winter. By switching between islands and mainland areas from winter to summer, a fox can stabilize fluctuations in hare numbers on the islands. This is dependent on how often the ice permits a fox to reach an island and the lack of numerical response by predators.     

Temporal dynamics of European brown hare syndrome infection in Northern Italian brown hares (Lepus europaeus)

The progressive decline in the hare population across Europe has been associated with the occurrence of European brown hare syndrome (EBHS), a highly contagious disease considered endemic in all European countries. This study aimed to evaluate the in-field temporal dynamics of European brown hare syndrome virus (EBHSV) infection in wild European brown hares (Lepus europaeus) and to test the influence of population density on EBHS seroprevalence. A total of 512 blood samples were collected from free ranging hares captured for restocking in seven different areas of the province of Brescia (Northern Italy) during seven consecutive years (2006–2013) and tested using a competitive ELISA. A generalized linear mixed model estimated the yearly effects of population density on EBHS prevalence. Of the 512 tested, 344 (67.2 %) tested positive for EBHSV antibodies, with the annual seroprevalence ranging from 94.3 to 35.8 %. The prevalence was 3.303 times higher in areas with a density of over 15 hares/km² and declined over the years. The results indicate the ongoing transmission of the virus in the tested brown hare population. Since the eradication of EBHS in a wild population is not feasible, a strategy aimed at promoting the endemic stability of the virus through density-dependent mechanisms could be applied; however, this seems more difficult in practice than in theory and would most likely require a very high density of brown hares.     

The changing status of the Brown hare (Lepus capensis L.) in Britain

Game bag records are used to examine temporal and geographical changes in numbers of Brown hares. Records are taken from a survey made by the Oxford Bureau of Animal Population in 1938, which covered over 400 estates, and from the Game Conservancy's National Game Census, which has been monitoring shooting records of between 400 and 500 estates since 1961. Eastern regions of Britain had higher bags of hares than western regions in the three periods; 1891–1910, 1917–1936, 1961–1978. Hare game bags were generally high in counties with large proportions of tilled land. There was no consistent temporal trend in bags from different areas before 1938, but all areas show a considerable reduction in bags since 1961. This reduction is statistically significant. Time-series analysis of one extended set of data did not show evidence of cyclic fluctuations.     

Techniques for assessing the abundance of Brown Hares Lepus europaeus

Over the last few decades, there have been significant declines in Brown Hare Lepus europaeus numbers throughout Europe, leading to concern for their status in many countries. In Britain, there were no quantified data on the extent of this decline, on current population levels, or any baseline against which to monitor future population changes. The need for a quantified national hare survey led to this evaluation of the techniques available to assess hare numbers. Published information on counting hares is reviewed, and various techniques compared by applying them to a number of sites in southern England. Three basic approaches are available: counts of inactive hares, counts of active hares and indirect methods. Counts of inactive hares include total clearance, wide belt and line transect counts. Total clearance counts give an absolute figure, but are labour intensive and can only be applied to restricted areas. Wide-belt assessments are difficult to apply in certain habitats and even in open areas tend to produce a substantial over-estimate. Line transect counts are easy to undertake and are not labour intensive but should only be applied to large areas, or data from several small areas combined. Counts based on active hares are more problematical, because it is difficult to determine what proportion of the population is inactive at any one time. Spotlight counts based on variable circular plots are the most accurate but difficult to apply widely, and twilight counts are very subjective in their interpretation, especially when surveying small areas or areas with a large proportion of concealing habitats. Of the indirect methods, dung pellet counts can be valuable in specific areas but are difficult to apply across a range of habitats. We concluded that, of the various techniques considered, line transect counts have the greatest potential for a national survey, but need to be stratified so that enough transects are undertaken within each habitat stratum to obtain a reliable mean population estimate for each stratum.     

Canada lynx occurrence and forest management in the Acadian Forest

We evaluated patterns of occurrence and non-occurrence for Canada lynx (Lynx canadensis) across a 16,530-km² study area in Maine to provide a better understanding of lynx habitat selection and habitat ecology on commercially managed forestlands in the Acadian Forest. Because of the influence of forest structure on lynx habitat selection and abundance of their primary prey, the snowshoe hare (Lepus americanus), and to improve our ability to build robust models, we used habitat information derived from a time series of Landsat satellite imagery spanning the period 1973–2004. We defined and mapped 10 forest types based on forest harvest history, time since harvest, and current forest condition. We compared a suite of models to evaluate relative influences of forest composition, habitat patch configuration, and hare density on habitat selection by lynx at the landscape scale. Occupied areas had greater average hare densities and percentage of mature conifer. Average hare density in occupied areas (0.74 hares/ha) was greater than in unoccupied areas (0.62 hares/ha), but was less than previous research has suggested may be necessary to support lynx populations in the southern portion of the species' range. No occupied areas occurred where average hare density was <0.5 hares/ha. Average hare density at the landscape-scale was strongly influenced by amount of high-quality hare habitat (i.e., conifer or mixedwood regenerating forest, 15–35 yr post-harvest). Edge density between mature conifer and high-quality hare habitat was substantially greater in occupied areas compared to unoccupied areas. Juxtaposition of those 2 forest types may provide edge habitat where lynx experience easier travel and improved access to prey in landscapes with extensive areas of high-quality hare habitat where travel and access may be somewhat limited by high understory stem density. Probability of occurrence declined nonlinearly with changes in hare density and percent mature conifer forest in the landscape; thus, suitability of currently occupied landscapes could change markedly with future changes in landscape-level hare densities and changing habitat associated with forest management. Where lynx conservation is a priority, we recommend that managers focus on creating and maintaining a minimum of 27% high-quality hare habitat within 100-km² areas to promote landscape-scale hare densities >0.5 hares/ha. © The Wildlife Society, 2013     

Limited effect of hare grazing and short-term climatic variations on the most common alpine vegetation community in the Snowy Mountains,Australia

Background: In the alpine zone of the Snowy Mountains, grazing by mammals is limited. However, introduced European hare numbers have increased since the 1970s.

Aims: To estimate the density of hares and hence grazing pressure among years. To assess the response of biomass, vegetation height and composition to a cessation of hare grazing.

Methods: We used indices of hare abundance based on spotlighting and counts of hare pellets on a transect. The effect of hare grazing on tall alpine herbfield was assessed by using 15 paired exclosure and control quadrats for six years.

Results: The indices of hare abundance suggested densities similar to those in upland areas of Britain. Grazing did not affect the composition, cover of herbs or graminoids or, for 2010, vegetation height or biomass. Variation in vegetation and hare numbers among years was not correlated with climatic variables. Observations of selective grazing suggested that impacts on vegetation may be localised and restricted to certain species. Prior analyses of hare pellets indicated that hares might spread seed of native and exotic species.

Conclusions: Hares are having no general effect on tall alpine herbfield but may affect certain plant species via selective grazing or by spread of viable seed.     

Museum samples provide novel insights into the taxonomy and genetic diversity of Irish red grouse

The taxonomic status of red grouse in Ireland has been the subject of considerable debate over the past century. Irish red grouse are usually classified as Lagopus lagopus scoticus, which is the same subspecies as that found in Britain, but some ornithologists believe that native Irish red grouse constitute an endemic subspecies, namely L. lagopus hibernicus. The considerable decline of Irish red grouse over the past century, along with possible hybridization with introduced grouse from Britain, have highlighted the need to resolve their taxonomic status as part of a biodiversity management plan. However, genetic analysis of samples from a single point in time will provide limited insight into potentially confounding historical events such as hybridization and introgression. We therefore compared mtDNA sequences from both current and historical samples of the two putative subspecies, scoticus and hibernicus, to see if they are or were genetically distinct. Red grouse from Britain and Ireland shared mitochondrial haplotypes, and our historical data suggest that this is unlikely to be the result of recent hybridization and introgression. These findings, combined with a general lack of documented differences in behaviour and ecology, suggest that Irish red grouse should remain classified as L. lagopus scoticus. At the same time, we found evidence that a significant amount of genetic diversity has been lost from Irish red grouse over the past century, presumably as a result of diminishing population sizes and fragmentation of extant populations. A loss of habitat, combined with the declining numbers and genetic diversity of Irish red grouse, justify their designation as an All-Ireland Priority (Red List) species and a Northern Ireland Priority Species for conservation.     

Molecular Approaches Revealing Prehistoric, Historic, or Recent Translocations and Introductions of Hares (Genus Lepus) by Humans

Although only of medium size, and thus of little nutritional value compared to big game such as mammoths and ungulates, hares (Lepus spp.) probably have always been a food source for humans, as documented in archaeological finds. Nowadays, hares, particularly such species as the brown hare (L. europaeus), are among the most important game species in many European countries. For hunting, perhaps religious reasons, and in connection with certain myths, hares have been and are still being intentionally translocated. Ancient translocations by humans can be inferred from the presence of hares on islands that had no mainland connections, at least during the Pleistocene, the major evolutionary period of the genus Lepus. We review some of the literature on anthropogenic translocations of hares. We focus on three examples [the brown hare (L. europaeus), the Corsican hare (L. corsicanus), and the Sardinian hare (L. capensis)], where some molecular data could be used to trace the translocation routes and possible origins of introduced hare populations. Certain molecular marker systems, such as sequences of the hypervariable part I (HV-1) of the mitochondrial control region, show high variability in hare species and are thus promising for tracing both recent and ancient origins of translocated hares. Some other molecular marker systems as well as caveats connected with the use of such marker systems in the genus Lepus are also discussed.     

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.