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/km(2). Game bag records indicate that hare densities may have been much higher in the past, with a maximum of 138 hares/km(2) 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 21000 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.

The diet and landclass affinities of the Irish hare Lepus timidus hibernicus

The diet of the Irish hare Lepus timidus hibernicus was investigated in Northern Ireland. Faecal pellets were collected at regular intervals from three contrasting study sites and from a number of randomly selected sites from known land classes. Microhistological techniques were employed to analyse the pellets. Grass species occurred in high percentage frequencies in all samples of faecal pellets and a wide variety of plant species were represented (up to 26 species in one study site). In a stratified, random survey of the presence of hares, areas of lowland intensive agriculture had fewer positive records than expected. It is contended that L. t. hibernicus may not be able to sustain viable populations in areas of intensive agriculture, which are almost exclusively composed of ryegrass Lolium spp.

Post-release GPS tracking of hand-reared Irish hare Lepus timidus hibernicus leverets, Slemish, Co. Antrim, Northern Ireland.

Animal rescue centres release large numbers of captive-bred, rehabilitated or translocated animals into the wild annually but little is known about their post-release survival and behaviour. We developed a novel and innovative coupling of traditional radio-tags with new GPS loggers to track hand-reared Irish hare Lepus timidus hibernicus leverets after release into the wild. Cyanoacrylate SuperGlue® proved a poor fixative with two out of three leverets managing to detach their tags within 24 hours. Nevertheless, a total of 2,505 GPS locations were recorded every 60 seconds for one leveret over three nights (approx. 835 per night). The leveret dispersed

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.

First record of minimum Irish hare (Lepus timidus hibernicus Bell 1837) longevity in the wild.

Average longevity of the mountain hare (Lepus timidus L., 1758) has been estimated at nine years in the wild (Macdonald D. and Barrett, P. 1993 Mammals of Britain and Europe. Harper Collins Publishers, London) with a maximum recorded age of 18 years for one marked animal (Angerbjörn, A. and Flux, J. E. C. 1995 Lepus timidus. Mammalian Species 495: 1–11). However, the longevity of the Irish hare (L. t. hibernicus Bell 1837) is entirely unknown. A total of 14 Irish hares was trapped and tagged at Belfast International Airport, Co. Antrim from February to April 2005. The sex, age (juvenile or adult) and weight of each animal were recorded. Adults were taken as those individuals >8-10 months old defined by the fusing of the notch between the apophysis and diaphysis of the tibia and humerus (Flux, J. E. C. 1970 Journal of Zoology 161: 75-123). Individual identification was made by a system of colourcoded ear tags (Roxan iD Ltd. Selkirk, Scotland) being inserted in the centre of the pinna of each ear. Each ear tag (6 × 34 mm) and puncture site was disinfected with 70 per cent ethanol prior to insertion. An adult male, #001/002 ‘Blue/Blue’, was tagged on 3 March 2005 weighing 3.8 kg and was sighted during a return site visit on 4 April 2007. An adult female, #026/003 ‘Green/Yellow’, was tagged on 15 April 2005 weighing 4.0 kg and was sighted during return visits on 25 March 2010 and 19 October 2010. The latest possible date of birth for both individuals was spring/summer 2004. Consequently, they were at least 3 years and 6.5 years old, respectively. This is the first record of minimum Irish hare longevity in the wild. These observations suggest that ear tagging does not compromise animal welfare and is an effective means of long-term monitoring. Future research may utilize capture-mark-recapture methods.

Range expansion and comparative habitat use of insular, congeneric lagomorphs: invasive European hares Lepus europaeus and endemic Irish hares Lepus timidus hibernicus

The European hare (Lepus europaeus) has declined throughout its native range but invaded numerous regions where it has negatively impacted native wildlife. In southern Sweden, it replaces the native mountain hare (L. timidus) through competition and hybridisation. We investigated temporal change in the invasive range of the European hare in Ireland, and compared its habitat use with the endemic Irish hare (L. timidus hibernicus). The range of the European hare was three times larger and its core range twice as large in 2012–2013 than in 2005. Its rate of radial range expansion was 0.73 km year−1 with its introduction estimated to have occurred ca. 1970. Both species utilised improved and rough grasslands and exhibited markedly similar regression coefficients with almost every land cover variable examined. Irish hares were associated with low fibre and high sugar content grass (good quality grazing) whilst the invader had a greater tolerance for low quality forage. European hares were associated with habitat patch edge density, suggesting it may be more suited to using hedgerows as diurnal resting sites than the Irish hare. Consequently, the invader had a wider niche breadth than the native but their niche overlap was virtually complete. Given the impact of the European hare on native species elsewhere, and its apparent pre-adaption for improved grasslands interspersed with arable land (a habitat that covers 70 % of Ireland), its establishment and range expansion poses a significant threat to the ecological security of the endemic Irish hare, particularly given their ecological similarities.

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.

Is naturalisation of the brown hare in Ireland a threat to the endemic Irish hare?

On islands, one of the greatest risks to native wildlife is the establishment of alien species. In Ireland, the Irish hare (Lepus timidus hibernicus), the only native lagomorph, may be at risk from competitive exclusion and hybridisation with naturalised brown hares (L. europaeus) that were introduced during the late nineteenth century. Pre- and post-breeding spotlight surveys during 2005 in the north of Ireland determined that brown hare populations are established in mid-Ulster and west Tyrone. In mid-Ulster, brown hares comprised 53%-62% of the hare population, with an estimated abundance of 700-2000 individuals between pre- and post-breeding periods. Comparison of habitat niches suggest that Irish and brown hares have comparable niche breadths that at times completely overlap, suggesting the potential for strong competition between the species. Anecdotal evidence suggests that both species may hybridise. Further research is urgently required to assess the degree of risk that naturalised brown hares pose to the Irish hare population and what action, if any, is needed to ensure the future ecological security and genetic integrity of the native species.

Factors associated with hare mortality during coursing

Hare coursing is a widespread but controversial activity. In an attempt to reduce hare mortality and mitigate the activity's impact on hare welfare, the Irish Coursing Club introduced measures including the compulsory muzzling of dogs in 1993. However, the efficacy of these measures remained the subject of heated debate. Official records, corroborated by independent video evidence, were used to assess the fate of individual Irish hares (Lepus timiclus hibernicus) during coursing events from 1988-2004. Muzzling dogs significantly reduced levels of hare mortality. In courses using unmuzzled dogs from 1988189-1992193 mean hare mortality was 15.8%, compared to 4.1% in courses using muzzled dogs from 1993194-2003104. Further reductions in mortality could not be accounted for by muzzling dogs, supporting the efficacy of other factors such as improved hare husbandry. The duration of the head start given to the hare prior to the release of the dogs significantly affected the outcome of the course. Hares that were killed had head starts of greater duration than those that were chased but survived, suggesting the former may have been slower. The selection of hares by assessment of their running ability may provide means to reduce hare mortality during courses further. Our findings support the efficacy of measures taken to mitigate the impact of coursing on individual hares. However, it is necessary to evaluate the impact of removing hares from the source population and of returning coursed hares to the wild before the wider impact of coursing on wild hare populations can be determined.

Homogeneous habitat can meet the discrete and varied resource requirements of hares but may set an ecological trap

In common with other farmland species, hares (Lepus spp.) are in widespread decline in agricultural landscapes due to agricultural intensification and habitat loss. We examined the importance of habitat heterogeneity to the Irish hare (Lepus timidus hibernicus) in a pastoral landscape. We used radio-tracking during nocturnal active and diurnal inactive periods throughout one year. In autumn, winter and spring, hares occupied a heterogeneous combination of improved grassland, providing food, and Juncus-dominated rough pasture, providing refuge. In summer, hares significantly increased their use of improved grassland. This homogeneous habitat can fulfil the discrete and varied resource requirements of hares for feeding and shelter at certain times of year. However, improved grassland may be a risky habitat for hares as silage harvesting occurs during their peak birthing period of late spring and early summer. We therefore posit the existence of a putative ecological trap inherent to a homogeneous habitat of perceived high value that satisfies the hares' habitat requirements but which presents risks at a critical time of year. To test this hypothesis in relation to hare populations, work is required to provide data on differential leveret mortality between habitat types.

Mammals and agri-environment schemes: hare haven or pest paradise?

1. Agri-environment schemes (AESs) are designed to create landscape-scale improvements in biodiversity. While the specific aims of AESs do not always include the enhancement of species of conservation concern, associated conservation strategies, such as the UK Biodiversity Action Plan, often rest on the assumption that AESs enhance environmental conditions and thereby improve the conservation status of target species. However, there is little evidence for the general efficacy of AESs in this respect. 2. To evaluate the effects of the Environmentally Sensitive Area (ESA) scheme, a widespread AES in Northern Ireland, a spotlight survey of the relative abundance of three mammal species, Irish hare Lepus timidus hibernicus, European rabbit Oryctolagus cuniculus and red fox Vulpes vulpes, was conducted. Of these, the Irish hare is a priority species for conservation action and the focus of a species action plan, while rabbit and fox are commonly considered agricultural pests. The effects of ESA designation and habitat on each species were assessed at 150 ESA and 50 non-ESA sites, matched for landscape characteristics. 3. The ESA scheme had no demonstrable effect on the abundance of Irish hares, and this agri-environment scheme did not target the landscape and habitat variables associated with hares. 4. In contrast, the abundance of rabbits and foxes was significantly greater within ESAs than the wider countryside. Agricultural factors such as reduced livestock stocking density, reduced overgrazing and field boundary enhancements may create more favourable conditions for both species. Aside from the implications for farm economics, the proliferation of rabbit populations within conservation areas may raise issues concerning the grazing of important plant communities, while increases in fox populations may adversely affect ground-nesting birds and other animal species of conservation concern. 5. Synthesis and applications: The abundance of rabbits and foxes corroborates recent work that suggests AESs may benefit common species but can not be relied upon to encourage rarer species. The Irish hare species action plan relies on agri-environment schemes to enhance the species’ status and realize the target of increasing the hare population by 2010 by promoting suitable habitat. However, the ESA scheme is unlikely to help in achieving these objectives. Targeted and evidence-based agri-environment prescriptions are clearly required in order to ensure the realization of species-specific conservation targets.

Potential Rôle of Hares in the Spread of Liver Fluke in the Netherlands.

Hares (Lepus europeanus) sharing pasture with cattle from six locations in the Netherlands were examined for the presence of liver fluke (Fasciola hepatica) and shown to have prevalences of infection ranging from zero to 41%. The mitochondrial haplotypes of liver flukes present in the hare populations were determined and compared with those found in cattle from a farm where triclabendazole resistance has been reported. Phylogenetic analysis indicated that the flukes present in the hares belonged to the same clades as those present in the cattle. A consideration of the life cycle of the liver fluke and the seasonal breeding pattern and ecology of hares supports the suggestion that hares may act as a refugia for liver fluke and as a vector for the spread of drug-resistant genotypes.

Biogeography, macroecology and species’ traits mediate competitive interactions in the order Lagomorpha

1. In addition to abiotic determinants, biotic factors, including competitive, interspecific interactions, limit species’ distributions. Environmental changes in human disturbance, land use and climate are predicted to have widespread impacts on interactions between species, especially in the order Lagomorpha due to the higher latitudes and more extreme environmental conditions they occupy.
2. We reviewed the published literature on interspecific interactions in the order Lagomorpha, and compared the biogeography, macroecology, phylogeny and traits of species known to interact with those of species with no reported interactions, to investigate how projected future environmental change may affect interactions and potentially alter species’ distributions.
3. Thirty-three lagomorph species have competitive interactions reported in the literature; the majority involve hares (Lepus sp.) or the eastern cottontail rabbit (Sylvilagus floridanus). Key regions for interactions are located between 30-50°N of the Equator, and include eastern Asia (southern Russia on the border of Mongolia) and North America (north western USA).
4. Closely related, large-bodied, similarly sized species occurring in regions of human-modified, typically agricultural landscapes, or at high elevations are significantly more likely to have reported competitive interactions than other lagomorph species.
5. We identify species’ traits associated with competitive interactions, and highlight some potential impacts that future environmental change may have on interspecific interactions. Our approach using bibliometric and biological data is widely applicable, and with relatively straightforward methodologies, can provide insights into interactions between species.
6. Our results have implications for predicting species’ responses to global change, and we advise that capturing, parameterizing and incorporating interspecific interactions into analyses (for example, species distribution modelling) may be more important than suggested by the literature.