Lepus europaeus Pallas 1778

53. European Hare Lepus europaeus French: Liévre d'Europe / German: Feldhase / Spanish: Liebre europea Other common names: Brown Hare, European Brown Hare Taxonomy. Lepus europaeus Pallas, 1778, type locality not given. Restricted by Esl. Trouessart in 1910 to “Poland.” It was formerly included in L. capensis based on a cline in morphological characteristics (mainly size) from north-east ern Africa eastward across the northern Arabian Peninsula, Middle East, and northward through Israel to Turkey. In Kazakhstan, where the large “ europaeus ” and the small “ capensis ” (= tolar) live in sympatry without hybridization, the interpretation was of the overlapping ends of a ring species. A reanalysis showed a discontinuity between smaller “ capensis ” and larger “ europaeus ” running from the eastern Mediterranean coast through Iran. This is the basis on which L. europaeusis separated from L. capensis and L. tolai. In Iran, east of the border of the distribution of L. europaeus, L. tolai occurs apparently in allopatry or parapatry with L. europaeus. Nevertheless, recent evidence suggests that the hypothesis of conspecific status of L. europaeus and L. capensis may be correct. A study of the nuclear gene pool of L. capensis, L. europaeus, and North African species of Lepus indicated that North African species and L. europaeus belong to L. capensis. Nevertheless, a study of mtDNA of these three groups indicated a significant degree of divergence supporting species-specific designation. These studies show that genetic differentiation between L. capensis and L. europaeus might be attributed to geographical separation rather than divergence. It has been speculated that gene flow may occur in the Middle East where distributions meet and may result in intergraded populations. Until conclusive evidence of a single species complex is available to support a change in taxonomic status of L. europaeus, it remains a distinct species. Subspecies are recognized based mainly in size and color. Studies conducted in the USSR on differences in summer fur color and fur structure showed a tendency for clinal variation. Geographic variation has been shown for winter fur color and metric characters of skulls of L. europaeus. Thus the reality of existence of all subspecific forms described for the USSR seems very doubtful. As taxonomists are still trying to clarify the species differentiation in Lepus, the subspecific taxonomy is not elaborated yet. Hence, the distinction in subspecies might be arbitrary and unreasonable throughout the range of L. europaeus. Molecular phylogenetic studies in Spain have shown that the Cantabrian population has unique mtDNA relative to other European populations. Sixteen subspecies recognized. Subspecies and Distribution. L.e.europaeusPallas,1778—WesternEurope. L. e. caspicus Hemprich & Ehrenberg, 1832 — Lower Volga, Kalmykia (Russia) and W Kazakhstan. JR e. connor Robinson, 1918 — NW Iran. e. creticus Barrett-Hamilton, 1903 — Crete (Greece). a e. cyprius Barrett-Hamilton, 1903 — Cyprus. e. cyrensis Satunin, 1905 — Azerbaijan, Transcaucasia. a e. hybridus Desmarest, 1822 — Baltic States, Belarus, Ukraine, Finland, W & C Russia. Sl e. judeae Gray, 1867 — Palestine. aE e. karpathorum Hilzheimer, 1906 — Carpathian Mts. all e. medius Nilsson, 1820 — Denmark. al e. occidentalis de Winton, 1898 — Great Britain. ul e. parnassius Miller, 1903 — C Greece. el. e. ponticus Ognev, 1929 — Black Sea coast (Russia). ul. e. rhodius Festa, 1914 — Rhodes (Greece). Bl e. syriacus Hemprich & Ehrenberg, 1832 — Syria. ab. e. transsylvanicus Matschie, 1901 — E & SE Europe. The European Hare originated from grasslands in the Middle East, and from there, it has spread into agricultural areas of Europe. The distribution today includes European lowland areas from S Scandinavia to the N & NE Iberian Peninsula and several Mediterranean Is (Crete, Rhodes, Cyprus), and extends E to W Siberian and W Mongolian lowlands and S to N Israel, N Syria, N Iraq, the Tigris-Euphrates Valley, and NW Iran. Ancient introduction into Great Britain (shaded in the map). In more recent times this species was widely introduced as a game species into S Sweden, N Ireland, Corsica, SE Canada to NE USA, South America (Argentina, Brazil, Chile, Uruguay, and Paraguay), Australia, New Zealand, and several islands, including Barbados, Réunion, and the Falkland (Malvinas) Is. Descriptive notes. Head—body 550-680 mm, tail 75-140 mm, ear 100-140 mm, hindfoot 124-185 mm; weight 3.5-5 kg. The European Hare has tawny or rusty fur. Dorsal pelage is darker than chest and sides. Fur on back is long and curled. Ventral pelage is white. Ears are pale gray, with large triangular black patches on backs of tips. Tail is black on top and white underneath. There is some white on sides of head and bases of ears, and haunches are gray during winter. Geographical variation is low among mainland populations due to high mobility and occupation of continuous habitats. Geographical variation is also obscured by translocations by humans. Habitat. Temperate open woodland, farmland with pastures, grasslands, moorland, saltmarsh, steppes, and sub-desert from sea level to elevations of ¢.2300 m. High densities of European Hares are reached on arable land. In Chile, Argentina, Australia, and New Zealand, where no other species of Lepus occurs, European Hares also inhabits pampas, sand dunes, marshes, and alpine fellfield (slopes). Many surveys have investigated habitat preferences of European Hares. In all studies,fallow land is preferred consistently, and residential areas are avoided. There is a positive association with abundance and habitat diversity. Food and Feeding. European Hares feed predominantly on cultivated crops, weeds, and grasses. A study investigating dietary preferences showed that they positively selected forbs and grasses. Furthermore, chemical analysis revealed that they select food for high-energy content (crude fat and crude protein) and avoid crude fiber. During feeding, dominance hierarchies among individuals have been demonstrated. Breeding. Reproduction of European Hares takes place from winter solstice until the end of summer, with a peak in spring (“March madness”). During the peak, males congregate around a female in estrus. The female boxes and chases off males. Female European Hares have 1-5 litters/year, with an average ofthree litters per year. Females have between 4-6 young/year in Argentina and 13 young/year in the Czech Republic, but in most populations, females are remarkably consistent at ¢.10 young/year. Litter size can vary with respect to season, with smaller litters produced earlier and larger litters later. Superfetation (a female ovulates while pregnant and then carries both embryos and fetuses) is common in captivity and reduces normal gestation of¢.42 days to ¢.38 days. In the wild, superfetation is less common. Nursing takes place once per day, about one hour after sunset, and lasts only a few minutes. Females reach maturity at 7-8 months old and males at ¢.6 months old. Young from the firstlitters of the year may breed in theirfirst year. Where sympatric, European Hares and Mountain Hares (L. timidus) can hybridize and produce fertile offspring. Activity patterns. European Hares are mostly nocturnal. In winter, activity starts with regularity shortly after sunset and ends shortly before sunrise. In summer, activity is less consistently tied to the dark period so that active European Hares can be regularly observed in full daylight. With GPS telemetry, it was possible to gain new knowledge on daily activity patterns of European Hares during summer. If sunset was early and sunrise late, evening and morning activity peaks occurred during the dark phase. This was the case in late spring and early autumn when nights are longer. When sunset was late and sunrise was early, activity peaked in full daylight. Movements, Home range and Social organization. Distances between areas used during activity and inactivity are 122-306 m in agricultural areas and are shorter when field sizes are small than when field sizes are large. Home ranges are 12-330 ha, depending on study area. A recent study using GPS-collars showed that European Hares in an agricultural landscape with small field sizes used a confined space during the daily resting phase (on average 200 m?*), and home ranges during the active phase were c.3 ha. The European Hare lives alone but can be observed in small groups during reproduction season or when densities are high. Status and Conservation. Classified as Least Concern on The IUCN Red List. The European Hare is widespread and mostly abundant throughout its distribution. Since 1910, however, populations evidently have decreased throughout Europe due to industrialization of agriculture. The European Hare is listed under Appendix III of the Bern Convention on the Conservation of European Wildlife and Natural Habitats. Several countries have placed the European Hare on their red lists as near threatened or threatened. There is growing concern regarding status of regional forms of the European Hare. A meta-study, reviewing literature from twelve European countries and comparing population densities related to habitat characteristics, concluded that the primary cause of declines of the European Hare was agricultural intensification. Abundance was negatively associated to predation. Field size, temperature, precipitation, and hunting had no effect on densities. In Greece, Spain, France, and Denmark, restocking of European Hares from other regions or countries to supplement densities for hunting has been identified as a threat to regional gene pools. Diseases such as the European Brown Hare Syndrome (EBHS), pasteurellosis, yersiniosis (pseudotuberculosis), coccidiosis, and tularaemia are additional threats because they have been identified as principle causes of mortalities of European Hares in some regions. Bibliography. Angermann (1972, 1983, 2016)

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