European hare

European hares are primarily herbivorous. They may forage for wild grasses and weeds but with the intensification of agriculture, they have taken to feeding on crops when preferred foods are not available.[1] During the spring and summer, they feed on soy, clover and corn poppy[27] as well as grasses and herbs.[20] During autumn and winter, they primarily choose winter wheat, and are also attracted to piles of sugar beet and carrots provided for them by hunters.[27] They also eat twigs, buds and the bark of shrubs and young fruit trees during winter.[20] Cereal crops are usually avoided when other more attractive foods are available, the species appearing to prefer high energy foodstuffs over crude fibre.[28] When eating twigs, hares strip off the bark to access the vascular tissues which store soluble carbohydrates. Compared to the European rabbit, food passes through the gut more rapidly in the hare, although digestion rates are similar.[18] They sometimes eat their own green, faecal pellets to recover undigested proteins and vitamins.[19] Two to three adult hares can eat more food than a single sheep.[20]

Hare pellets

European hares forage in groups. Group feeding is beneficial as individuals can spend more time feeding knowing that other hares are being vigilant. Nevertheless, the distribution of food affects these benefits. When food is well-spaced, all hares are able to access it. When food is clumped together, only dominant hares can access it. In small gatherings, dominants are more successful in defending food, but as more individuals join in, they must spend more time driving off others. The larger the group, the less time dominant individuals have in which to eat. Meanwhile, the subordinates can access the food while the dominants are distracted. As such, when in groups, all individuals fare worse when food is clumped as opposed to when it is widely spaced.[29]

Hares fighting (above) and running during "March madness"

European hares have a prolonged breeding season which lasts from January to August.[30][31] Females, or does, can be found pregnant in all breeding months and males, or bucks, are fertile all year round except during October and November. After this hiatus, the size and activity of the males' testes increase, signalling the start of a new reproductive cycle. This continues through December, January and February when the reproductive tract gains back its functionality. Matings start before ovulation occurs and the first pregnancies of the year often result in a single foetus, with pregnancy failures being common. Peak reproductive activity occurs in March and April, when all females may be pregnant, the majority with three or more foetuses.[31]

The mating system of the hare has been described as both polygynous (single males mating with multiple females) and promiscuous.[32] Females have six-weekly reproductive cycles and are receptive for only a few hours at a time, making competition among local bucks intense.[30] At the height of the breeding season, this phenomenon is known as "March madness",[31] when the normally nocturnal bucks are forced to be active in the daytime. In addition to dominant animals subduing subordinates, the female fights off her numerous suitors if she is not ready to mate. Fights can be vicious and can leave numerous scars on the ears.[30] In these encounters, hares stand upright and attack each other with their paws, a practice known as "boxing", and this activity is usually between a female and a male and not between competing males as was previously believed.[20][33] When a doe is ready to mate, she runs across the countryside, starting a chase that tests the stamina of the following males. When only the fittest male remains, the female stops and allows him to copulate.[30] Female fertility continues through May, June and July, but testosterone production decreases in males and sexual behaviour becomes less overt. Litter sizes decrease as the breeding season draws to a close with no pregnancies occurring after August. The testes of males begin to regress and sperm production ends in September.[31]

Newborn leverets in a depression

Does give birth in hollow depressions in the ground. An individual female may have three litters in a year with a 41- to 42-day gestation period. The young have an average weight of around 130 grams (4.6 oz) at birth.[34] The leverets are fully furred and are precocial, being ready to leave the nest soon after they are born, an adaptation to the lack of physical protection relative to that afforded by a burrow.[20] Leverets disperse during the day and come together in the evening close to where they were born. Their mother visits them for nursing soon after sunset; the young suckle for around five minutes, urinating while they do so, with the doe licking up the fluid. She then leaps away so as not to leave an olfactory trail, and the leverets disperse once more.[20][35] Young can eat solid food after two weeks and are weaned when they are four weeks old.[20] While young of either sex commonly explore their surroundings,[36] natal dispersal tends to be greater in males.[32][37] Sexual maturity occurs at seven or eight months for females and six months for males.[1]

Golden eagle with a freshly caught European hare

European hares are large leporids and adults can only be tackled by large predators such as canids, felids and the largest birds of prey.[19] In Poland it was found that the consumption of hares by foxes was at its highest during spring, when the availability of small animal prey was low; at this time of year, hares may constitute up to 50% of the biomass eaten by foxes, with 50% of the mortality of adult hares being caused by their predation.[38] In Scandinavia, a natural epizootic of sarcoptic mange which reduced the population of red foxes dramatically, resulted in an increase in the number of European hares, which returned to previous levels when the numbers of foxes subsequently increased.[39] The golden eagle preys on the European hare in the Alps, the Carpathians, the Apennines and northern Spain.[40] In North America, foxes and coyotes are probably the most common predators, with bobcats and lynx also preying on them in more remote locations.[34]

European hares have both external and internal parasites. One study found that 54% of animals in Slovakia were parasitised by nematodes and over 90% by coccidia.[41] In Australia, European hares were reported as being infected by four species of nematode, six of coccidian, several liver flukes and two canine tapeworms. They were also found to host rabbit fleas (Spilopsyllus cuniculi), stickfast fleas (Echidnophaga myrmecobii), lice (Haemodipsus setoni and H. lyriocephalus), and mites (Leporacarus gibbus).[42]

European brown hare syndrome (EBHS) is a disease caused by a calicivirus similar to that causing rabbit haemorrhagic disease (RHD) and can similarly be fatal, but cross infection between the two mammal species does not occur.[43] Other threats to the hare are pasteurellosis, yersiniosis (pseudo-tuberculosis), coccidiosis and tularaemia, which are the principal sources of mortality.[44]

In October 2018, it was reported that a mutated form of the rabbit haemorrhagic disease virus (RHDV2) may have jumped to hares in the UK. Normally rare in hares, a significant die-off from the virus has also occurred in Spain.[45][46]

In Europe, the hare has been a symbol of sex and fertility since at least Ancient Greece. The Greeks associated it with the gods Dionysus, Aphrodite and Artemis as well as with satyrs and cupids. The Christian Church connected the hare with lustfulness and homosexuality, but also associated it with the persecution of the church because of the way it was commonly hunted.[47]

In Northern Europe, Easter imagery often involves hares or rabbits. Citing folk Easter customs in Leicestershire, England, where "the profits of the land called Harecrop Leys were applied to providing a meal which was thrown on the ground at the 'Hare-pie Bank'", the 19th-century scholar Charles Isaac Elton proposed a possible connection between these customs and the worship of Ēostre.[48] In his 19th-century study of the hare in folk custom and mythology, Charles J. Billson cites folk customs involving the hare around Easter in Northern Europe, and argues that the hare was probably a sacred animal in prehistoric Britain's festival of springtime.[49] Observation of the hare's springtime mating behaviour led to the popular English idiom "mad as a March hare",[47] with similar phrases from the sixteenth century writings of John Skelton and Sir Thomas More onwards.[50] The mad hare reappears in Alice's Adventures in Wonderland by Lewis Carroll, in which Alice participates in a crazy tea party with the March Hare and the Mad Hatter.[51]

Sir John Tenniel's March Hare with Alice, the Dormouse, and the Hatter from Alice's Adventures in Wonderland, 1865

Any connection of the hare to Ēostre is doubtful. John Andrew Boyle cites an etymology dictionary by A. Ernout and A. Meillet, who wrote that the lights of Ēostre were carried by hares, that Ēostre represented spring fecundity, love and sexual pleasure. Boyle responds that almost nothing is known about Ēostre, and that the authors had seemingly accepted the identification of Ēostre with the Norse goddess Freyja, but that the hare is not associated with Freyja either. Boyle adds that "when the authors speak of the hare as the 'companion of Aphrodite and of satyrs and cupids' and 'in the Middle Ages [the hare] appears beside the figure of [mythological] Luxuria', they are on much surer ground."[52]

The hare is a character in some fables, such as The Tortoise and the Hare of Aesop.[53] The story was annexed to a philosophical problem by Zeno of Elea, who created a set of paradoxes to support Parmenides' attack on the idea of continuous motion, as each time the hare (or the hero Achilles) moves to where the tortoise was, the tortoise moves just a little further away.[54][55] The German Renaissance artist Albrecht Dürer realistically depicted a hare in his 1502 watercolour painting Young Hare.[56]

Painting of gentlemen hunting hares by Richard Ansdell

Across Europe, over five million European hares are shot each year, making it probably the most important game mammal on the continent. This popularity has threatened regional varieties such as those of France and Denmark, through large-scale importing of hares from Eastern European countries such as Hungary.[5] Hares have traditionally been hunted in Britain by beagling and hare coursing. In beagling, the hare is hunted with a pack of small hunting dogs, beagles, followed by the human hunters on foot. In Britain, the 2004 Hunting Act banned hunting of hares with dogs, so the 60 beagle packs now use artificial "trails", or may legally continue to hunt rabbits.[57] Hare coursing with greyhounds was once an aristocratic pursuit, forbidden to lower social classes.[58] More recently, informal hare coursing became a lower class activity and was conducted without the landowner's permission;[59] it is also now illegal.[60] In Scotland concerns have been raised over the increasing numbers of hares shot under license.[61]

Hare is traditionally cooked by jugging: a whole hare is cut into pieces, marinated and cooked slowly with red wine and juniper berries in a tall jug that stands in a pan of water. It is traditionally served with (or briefly cooked with) the hare's blood and port wine.[62][63] Hare can also be cooked in a casserole.[64] The meat is darker and more strongly flavoured than that of rabbits. Young hares can be roasted; the meat of older hares becomes too tough for roasting, and may be slow-cooked.[63][65]

Photograph of a hare on farmland

The European hare has a wide range across Europe and western Asia and has been introduced to a number of other countries around the globe, often as a game species. In general it is considered moderately abundant in its native range,[13] but declines in populations have been noted in many areas since the 1960s. These have been associated with the intensification of agricultural practices.[66] The hare is an adaptable species and can move into new habitats, but it thrives best when there is an availability of a wide variety of weeds and other herbs to supplement its main diet of grasses.[1] The hare is considered a pest in some areas; it is more likely to damage crops and young trees in winter when there are not enough alternative foodstuffs available.[20]

The International Union for Conservation of Nature has evaluated the European hare's conservation status as being of least concern. However, at low population densities, hares are vulnerable to local extinctions as the available gene pool declines, making inbreeding more likely. This is the case in northern Spain and in Greece, where the restocking by hares brought from outside the region has been identified as a threat to regional gene pools. To counteract this, a captive breeding program has been implemented in Spain, and the relocation of some individuals from one location to another has increased genetic variety.[1] The Bern Convention lists the hare under Appendix III as a protected species.[25] Several countries, including Norway, Germany, Austria and Switzerland,[1] have placed the species on their Red Lists as "near threatened" or "threatened".[67]