|European hare range
(dark red - native, red - introduced)
The European hare (Lepus europaeus), also known as the brown hare, is a species of hare native to Europe and parts of Western Asia and Central Asia. It is a mammal adapted to temperate, open country. It is related to and looks very similar to the European rabbit, which is in the same family but in a different genus. Hares are larger than the European rabbit, have longer ears and hind legs and breed on the ground rather than in a burrow. They rely on speed to escape from predators.
Generally nocturnal and shy in nature, hares change their behaviour in the spring, when they can be seen in broad daylight chasing one another around fields and meadows. During this spring frenzy, they can be seen striking one another with their paws ("boxing"). For a long time, this had been thought to be competition between males, but closer observation has revealed it is usually a female hitting a male, either to show she is not yet ready to mate or as a test of his determination. This species has a fairly long breeding season which lasts from January to August. Hares are herbivorous and feed on grasses, herbs, twigs, buds, bark and field crops. Their natural predators include birds of prey, canids and felids.
The European hare is listed as being of Least Concern by the IUCN. However it is declining in mainland Europe because of changes in farming practices. The hare has been a traditional symbol of fertility and reproduction in some cultures, and its courtship behaviour in the spring inspired the English idiom mad as a March hare.
Taxonomy and genetics
The European hare was first described by German zoologist Peter Simon Pallas in 1778. It shares the genus Lepus with other hares and jackrabbits. These animals are distinguished from other leporids by their longer legs, wider nostrils and precocial young. The former two are adaptations for long distance running. Historically, up to 30 subspecies of European hare have been classified, although their status has been variable. These subspecies have been distinguished by differences in pelage colouration, body size, external body measurements, and skull and tooth shape. The Corsican hare, Broom hare and Granada hare were at some points considered to be subspecies of the European hare, however DNA sequencing and morphological analysis has supported them as separate species.
There is some debate as to whether the European hare and Cape hare are the same species. A 2005 nuclear gene pool study supported this position. However, a 2006 study of the mtDNA of these same animals concluded they had diverged enough to be considered separate species. A 2008 study claims that in the case of Lepus hares, with their rapid evolution, species designation cannot be based solely on mtDNA but should also include an examination of the nuclear gene pool. It is possible that the genetic differences between the European and Cape hare is due to geographic distance rather than actual divergence. It has been speculated that in the Near East, hare populations are intergrading and experiencing gene flow. Another 2008 study suggests that more research is needed before a conclusion is reached as to whether there is a species complex. As of 2008, the European hare remains classified as a single species until further data shows otherwise.
Cladogenesis analysis suggests that European hares survived the Last glacial period during the Pleistocene via refugia in southern Europe (Italian peninsula and Balkans) and Asia Minor. Subsequent colonisations of Central Europe appear to have been initiated by human-caused environmental changes. Genetic diversity in current populations is high with no signs of inbreeding. In addition, gene flow appears to be bias towards males and but overall populations are matrilineally structured. There appears to be particularly great genetic diversity in hares in the North Rhine-Westphalia region of Germany. However, it is possible that restricted gene flow could change this within populations that become isolated. Based on molecular phylogenetic studies, the Cantabric population in Spain has unique mtDNA in relation to other European populations.
The European hare is one of the largest living members of Lagomorpha. Its head and body length can range from 48 to 75 cm (19 to 30 in) with a tail length of 7 to 13 cm (2.8 to 5.1 in). The body mass can range from 2.5 to 7 kg (5.5 to 15.4 lb). As with all leporids, the hare has elongated ears which, in this species, ranges from 9.4 to 11.0 cm (3.7 to 4.3 in) from the notch. It also has long hind feet that have a length from 14 to 16 cm (5.5 to 6.3 in). The fur colour is grizzled yellow-brown on the back; rufous on the shoulders, legs, neck and throat; white on the underside and black on the tail and ear tips. The European hare’s fur does not turn completely white in the winter, although the sides of the head and base of the ears do develop white areas. The limb musculature of hares is adapted for high-speed endurance running in open areas. By contrast, cottontail rabbits are built for short bursts of speed in more covered habitats.
Range and habitat
European hares are native to much of continental Europe. Their range extends from northern Spain to southern Scandinavia, eastern Europe and northern parts of the Middle East and Central Asia. They have been extending their range into Siberia. They may have been introduced to Britain by the Romans (circa 2000 years ago) as there are no records of them from earlier sites. They have also been introduced, mostly as game animals, to North America (in Ontario and New York State, and unsuccessfully in Pennsylvania, Massachusetts, and Connecticut), South America (Brazil, Chile, Argentina, Uruguay, Paraguay, Bolivia, Peru and the Falkland Islands), Australia, both islands of New Zealand and the south Pacific coast of Russia.
Hares primarily live in open fields with scattered brush for shelter. They are very adaptable and thrive in mixed farmland. According to a study done in the Czech Republic, the mean hare densities were highest at altitudes below 200 metres (660 ft), 40 to 60 days of annual snow cover, 450 to 700 millimetres (18 to 28 in) of annual precipitation, and a mean annual air temperature of around 10 °C (50 °F). With regards to climate the study found that hare densities were highest in: "A warm and dry district with mild winter and longer duration of sunshine; a warm and dry district with mild winter and shorter duration of sunshine; a warm and moderately dry district with mild winter".
In the United Kingdom, hares are seen most frequently on arable farms, especially those with fallow land, wheat and sugar beet crops. In mainly grass farms their numbers are raised when there are improved pastures, some arable crops and patches of woodland. They are seen less frequently where foxes are abundant or where there are many buzzards. They also seem to be fewer in number in areas with high rabbit populations. Although they are shot as game when they are plentiful, this is a self-limiting activity and is less likely to occur in localities where they are scarce.
Behaviour and life history
Hares are primarily nocturnal and spend a third of their time foraging. During daytime, a hare hides in a depression called a "form" where it is partially hidden. Hares can run at 70 km/h (43 mph) and when confronted by predators they rely on outrunning them in the open. Hares may be preyed on by canids, felids and birds of prey. They are generally thought of as asocial but can be seen in both large and small groups. They do not appear to be territorial, living in shared home ranges of around 300 ha (740 acres). Hares communicate with each other by a variety of visual signals. To show interest they raise their ears, while lowering the ears warns others to keep away. When challenging a conspecific, a hare thumps its front feet; the hind feet are used to warn others of a predator. A hare squeals when hurt or scared and a female makes "guttural" calls to attract her young.
Food and foraging
European hares are primarily herbivorous. They may forage wild grasses and weeds but with the intensification of agriculture, they have taken to feeding on crops. During the spring and summer, they feed on soy, clover and corn poppy as well as grasses and herbs. During autumn and winter, hare primarily select winter wheat as well as tubers of sugar beet and carrot which are provided by hunters. They also eat herbage, twigs, buds and the bark of shrubs and young fruit trees during winter. Cereal crops are usually avoided. The species appears to prefer high energy content over crude fibre. They have been known to eat their own green, pellet feces to recover proteins and vitamins. Two to three adult hares can eat more food than a single sheep.
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.
Mating and reproduction
European hares have a prolonged breeding season which lasts from January to August. Sexual maturity occurs at seven or eight months for females and six months for males. 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 autumn, the resting period for breeding activity, the size and activity of the males' testes increase, signalling the start of a new reproductive cycle. This continues through December, January and February and the reproductive tract gains back its functionality. Matings start before ovulation with the first pregnancies containing one foetus and pregnancy failures being common. Full reproductive activity begins in March and April, when all the females may become pregnant, the majority with three or more foetuses.
Females have six-weekly reproductive cycles and are receptive for only a few hours in one day. Thus competition among local bucks is intense. This phenomenon is known as "March madness" as it observed in March as the nights, the bucks' preferred time for activity, are shorter and thus forces them to be active in the daytime. In addition to dominants 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. 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 males as previously believed. When a doe is ready to mate, she runs across the countryside, starting a chase that tests the fitness of the following males. When only the most dominant male remains, the female stops and allows the male to copulate.
Female fertility continues through May, June and July, however 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. 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 weigh about 100 grams (3.5 oz) at birth. The leverets (young hares) are fully furred and are precocial, being ready to leave the nest as soon as they are born, an adaption to the lack of physical protection relative to that afforded by a burrow. A mother visits the nest to nurse her leverets for five minutes a day. Young can eat solid food after two weeks and are weaned when they are four weeks old. Hares can live for as long as twelve years.
The European hare is listed as being of Least Concern by the IUCN as it is considered to be wide-ranging and moderately abundant throughout its geographic distribution. However, population declines have occurred since the 1960s, possibly caused by the intensification of agricultural practices. In low population densities, hare are vulnerable to local extinctions. The Bern Convention lists the hare under Appendix III as a protected species. Several countries have placed the species on their Red Lists as "near threatened" or "threatened". The hare is considered a pest in some areas and is known to damage crops. They are also hunted as game animals. Additional threats to the hare are the diseases European brown hare syndrome, pasteurellosis, yersiniosis (pseudo-tuberculosis), coccidiosis and tularaemia, which are the principal sources of mortality.
In human culture
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 with the persecution of the church as it was commonly hunted.
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. 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. 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 nothing is known about Ēostre outside of Bede's single passage, that the authors had seemingly accepted the identification of Ēostre with the Norse goddess Freyja, yet 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."
In art and literature
Observation of the hare's springtime mating behaviour led to the popular English idiom mad as a March hare. The hare is a character in some fables, such as The Tortoise and the Hare of Aesop. It also appears in Alice in Wonderland by Lewis Carroll, in which Alice participates in a crazy tea party with the March Hare and the Mad Hatter. The German Renaissance artist Albrect Durer realistically depicted a hare in his 1502 watercolour painting Young Hare.
Hare is traditionally cooked by jugging: a whole hare is cut into pieces, marinated and cooked 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.
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