The European hare (Lepus europaeus), also known as the brown hare, eastern jackrabbit and eastern prairie hare, is a species of hare native to northern, central, and western Europe and western Asia. It is a mammal adapted to temperate, open country. It is related to the similarly appearing rabbit, which is in the same family but a different genus. It breeds on the ground rather than in a burrow and relies on speed to escape.

Normally shy animals, hares change their behaviours in the spring, when they can be seen in broad daylight chasing one another around meadows. During this spring frenzy, hares can be seen "boxing", where hares strike one another with their paws. 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 quite ready to mate or as a test of his determination.^[3]

The hare is declining in mainland Europe due to changes in farming practices. Its natural predators include the golden eagle and carnivorous mammals, such as the red fox and wolf. Smaller hares native to southern Europe previously regarded as European hares have been split off as separate species in recent years, including the broom hare in northern Spain.

Taxonomy and genetics

One study of the nuclear gene pool of the European hare and the Cape hare suggests they may well be the same species.^[4] However, a later study of the mtDNA of these same animals concluded they diverged significantly and are thus different species.^[5] It has been suggested that in the case of hares whose evolution is rapid, species designation cannot be based solely on mtDNA and there needs to be examination of the nuclear gene pool.^[6] It seems the genetic differentiation between the European and Cape hare is caused by geographic distance rather than divergence.^[6] It has been speculated that in Near Eastern, hares are experiencing gene flow where populations meet and this may leave to intergraded populations.^[6] It has been proposed that "a combined phylogenetic, phylogeographic, and population genetic approach,…, based on various nuclear and mitochondrial markers and including other biological characters, such as phenotypic and morphometric data," are needed for conclusive evidence of a single species complex.^[7] However, the European hare remains classified as a true species until there is data found supporting a change its taxonomic status.^[2]

There appears to be genetic diversity in the European hare in the North Rhine-Westphalia region of Germany.^[8] However, it is possible that restricted gene flow could change this within populations that become isolated.^[8] Based, on molecular phylogenetic studies the Cantabric population in Spain has unique mtDNA in relation to other European populations.^[9]

Subspecies

A European hare (above) compared with a mountain hare

There are 15 recognized subspecies of European hare.^[10]

• Lepus europaeus caspicus
• Lepus europaeus connori
• Lepus europaeus creticus
• Lepus europaeus cyprius
• Lepus europaeus cyrensis
• Lepus europaeus europaeus
• Lepus europaeus hybridus
• Lepus europaeus judeae
• Lepus europaeus karpathorum
• Lepus europaeus medius
• Lepus europaeus occidentalis
• Lepus europaeus parnassius
• Lepus europaeus ponticus
• Lepus europaeus rhodius
• Lepus europaeus syriacus
• Lepus europaeus transsylvanicus

Description

Portrait of hare

The European hare has a head and body length ranging from 600-750 mm with a tail length of 72–110 mm.^[11] There is no noticeable sexual dimorphism in the species. As with all leporids, the hare has elongated ears which in this species ranges from 94–102 mm from the notch. The ears of the European hare are greyish white inside and have black tips on the top ends. It also has long hind feet that have a length from 142 to 161 mm. Most of the hare’s body is covered in yellowish-brown to greyish-brown fur but has greyish-white fur on the underside. In addition its face is brown with black rings around the eyes. Unlike some other leporids, the European hare’s fur does not turn white in the winter, but it does get slightly more grey.^[11][12] The hare’s skull has a length from 96 to 104 mm and a width from 44 to 51 mm.^[11] The skull has nasal bones that are short, broad and heavy as well as prominent anterior and posterior lobes of the supraorbital processes.^[13] In addition, the skull has a prominent subcutaneous process of the lacrimal bone, projecting from the anterior wall of the orbit.^[14]

Range and habitat

Hare running in open field

The European hare ranges from continental Europe though the Middle East and into central Asia. It was probably introduced to Great Britain in ancient times, partially replacing its close relative, the Mountain hare.^[15] In more recent centuries the hare has been introduced to many other areas around the world: Eastern North America, South America, Australia, New Zealand and many islands including Tasmania, the Falklands, Barbados and Reunion.^[16] The species was imported to North America from Germany by a farmer living near Cambridge, Ontario, Canada in 1912. It escaped from the farm, successfully colonized fields and woodland edges, and quickly made the "Jackrabbit" a common sight in southern Ontario, New York State and New England.

Hares primarily live in open fields and pasture usually near agricultural areas and bordered by hedgerows and woodlots. They prefer to live in shallow forms like clumps of grass, weeds or brush.^[11] According to a study done in the Czech Republic the mean hare densities were highest in habitat with elevations from sea level to 200 m (231.47/10 km²), annual snow cover duration from 40–60 days (183.95/10 km²); mean annual precipitation: 450–700 mm (174.71/10 km²), annual sunshine duration: 1801-2000 (169.72/10 km²) and mean annual air temperature of around 10.0˚C (245.00/10 km²).^[17] Climatic areas with the highest mean densities were found to have been "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".^[17]

Ecology and behavior

European hare displaying typical hiding behaviour at Uitkerkse Polders, Belgium

Outside of the mating season, the European hare lives a largely solitary lifestyle. It is mostly nocturnal and crepuscular and forages between 7:00 p.m. and 7:00 a.m.^[13] During daytime, a hare will hide in a depression called a "form" where they are partially hidden.^[14] Hares are capable of running in speeds of up to 35 mph (56 km/h) in a straight line.^[13] When running from its predators, the hare can dodge and change direction quickly. They will even dive into streams and can swim.^[14] Little evidence shows that hares stay within a restricted home range.^[14] Predators of the hare include the red fox, wolf, coyote, wild cats and birds of prey.^[14] Although they are usually quiet, hares will make low grunts and females will make "guttural" calls to her young. They emit a shrill call when caught or hurt.^[11][14]

Food and foraging

Hare pellets

European hares are primarily herbivorous. During the summer, they eat grasses, herbs and field crops.^[13] During the winter, they eat twigs, buds and the bark of shrubs and young fruit trees.^[13] They have been known to eat their own green, pellet feces to recover proteins and vitamins.^[13] Two to three adult hares can eat as much as one sheep.^[11][14] Though normally comparatively solitary, European hares will forage in groups. Group feeding is beneficial as individuals can spend more time feeding, as there is increased corporate vigilance.^[18] Nevertheless, the distribution of food affects the benefits of group foraging depend. When food is well-spaced, all hares are able to access it. When food is clumped together, dominant hares are able to monopolize this resource.^[18] In small gatherings, dominants are able to exclude subordinates, but in larger gatherings, they must devote more time to defending the patch.^[18] Therefore, dominants spend less time feeding as the group size increases, while subordinates feed more since they have more opportunities, since the dominants are chasing off others. As such, when in groups, all individuals fare worse when food is clumped as opposed to when it is spaced.^[18]

Mating and reproduction

Hares during "March madness"

European hares have a prolonged breeding season which lasts from January to August.^[3][19] At least some females, or does, have been found pregnant in all breeding months and males, or bucks, are fertile in all months of the year except in October and November.^[19] After the rest period in autumn, the size and activity of the males' testes increase in November, the first indication of a new reproductive cycle. This continues though December, January and February and the reproductive tract becomes fully functional again. Matings proceed ovulation and the first pregnancies usually have one fetus, although pregnancy failures are common during time.^[19] Full reproductive activity occurs by March and April and nearly 100% of females may become pregnant with most carrying three or more fetuses.^[19] During breeding, females are receptive for just a few hours on one day in each of their six-weekly cycles.^[3] Thus local bucks compete for a doe's favor with dominant males striving to keep the others at bay. In addition the female will fight off any male that approaches her before she is ready.^[3] This phenomenon is known as "March madness". This is because the behavior is more often observed in March as the nights, the bucks preferred time for activity, are shorter and thus forces them to be active in the daytime.^[3] A female will viciously fight off her suitors, giving them scarred ears. Hares have been observed to stand on their hind legs and hit 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 will start a wild chase across the countryside, shaking off following males until only one remains. After this the female will stop and allow the remaining male to mate with her.^[3]

Young hare hiding

The does continue to be highly fertile through May, June and July. However, the overt behaviour characteristic of spring reduces and testicular testosterone production drops.^[19] By July and August the reproductive cycle ends and there is a reversal of the changes that took place earlier in the year. In August, the testes regress rapidly and by September sperm production ceases with the sperm reserves in the epididymis becoming progressively depleted and the sperm degenerate by October. Litter sizes reduce in numbers towards the end of the breeding season and by the end of August there appear to be no more pregnancies.^[19] Young or leverets are born precocial from birth and have long, silky fur.^[11] This is because hares do not give birth to their young below ground in a burrow but rather in a form. Thus hares are adapted to the lack of physical protection, relative to that afforded by a burrow. A mother will disperse her young over a moderately large area to lessen the chance of a predator preying on the whole litter. She will make the rounds to nurse them.^[14] Young become independent at around a month old.

Status and human interactions

The March Hare with Alice, the Dormouse, and the Hatter from Alice's Adventures in Wonderland

The European hare is widespread and abundant across its geographic range.^[20] However, population declines since in the 1960s have possibly been caused by the intensification of agricultural practices.^[21] The Bern Convention of Europe list the hare under Appendix III.^[22] Several countries have placed L. europaeus on their Red List as "near threatened" or "threatened".^[23] The hare is considered a pest in some places, such as Argentina, Australia and North America. It causes damage to agriculture, particularly apple orchards.^[14] European hares are also hunted as game animals and their meat is considered white and delicious.^[14] Additional threats to the hare are the diseases European brown hare syndrome, pasteurellosis, yersiniosis (pseudo-tuberculosis), coccidiosis and tularaemia, which are principal sources of mortality.^[24]

In culture

In modern paganism, the hare (or rabbit) is associated with the spring goddess Eostre. In Germany, the Netherlands, Belgium and some other European mainland countries, it still is the Easter hare rather than the Easter Bunny. The phase "mad as a March hare" was derived from observations of the hare’s breeding behavior. The hare is a character of 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 tea party with the March Hare and the Mad Hatter.

References