European hare

European hare[1]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Lagomorpha
Family: Leporidae
Genus: Lepus
Species: L. europaeus
Binomial name
Lepus europaeus
Pallas, 1778
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

A European hare (top) compared with a mountain hare

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.[3] These subspecies have been distinguished by differences in pelage colouration, body size, external body measurements, and skull and tooth shape.[4] 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.[5][6]

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.[7] However, a 2006 study of the mtDNA of these same animals concluded they had diverged enough to be considered separate species.[8] 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.[9] 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.[9] Another 2008 study suggests that more research is needed before a conclusion is reached as to whether there is a species complex.[10] As of 2008, the European hare remains classified as a single species until further data shows otherwise.[2]

There appears to be genetic diversity in the European hare in the North Rhine-Westphalia region of Germany.[11] However, it is possible that restricted gene flow could change this within populations that become isolated.[11] Based on molecular phylogenetic studies, the Cantabric population in Spain has unique mtDNA in relation to other European populations.[12]

Description

Portrait of a European hare

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).[13] 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,[14] although the sides of the head and base of the ears do develop white areas.[3] 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.[15]

Range and habitat

Hare running in open field

European hares are native to much of continental Europe. Their range extends from northern Spain to southern Scandinavia, from Ireland to eastern Europe and northern parts of the Middle East and Central Asia. They have been extending their range into Siberia.[3] They may have been introduced to Britain by the Romans.[16] 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), Brazil, Chile, Argentina, Uruguay, Paraguay, Bolivia, Peru, the Falkland Islands, Australia, both islands of New Zealand and the south Pacific coast of Russia.[3][14] [17]

Hares primarily live in open fields with scattered brush for shelter. They are very adaptable and thrive in mixed farmland.[3] 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".[18]

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.[19]

Behaviour and life history

European hare displaying typical hiding behaviour.

Hares are primarily nocturnal and spend a third of their time foraging.[3] During daytime, a hare will hide 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.[14] Hares may be preyed on by canids, felids and birds of prey.[20] 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 will thump its front feet; the hind feet are used to warn others of a predator. A hare will squeal when hurt or scared and a female will make "guttural" calls to attract her young.[14]

Food and foraging

Hare pellets

European hares are primarily herbivorous. During the summer, they eat grasses, herbs and field crops.[14] Their preference is for wild grasses and weeds but with the intensification of agriculture, they have taken to feeding on crops.[2] During the winter, they eat herbage, twigs, buds and the bark of shrubs and young fruit trees.[14] They have been known to eat their own green, pellet feces to recover proteins and vitamins.[20] Two to three adult hares can eat more food than a single sheep.[14]

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.[21]

Mating and reproduction

Hares during "March madness"
Young European hare

European hares have a prolonged breeding season which lasts from January to August.[22][23] Sexual maturity occurs at seven or eight months for females and six months for males.[2] 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.[23]

Females have six-weekly reproductive cycles and are receptive for only a few hours in one day. Thus competition among local bucks is intense.[22] This phenomenon is known as "March madness"[23] 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 will fight off her numerous suitors if she is not ready to mate. Fights can be vicious and can leave numerous scars on the ears.[22] Hares will 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.[14] When a doe is ready to mate, she will run across the countryside, starting a chase that will test the fitness of the following males. When only the most dominant male remains, the female will stop and allow copulation.[22]

Young hare hiding

Female fertility continues through May, June and July, however testosterone production decreases in males and sexual behaviour becomes less overt.[23] 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.[23] 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.[2] The leverets (young hares) are fully furred and are precocial, being ready to leave the nest as soon as they are born,[14] an adaption to the lack of physical protection relative to that afforded by a burrow. A mother will visit the nest to nurse the leverets for five minutes a day. Young can eat solid food after two weeks and are weaned when they are four weeks old.[14] Hares can live for as long as twelve years.[2]

Status and human interactions

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

The European hare is listed as being of Least Concern by the IUCN[2] as it is considered to be wide-ranging and moderately abundant throughout its geographic distribution.[11] However, population declines have occurred since the 1960s and have possibly been caused by the intensification of agricultural practices.[24] In low population densities, hare are vulnerable to local extinctions.[2] The Bern Convention lists the hare under Appendix III as a protected species.[19] Several countries have placed the species on their Red List as "near threatened" or "threatened".[25] The hare is considered a pest in some areas and is known to damage crops.[14] They are also hunted as game animals.[3] 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.[26]

In culture

In Anglo-Saxon paganism, the hare is associated with reproduction and fertility and is a symbol for the spring goddess Eostre. Its connection with Easter eggs was based on a misconception by the Europeans that lapwings laid their eggs in the homes of hares. Germanic cultures noticed the high activity of hares during the spring and it was believed that their "mating dance" helped the earth grow.[27] This observation would also lead to the popular English idiom mad as a March hare.[28] The hare is a character in some fables, such as The Tortoise and the Hare of Aesop.[29] 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.[30]

References

  1. ↑ Hoffman, R.S.; Smith, A.T. (2005). "Order Lagomorpha". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 198–199. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. 1 2 3 4 5 6 7 8 Smith, A. T.; Johnston, C. H. (2008). "Lepus europaeus". IUCN Red List of Threatened Species. Version 2011.1. International Union for Conservation of Nature. Retrieved 2011-08-18.
  3. 1 2 3 4 5 6 7 Chapman, J. A.; Flux, J. E. C. (1991). Rabbits, Hares and Pikas: Status Survey and Conservation Action Plan. IUCN. pp. 62, 76–78. ISBN 2831700191.
  4. ↑ Suchentrunk, F.; Mamuris, Z.; Sfougaris, A. I.; Stamatis, C. (2003). "Biochemical genetic variability in brown hares (Lepus europaeus) from Greece". Biochemical Genetics 41 (5–6): 127–40. doi:10.1023/A:1023354709392. PMID 12834043.
  5. ↑ Palacios, F. (1989). "Biometric and morphologic features of the species of the genus Lepus in Spain". Mammalia 53 (2): 227–64. doi:10.1515/mamm.1989.53.2.227.
  6. ↑ Riga, F.; Trocchi, V.; Randi, E.; Toso, S. (2001). "Morphometric differentiation between the Italian hare (Lepus corsicanus De Winton, 1898) and the European brown hare (Lepus europaeus Pallas, 1778)". Journal of Zoology 253 (2): 241–52. doi:10.1017/S0952836901000218.
  7. ↑ Ben Slimen, H.; Suchentrunk, F.; Memmi, A.; Ben Ammar Elgaaied, A. (2005). "Biochemical genetic relationships among Tunisian hares (Lepus sp.), South African Cape hares (L. capensis), and European brown hares (L. europaeus)". Biochemical Genetics 43 (11–12): 577–596. doi:10.1007/s10528-005-9115-6. PMID 16382363.
  8. ↑ Ben Slimen, H.; Suchentrunk, F.; Memmi, A.; Sert, H.; Kryger, U.; Alves, P. C.; Elgaaied, A. B. A. (2006). "Evolutionary relationships among hares from North Africa (Lepus sp. or Lepus spp.), Cape Hares (L. capensis) from South Africa, and brown hares (L. europaeus), as inferred from mtDNA PCR-RFLP and allozyme data". Journal of Zoological Systematics and Evolutionary Research 44 (1): 88–99. doi:10.1111/j.1439-0469.2005.00345.x.
  9. 1 2 Ben Slimen, H.; Suchentrunk, F.; Ben Ammar Elgaaied, A. (2008). "On shortcomings of using mtDNA sequence divergence for the systematics of hares (genus Lepus): An example from Cape hares". Mammalian Biology 73 (1): 25–32. doi:10.1016/j.mambio.2007.02.003.
  10. ↑ Ben Slimen, H.; Suchentrunk, F.; Stamatis, C.; Mamuris, Z.; Sert, H.; Alves, P. C.; Kryger, U.; Shahin, A. B.; Ben Ammar Elgaaied, A. (2008). "Population genetics of Cape and brown hares (Lepus capensis and L. europaeus): A test of Petter's hypothesis of conspecificity". Biochemical Systematics and Ecology 36 (1): 22–39. doi:10.1016/j.bse.2007.06.014.
  11. 1 2 3 Fickel, J.; Schmidt, A.; Putze, M.; Spittler, H.; Ludwig, A.; Streich, W. J.; Pitra, C. (2005). McCorquodale, ed. "Genetic structure of populations of European brown hare: Implications for management". Journal of Wildlife Management 69 (2): 760–770. doi:10.2193/0022-541X(2005)069[0760:GSOPOE]2.0.CO;2.
  12. ↑ Palacios, F.; Estonba, A.; Perez-Suarez, G.; Alonso-Campos, G.; Sanz, A.; Galan, L. (2004). Report on the restoration program of the Cantabrian population of brown hare (Lepus europaeus Pallas, 1778) in the Basque Country, Spain. Vairao, Portugal: Research Center in Biodiversity and Genetic Resources.
  13. ↑ Burnie, D.; Wilson, D. E., ed. (2005). Animal: The Definitive Visual Guide to the World's Wildlife. DK Adult. p. 143. ISBN 0789477645.
  14. 1 2 3 4 5 6 7 8 9 10 11 Naughton, D. (2012). The Natural History of Canadian Mammals. University of Toronto Press. pp. 235–38. ISBN 1442644834.
  15. ↑ Schnurr, D. L.; Thomas, V. G. (1984). "Histochemical properties of locomotory muscles of European hares and cottontail rabbits". Canadian Journal of Zoology 62 (11): 2157–2163. doi:10.1139/z84-313.
  16. ↑ Hutchings, M.R.; Harris, S (1996). "The current status of the brown hare (Lepus europaeus) in Britain" (PDF). Joint Nature Conservation Committee. Retrieved 30 May 2015.
  17. ↑ http://www.seb-ecologia.org.br/2009/resumos_ixceb/1789.pdf
  18. ↑ Pikula, J.; Beklová, M.; Holešovská, Z.; Treml, F. (2004). "Ecology of European brown hare and distribution of natural foci of Tularaemia in the Czech Republic" (pdf). Acta Veterinaria Brno 73 (2): 267–273. doi:10.2754/avb200473020267.
  19. 1 2 Vaughan, N.; Lucas, E.; Harris, S.; White, P. C. L. (2003). "Habitat associations of European hares Lepus europaeus in England and Wales: Implications for farmland management". Journal of Applied Ecology 40 (1): 163–175. doi:10.1046/j.1365-2664.2003.00784.x. JSTOR 827268.
  20. 1 2 "Lepus europaeus European hare". Animal Diversity. University of Michigan.
  21. ↑ Monaghan. P.; Metcalfe, N. B. (1985). "Group foraging in wild brown hares: Effects of resource distribution and social status". Animal Behaviour 33 (3): 993–999. doi:10.1016/S0003-3472(85)80033-6.
  22. 1 2 3 4 Holly, T. (2001). "Mad World of the European Hare". In MacDonald, D. The New Encyclopedia of Mammals. Oxford: Oxford University Press. pp. 710–711. ISBN 0198508239.
  23. 1 2 3 4 5 Lincoln, G. (1974). "Reproduction and March madness in the brown hare, Lepus europaeus". Journal of Zoology 174 (1): 1–14. doi:10.1111/j.1469-7998.1974.tb03140.x. PMID 4468894.
  24. ↑ Smith, R. K.; Jennings, N. V.; Harris, S. (2005). "A quantitative analysis of the abundance and demography of European hares Lepus europaeus in relation to habitat type, intensity of agriculture and climate". Mammal Review 35 (1): 1–24. doi:10.1111/j.1365-2907.2005.00057.x.
  25. ↑ Reichlin, T.; Klansek, E.; Hackländer, K. (2006). "Diet selection by hares (Lepus europaeus) in arable land and its implications for habitat management". European Journal of Wildlife Research 52 (2): 109–118. doi:10.1007/s10344-005-0013-3.
  26. ↑ Lamarque, F.; Barrat, J.; Moutou, F. (1996). "Principal diagnoses for determining causes of mortality in the European brown hare (Lepus europaeus) found dead in France between 1986 and 1994". Gibier Faune Sauvage 13 (1): 53–72.
  27. ↑ Skye, M. (2007). Goddess Alive!: Inviting Celtic & Norse Goddesses Into Your Life. Llewellyn Worldwide. p. 59. ISBN 0738710806.
  28. ↑ Carnwell, S. (2010). Hare. Reaktion Books. p. 181. ISBN 9781861894311.
  29. ↑ Aesop (2012). Aesop's Fables. Simon & Brown. ISBN 1613823584.
  30. ↑ Carroll, Lewis (1993). Alice's Adventures in Wonderland. Dover Publications. ISBN 0486275434.

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