Lepus europaeus Pallas, 1778

Dataset
GBIF Backbone Taxonomy: Rank
SPECIES: Published in
Nova Spec. Quad. Glir. Ord. p.30

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Classification

kingdom
Animalia: phylum
Chordata: class
Mammalia: order
Lagomorpha: family
Leporidae: genus
Lepus: species
Lepus europaeus

native range

Africa

native range

Asia-Temperate

native range

China

native range

Europe

eunis habitat

Grassland and tall forb habitats (E level 1)

eunis habitat

Heathland, scrub and tundra habitats (F level 1)

native range

Northern Africa

ecofunctional group

Phytophagous

eunis habitat

Regularly or recently cultivated agricultural, horticultural and domestic habitats (I level 1)

biotope

agriculture

cuddliness

cuddly

nutrient level

eutrophic

lifespan

longlived

mobility

mobile

biome

terrestrial

spine

vertebrate

biology_ecology

BEHAVIOR Grouping behavior. — Lepus europaeus has long been supposed to be a solitary animal except during the breeding season (Flux 1990) but actually keeps in contact with neighbors even in areas with low densities, and in pairs or groups of two or three at higher densities (Schneider 1979; Zörner 1981; A. Bock, in litt.). If food is unevenly spaced it benefits from higher group sizes allocating more time to feeding with increased corporate vigilance. However, when food is clumped in small patches dominant L. europaeus attempt to monopolize the resource. When breeding, it gathers in groups of up to 13 (Lincoln 1974; Zörner 1981; Monaghan and Metcalfe 1985; Flux 2009; Bock 2013). Grouping is dynamic and hierarchies do not exist as in herds with leaders. There is a high tolerance among conspecifics and only under harsh conditions does it rebuff other approaching L. europaeus. In the breeding season, males fight for females; however, serious fights are rare. Rather, it shows a series of displays; “ showing off, ” humility gestures, and conciliation gestures. Between mother and litters social contact is restricted to suckling once a day and possibly also support in dangerous situations, when females approach their squealing young (Schneider 1979; Zörner 1981; Krapp 2003). Reproductive behavior. — The breeding season for Lepus europaeus starts around mid-December in the northern hemisphere and in June in the southern hemisphere. Sexual hormone production (e. g., testosterone) increases, leading to increased activity during daylight hours (Flux 1965; I have observed this daylight activity during two field seasons, June 2015 and 2019). L. europaeus gathers in mixed-sex groups, some may leave the group, others arrive. Courtship includes several changing sequences. Males seek to win a female by chasing; it makes conciliation sounds, approaching the female, trying cautious contact, chasing away or fighting with other males. Boxing between males establishes rank and the right for a female. Fights can be distinguished as “ distance fights ” and “ breast fights. ” In both cases L. europaeus stands on hind legs. In distance fights however, they stretch the back and touch only each other’s front paws; in breast fights front paws touch also each other’s face. If males come too close, females show defense either in turning around and sitting in a threatening position or by directly rebuffing the male. Active phases are often interrupted by resting phases with grooming and feeding. Mating games in the group continue until male – female pairs separate from the group. If the female does not respond to the males’ courtship the pair separates; however, if the female does respond chasing is prolonged. Courtship can continue several days, body contacts increase, finally leading to more intensive contacts where the male touches repeatedly the belly of the female thus triggering ovulation. Eventually, the male jumps on the back of the female for mating several times in succession (Krapp 2003). Either the couple separates or the procedure starts anew. Often, there is a so-called satellite-male nearby that observes the pair over days, approaches, and tries to mate with the female (Schneider 1979; Flux 1981; Krapp 2003). However, I have observed that if a male wins a female he defends her by chasing away other competitors coming closer than 3 – 4 m. Communication. — Vocalizations play a minor role in Lepus europaeus communication. There is a loud squeal when injured or threatened. It growls while chasing and mating (like rabbits). L. europaeus grinds its teeth which is considered to be a threat or fear sound. However, rabbits grind when lying peacefully and relaxed, so this might be the same in L. europaeus. Like the European rabbit, L. europaeus uses a special way of urinating. First, there is so-called mark urination, where the back and tail is lifted during urination; second, it sprays urine deliberately toward conspecifics (Zörner 1981; Krapp 2003). Miscellaneous behavior. — Lepus europaeus is mainly nocturnal or crepuscular. However, physiology does not indicate nocturnal activity based on its visual anatomy and cornea size. L. europaeus is actually classified as cathemeral (Kirk 2006). This adaptation allows them to avoid aerial predators by feeding at night, and run fast to escape predators hunting by day. In areas with no disturbance L. europaeus is also active by day. Furthermore, patterns shift with day length and breeding activity (Krapp 2003). Lepus europaeus does not hibernate, but in snow it builds hollows for shelter. On days with temperatures over 25 ° C it seeks shade and postpones activity to nightfall (Zörner 1981; Holley 2001; Schai-Braun et al. 2012). Nevertheless, resting takes most of the time in the 24 - h cycle, even active phases are interrupted by spontaneous resting phases. If active, it spends the time foraging, avoiding predators, grooming, and mating. During the breeding season, lactating females may spend 11 h feeding (Flux 1981). Mutual grooming could be observed in captivity, thus can be expected in the wild, too (Zörner 1981).

biology_ecology

Reproduction. — The start of sexual activity in the breeding season is the result of a complex system involving photoperiod and resulting production of hormones in the hypothalamus and pituitary gland, climatic conditions, and individual nutritional status (Nelson et al. 1990; Krapp 2003). Lepus europaeus is a polyestrous seasonal breeder with a reproduction period of about 215 – 256 days. Generally, breeding season starts in midwinter (December – February) in males followed by females about a month later (Zörner 1981; Caillol et al. 1986). In New Zealand and other places south of the equator, breeding season starts soon after the shortest day of year in June and in New Zealand over 90 % of adult females are pregnant from August to February (Flux 1990). In males from the northern hemisphere, inhibition of spermatogenesis starts in September and lasts to mid-December; in October and November gonads are entirely inactive (April – May in New Zealand — Flux 1990). Testes are relocated into the inguinal region, and their weight reaches a minimum for the year. Production of sexual hormones testosterone and luteinizing is low. At the end of December testes relocate to the scrotum, becoming visible. Their weight increases rapidly and spermatogenesis probably starts at a weight of 5 g (Zörner 1981). In females, the cycle of gonad development starts in January, about a month later than in males; lowest weight of ovaries ranges from November to December. As L. europaeus is a strictly induced ovulator, stimulation by copulation is considered a necessity. These cycles depend on climatic conditions, and in a warm winter the hormonal resting period might be shorter (Roellig et al. 2010). Considered to be an r-selected species with early maturity and relatively short life span, reproduction within the breeding season can be increased if a new conception overlaps an ongoing pregnancy (superfetation). Thus, the normally 42 - to 43 - day gestation period can be reduced to only 38 days. However, this can be seen mainly in captive hares (Flux 1990). Ultrasonography revealed a new set of corpora lutea near the active corpora lutea of the current pregnancy up to 2 days before delivery. Prepartum conception shortens the reproductive cycle, increasing the average number of litters within the breeding season, thus enhancing reproductive success (Roellig et al. 2010). A female gives birth to on average eight leverets per year. If conditions are good, 11 leverets are possible; unfavorable conditions can lead to a remarkable decrease in offspring due to embryonic resorption before mid-term (embryo about 25 mm in length — Roellig et al. 2010). Maximum reproductive performance was 12 in Denmark (Hansen 1992 [not seen, cited in Krapp 2003]) and 20 in the South of France (Bray 1998). Females that do not take part in reproduction show changes in the uterus that lead to sterility more frequently in old females (Krapp 2003). In the northern hemisphere young females born at the beginning of the breeding season normally do not reproduce in the same year; however, young females in Argentina and New Zealand may breed in their year of birth. This can be explained by the food which is available and of good quality (high protein content) year-round (Dietrich 1985). Number of litters during one breeding season averages 3 (range 1 – 4), and average litter size is 3 (range 1 – 5). Prenatal mortality determines the litter size; the highest death rate is in spring and autumn (23.4 – 58 % and 30 – 55.6 %, respectively), the lowest in summer (12.8 – 28 % — Zörner 1981; Krapp 2003). Observations of parturition in free-ranging L. europaeus are rare. If not disturbed the female gives birth to the leverets at one site. If disturbed, she might split places for parturition. However, dispersal of the litter is induced by the leverets themselves after about 3 days, probably a precaution against predation on the whole litter (Zörner 1981). ECOLOGY Population characteristics. — Estimates of population density depend on the method of data acquisition and the time of year (see “ Miscellaneous ”). According to Pikula et al. (2004) the highest mean densities can be reached in warm and dry or moderately dry districts with mild winters and shorter duration of sunshine. Densities of Lepus europaeus vary with availability of food and shelter and number of predators. Optimal conditions can lead to densities up to 100 individuals / km 2 (Klansek 1996) but also densities of 1 individual / km 2 are viable. High densities in Austria can exceed 100 individuals / km 2; but in Switzerland (adjacent to Austria) densities reach only 6 – 19 individuals / km 2 (Pfister et al. 2002). On islands with almost no predation pressure, for example, Illumø in Denmark, density reached 339 individuals / 100 ha without hunting (Abildgård et al. 1972). Nevertheless, even with hunting, density is higher on predator-free islands than in areas where predators exist (e. g., island Ven, South Sweden — Frylestam 1979). In Germany densities fluctuated from 5 to 26 individuals / km 2 averaging 11 individuals / km 2 in 2013 (Wildtier-Informationssystem der Länder Deutschlands [WILD], https: // www. jagdverband. de / forschung-aufklaerung / wild-monitoring). In Argentina L. europaeus reached densities of 100 individuals / km 2 (Dietrich 1985) compared to only 20 – 30 individuals / km 2 in New Zealand (Flux 1990). Densities vary depending on landscape, climate, predation, and diseases. Survival and mortality are influenced by a complex of interacting factors including climate and geographic area, diseases, predators, and anthropogenic factors like agriculture, hunting, or traffic. Generally, mortality is high in leverets; about 75 % of them die within the first 8 months. Average adult mortality is about 38 % (Broekhuizen and Maaskamp 1979; Gehle 2002). Despite high leveret mortality young L. europaeus dominate the population with about 55 %, varying from 30 % to 70 % depending on conditions. L. europaeus that are 1 – 3 years of age form about 30 %, of the population, and those 3 years and older about 14 % (Zörner 1981; Krapp 2003). Lepus europaeus can reach an age of 8 – 13 years. A markrecapture analysis revealed that females outnumber males when older; thus, females have a higher life expectancy than males. Generally, L. europaeus that dominate and effectively contribute to the population are 8 months to 5 years of age. The sex ratio can vary with annual climatic and ecological conditions, and changes in population dynamics. However, overall it is close to 1: 1 (Zörner 1981; Dietrich 1985; Flux 1990; Krapp 2003). Space use. — Originally, Lepus europaeus inhabited temperate savanna or grassland. After forest clearance and reclamation of wetlands for cultivation and agriculture L. europaeus followed to take advantage of the food supply. In this situation, L. europaeus prefers open fields and pastures bordered by hedgerows and woodlots (Fig. 5). Investigation of movements of radiotracked L. europaeus in England and Germany found a preference for low-growing crops, different areas used for feeding during night and sheltering during day, preference for woodlands and hedges and shifting activity according to crop development (Tapper and Barnes 1986; Rühe and Hohmann 2004). L. europaeus favors high diversity of plants with different crops, vegetables, and high-energy weeds like clover and corn poppy to cover physiological and reproductive needs (Jennings et al. 2006; Reichlin et al. 2006). Studies by Tapper and Barnes (1986) and Rühe and Hohmann (2004) found that night ranges were larger than day ranges and ranges increased as the number of habitats increased. The home range of L. europaeus is about 21 – 190 ha (Rühe and Hohmann 2004; Flux 2009). Males have larger home ranges than females but once a home range is established L. europaeus shows high site fidelity (Bray et al. 2007). In Australia, L. europaeus home ranges averaged 149 ha, compared to 10 ha for the European rabbit. L. europaeus can show extraordinary homing and sense of direction. In two relocation projects in Poland two of 500 and two of 300 L. europaeus returned “ back home ” over a distance of 229 km and 464 km, respectively (Zörner 1981). L. europaeus has great mobility that enables it to travel 20 km in a night and hundreds of kilometers in heavy snow or drought to look for food (Angermann et al. 1990). Also, if densities are high, it moves to find other suitable habitats, even up to elevations of 2,000 m (Flux 1990). Lepus europaeus lives aboveground year-round, exposed to all climatic conditions. Even when giving birth, females do not build nests but choose dry and sheltered places. It scratches shallow “ forms ” in places that meet its requirements, for example, exposed to sunlight (in winter), dry elevations in wetlands or in places that provide enough cover. The size of the form equals the size of the resting L. europaeus (Zörner 1981). Diet. — Lepus europaeus is exclusively herbivorous and generally the diet depends on what is available. The diet consists of a variety of sweet grass (Poaceae), weeds (Asteraceae, Brassicaceae, Fabaceae, Plantaginaceae), crops like rye (Secale cereale), wheat (Triticum vulgare), barley (Hordeum), and oat (Avena sativa); however, weeds and wild grasses are selected over crops when available (Reichlin 2006). A study of L. europaeus in southern Sweden revealed that rape, widely represented in agriculture, was less eaten in mixed farmland. It was only selected in rape monocultures; however, L. europaeus avoided it in late autumn possibly due to its content of glucosinulates (Frylestam 1986). The autumn diet in Hungarian L. europaeus included 24 species, among the cultivated species mainly wheat, Triticum aestivum, but grasses and shrubs (and a high consumption of elderberry, Sambucus) were also important food components in every area (Katona et al. 2010). Generally, food intake shifts to bark, young branches, and buds when weeds and crops are scarce in winter; thus, L. europaeus can damage reforestation or fruit-growing areas. In winter, L. europaeus generally prefers crops over weeds and also feeds on sugar beet and carrots if provided by hunters (Zörner 1981; Reichlin et al. 2006). In late autumn and winter, foraging takes place at night, dusk, or dawn; with increasing day length and vegetation growth, and the onset of breeding season, foraging extends to daytime. Food in other parts of the world, for example, alpine grassland in New Zealand, may comprise species never previously encountered such as Aristotelia fruticosa, Celmisia allanii, Celmisia coriacea, Chionochloa rubra, C. pallens, C. flavescens, Dracophyllum uniflorum, Hymenanthera alpina, Poa colensoi, and Schoenus pauciflorus (Flux 1990). In captivity L. europaeus drinks water on a regular basis; the amount of food (hay, hare pellets) intake is approximately 500 g. If temperature decreases food intake increases (Zörner 1981).

diagnosis

DIAGNOSIS Lepus europaeus (Fig. 1) and the sympatric Lepus timidus (mountain hare) in the northern hemisphere are very similar in size. Although some arctic subspecies of L. timidus may be as large as, or even larger, than the largest L. europaeus, L. europaeus is generally heavier at 3.5 – 5.0 kg (L. timidus averages from 2.4 kg in the Alps to 3.4 kg in Siberia — Angerbjorn 2018). Headbody length of L. europaeus is 550 – 650 mm (460 – 560 mm in L. timidus). The ears are longer from notch 110 – 140 mm and when bent forward they are longer than forehead (versus 67 – 106 mm in L. timidus, ears when bent forward are equal in length to forehead); the tail is also longer at 75 – 140 mm (mean 95 mm) compared with 53 – 70 mm in L. timidus; length of hind foot is 130 – 164 mm (mean 147 mm) which is shorter than in L. timidus (145 – 172 mm — Krapp 2003; Angerbjorn 2018). Fig. 1. — Adult Lepus europaeus, gender unknown, industrial area near Hamburg, Germany, July 2017. Photograph by A. Bock. The length of the skull (from the most rostral to the most caudal point of the occipital) averages 97.1 mm in L. europaeus (L. timidus averages 90 mm). The facial region of the skull is extended with markedly raised nasalia (Fig. 2) compared to a shorter facial and a more concave frontal region in L. timidus. Dentale is softer, processus condyloideus is inclined more posteriorly than in L. timidus. Diastema relatively long (average 31.2 mm), and at over 80 % of lower toothrow, longer than in L. timidus, corpus mandibulae with incisivus (I) inclined anteroventrally (anterodorsal in L. timidus). Incisivus 1 (I 1) of L. europaeus are rectangular in section; ratio of length to width 0.74: 0.80 compared to the more quadratic dimension in L. timidus of 0.84: 0.95; incisors are additionally more curved than in L. timidus. Root of I 1 far anterior of sutura premaxillo-maxillaris with a frequency of 78.3 – 100 % compared to 98.6 – 100 % in L. timidus. Zygomatic length (34.7 – 41.8 mm) and zygomatic width (44 – 49 mm) are longer than in L. timidus (length = 33.4 – 37.9; width = 42.5 – 45.6 mm). Front and rear wings of supraorbital appendix are well developed and mostly free (not as well developed in L. timidus), osseous palate shorter (compared to L. timidus), choanae wide. Coat color of L. europaeus does not change to white in winter as in L. timidus, although animals in the northern latitudes may turn slightly lighter. Teat formula 2 p + 4 a = 6 compared to 2 p + 6 a = 8 for L. timidus (Krapp 2003). Fig. 2. — Dorsal, ventral, and lateral views of skull and lateral view of mandible of an adult male of Lepus europaeus, collection locality Schwarzenbek, Germany, 02.12.1997, specimen number 8933 (468), Zoological Museum, University of Hamburg, greatest length of skull 95 mm. Photograph by A. Bock. Although similar in appearance to L. europaeus, Lepus capensis (Cape hare) is overall smaller and lighter (with ear length 97 – 114 mm; skull length 27.5 – 30 mm; length of hind foot 93 – 103 mm, weight 1.5 – 2.5 kg). When compared to the sympatric Oryctolagus cuniculus (European rabbit) the difference is obvious. The European rabbit is smaller (body weight 0.9 – 1.8 kg, head – body length 375 [female] and 406 [male] mm), and has a more compact shape. The European rabbit burrows and its leverets are born nidicolous (altricial) compared to the L. europaeus which lives aboveground with nidifugous (precocial) leverets. Ears are shorter (when bent forward they do not extend to the nose) and have no black tips. Furthermore, the eyes do not have a light-brown iris but look black and the pelage has more grayish than yellowish base hair and longer guard hair with black ends (Zörner 1981; Krapp 2003).

Name

Synonyms: Leppus europaeus; Lepus capensis europaeus; Lepus cyanotis Blanchard, 1957; Lepus europaeus ghigii de Beaux, 1927; Lepus europaeus mediterraneus Wagner, 1841; Lepus hispanicus; Lepus hybridus; Lepus karpathorum; Lepus meridionalis; Lepus occidentalis; Lepus parnassius; Lepus pyrenaicus; Lepus tesquorum; Lepus transsylvanicus
Homonyms: Lepus europaeus Pallas, 1778
Common names: Brown Hare in English; Brown Hare in English; Brown hare in English; Brown hare in language.; Common Hare in English; Common Hare in English; European Hare in English; European Hare in English; European Hare in English; European Hare in English; European Hare in English; European brown hare in language.; European hare in language.; European hare in English; European hare in language.; European hare in language.; Europski zec in Croatian; Feldhase in German; Feldhase in German; Feldhase in German; Fälthare in Swedish; Gráhéri in Icelandic; Haas in Dutch; Halljänes in Estonian; Hare in Danish; Hare in Danish; Hues in Luxembourgish; Iepure in Romanian; Iepurele-de-câmp in Romanian; Lebre-europeia in Portuguese; Lepre europea in Italian; Lepuri i eger in Albanian; Liebre europea in Spanish; Liebru in Maltese; Lièvre d'Europe in French; Mezei nyúl in Hungarian; Pilkasis kiškis in Lithuanian; Poljski zajec in Slovenian; Rusakko in Finnish; Sørhare in Norwegian; Zajíc polní in Czech; brown hare in language.; brown hare, european hare in English; haas in Dutch; liebre Europea in language.; lièvre in French; lièvre d'Europe in language.; Заєць сірий in Ukrainian; Brown Hare in English; European Hare in English

Bibliographic References

Hall, R. (2004) Däggdjur
Hall, R. (2016-05-02 00:00:00) Däggdjur
Smith, Andrew T., Charlotte H. Johnston, Paulo C., 2018: null. Lagomorphs. Pikas, Rabbits, and Hares of the World. 266.
Wilson, Don E., and DeeAnn M. Reeder, eds., 1992: null. Mammal Species of the World: A Taxonomic and Geographic Reference, 2nd ed., 3rd printing. xviii + 1207.
Wilson, Don E., and DeeAnn M. Reeder, eds., 2005: null. Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd ed., vols. 1 & 2. 2142.
Wilson, Don E., and F. Russell Cole, 2000: null. Common Names of Mammals of the World. xiv + 204.