Hares and rabbits

(Leporidae)

Class Mammalia

Order Lagomorpha

Family Leporidae

Thumbnail description
Small- to medium-sized mammals with long ears, short tails, and long hind legs

Size
10–30 in (25–75 cm); 14 oz–13.2 lb (400–6,000 g); females usually larger than males

Number of genera, species
11 genera; 61 species

Habitat
Forest, woodland, savanna, scrub, and tundra

Conservation status
Critically Endangered: 2; Endangered: 7; Vulnerable: 3; Data Deficient: 9

Distribution
Africa, Asia, Europe, North America, South America, and introduced to Australia, New Zealand, Java, and other islands

Evolution and systematics

The Leporidae is sometimes divided into subfamilies Paleolaginae (Pentalagus, Pronolagus, and Romerolagus) and Leporinae (remaining genera). There are 11 genera: 32 Lepus, true hares; 1 Oryctolagus, European rabbit; 17 Sylvilagus, cotton-tails; 1 Brachylagus, pygmy rabbit; 3 Pronolagus, red rock-hares; 1 Bunolagus, riverine rabbit; 1 Pentalagus, Amami rabbit; 1 Caprolagus, hispid hare; 1 Poelagus, Bunyoro rabbit; 2 Nesolagus, striped rabbits; and 1 Romerolagus, volcano rabbit.

Early leporids were found in both the Old and New World before Lepus evolved, but had a probable origin in Asia. The Leporidae arose in the Pliocene and Lepus in the early Middle Pleistocene. Whereas Pentalagus, Pronolagus, and Romerolagus are ancient forms, Lepus is a young expanding genus.

The monotypical genera in Leporidae are taxonomically distinct with the exception of Brachylagus, which sometimes is included in Sylvilagus. However, the Lepus genus is in a state of confusion regarding both phylogeny and number of species.

Physical characteristics

All species in the family Leporidae have basic brown or gray soft fur with relatively long legs, feet, and ears. The tail is short and bushy. Some northern forms turn white for winter, and two are striped. The family can be split into two groups, hares and rabbits, based on morphology and ecology. The hare group includes all hares and jackrabbits in the genus Lepus, hereafter called hares. The rabbit group consists of the remaining genera in the family, despite that they are sometimes also known as hares (for example, hispid hares) hereafter called rabbits. Hares have long legs and can reach considerable running speed. They also have longer ears, usually with black tips, and are, overall, larger than rabbits. The body length of hares is 15–30 in (40–75 cm), with a weight of 3–13.2 lb (1,350–6,000 g). Rabbits, on the other hand, have shorter legs and are generally smaller: 10–20 in (25–50 cm); 14 oz–6.6 lb (400–3,000 g).

Distribution

Altogether, hares and rabbits are distributed over most areas of the world: Africa, the Americas, Asia, and Europe, and have also been introduced into southern South America, Australia, New Zealand, Java, and other islands.

The natural habitat of European hares is open terrain ranging from forest steppes to arid steppes. They therefore benefit from cultivation of fields. In connection to agriculture, they spread rapidly to the east and northeast during the nineteenth and twentieth centuries. They were also successfully introduced in agricultural areas in Argentina, Ireland, Scandinavia, New Zealand, Australia, and southern Siberia.

The European rabbit has a similar increase in distribution due to many introductions in suitable habitats. Before the Ice Age, it inhabited parts of Western Europe, as indicated by fossils. However, its natural post-Ice Age distribution includes only Spain and northwest Africa. But already the Romans had introduced rabbits to different parts of Europe. There were several introductions in Australia, but the successful ones came from 24 rabbits in 1859 in Victoria and in 1864 to New Zealand. Both Australia and New Zealand suffered tremendously from a gigantic rabbit population explosion. In spite

of the disastrous consequences, wild rabbits were later introduced into Chile and became a plague there as well.

Habitat

Since hares and rabbits are found almost all over the world, they are also both found in a variety of habitats, from deep forests to open deserts. But hares have different basic habitat requirements than rabbits.

Hares are mostly found in open areas where their running speed is a good adaptation to escape predators. They can thus be found from arctic tundra to productive grasslands or deserts. In these open areas, they take advantage of shrubs and rocks to hide, and rely on their well-camouflaged fur. But snowshoe hares and partly mountain hares and Manchurian hares prefer coniferous or mixed forests.

However, rabbits are confined to forests and shrubs where they can hide in the vegetation or in burrows. Some rabbits are found in deep tropical forests, such as the striped rabbits, and others use specific habitats like riverine shrubs, as do the riverine rabbit and hispid hare. Cottontails inhabit a large variety of habitats, including swamps, forests, parks, agricultural field, shrubs, and deserts.

Behavior

Predation is an important aspect in the life of hares and rabbits. Hares escape predators by running away and try to confuse predators by backtracking themselves. Rabbits try to

escape predators by running into burrows or holes. Therefore, hares can travel considerable distances and have large home ranges, while rabbits mostly stay in the vicinity of safe hiding places in small home ranges or territories. Many species use distress calls or thump their hind feet on the ground to warn for predators. They also use a soft sound produced by grinding their teeth.

Hearing is the most important sense for leporids, but scent marking is also an central way to communicate. They have scent glands on their nose, chin, and around the anus. This is important in sexual situations, especially for the social European rabbit.

Feeding ecology and diet

All hares and rabbits are strictly herbivores. Their diets include green plant parts, especially grasses, clover, and, to a lesser extent, cruciferous and composite plants (e.g., dandelions). In winter, the diet also includes dry twigs, buds, young tree bark, roots, and seeds. In steppe regions, the winter diet consists of dry weeds and seeds. Most like cultivated plants such as winter grains, rapeseed, cabbage, parsley, and carnations. Both hares and rabbits may damage cereals, brassica crops, fruit trees, and tree plantations, especially in winter. Hares rarely drink when there are enough moist plants around, but sometimes eat snow in the winter.

Reproductive biology

Most hares and rabbits are solitary, although they can be seen in groups around mating season when males fight each other to build social hierarchies and try to get access to females in estrous. But the European rabbit forms highly social groups with one to four males and one to nine females. They can dig complex burrows that they defend against other social groups. Both males and females are promiscuous. Like many leporids, they mate again soon after giving birth. Since

gestation length for the European rabbit is about 30 days, they give birth to five to seven litters a year. European hares increase reproductive output by fertilizing a second litter before the first one is born (superfetation).

Hares give birth, after a long gestation period, to well-developed fur-covered leverets that have open eyes and that can already move around at birth. Rabbits have a shorter gestation period and build nests with fur in burrows or hollows underground or in dense cover. The newborn rabbit kittens are poorly developed with eyes closed and no fur. After birth,

both hare and rabbit mothers suckle their leverets only once a day with highly nutritious milk. This lack of social contact makes it difficult for predators to find juvenile hares and rabbits. Litter size of both hares and rabbits vary with geography and climate.

Conservation status

Typically, the ancient rabbit species are threatened with extinction because of low population sizes and restricted distributions, including the riverine rabbit, volcano rabbit, hispid hare, and the two striped rabbits. Some species on islands are also Endangered, such as the Hainan hare, Tres Marias cottontail, and Amami rabbit.

Many species have not been evaluated by IUCN and some of those will undoubtedly be listed as threatened such as the newly described Annamite striped rabbit. Another problem is the taxonomy where relationships between taxa are not well described.

Significance to humans

Many hares and rabbits are economically important for humans. They are hunted for meat, skin, and for sport. Cottontails and some of the hares are hunted in millions every year. However, they may also damage agricultural crops and forestry plantations. When introduced into new areas, they can also threaten native species.

Hares and rabbits are known from tales dating back centuries, mostly as tricksters and fertility symbols. The hare was, for example, important in the Germanic heathen mythology as the fertility goddess Ostara, and was transformed by Christians into the Easter celebration.

The domestic European rabbit is well known all over the world and used extensively in laboratory research and as pet animals. The first descriptions of domestic breeds of European rabbits come from France in the sixteenth century.

Species accounts

List of Species

Snowshoe hare

Lepus americanus

taxonomy

Lepus americanus Erxleben, 1777, Fort Severn, Ontario, Canada. Fifteen subspecies.

other common names

French: Lièvre d'Amérique; German: Schneeschuhhase; Spanish: Liebre nival.

physical characteristics

The smallest of the hares with large hind feet that turn white during the winter.

distribution

From Alaska to Newfoundland and south through the coastal range; the Rockies and the Appalachians to northern California, northern New Mexico, and Tennessee.

habitat

From conifer forests to mixed forests, with a preference for transition zones.

behavior

Populations show large synchronized fluctuations with a peak every eight to 11 years (10-year cycles). They use forms for cover during the day and make regular trails to feeding areas.

feeding ecology and diet

Summer diet is grasses, sedges, and various herbs, but changes to birch, aspen, willow, spruce, and pine during the winter.

reproductive biology

Promiscuous. Most reproductive characters are related to the 10-year population cycle. Normally gives birth to two litters per year, but this can increase to four, dependent on geography and population phase. Litter sizes vary from one to 10.

conservation status

Common; not threatened.

significance to humans

Important game species; may damage tree plantations.

European hare

Lepus europaeus

taxonomy

Lepus europaeus Pallas, 1778, southwest Poland. Thirty sub-species.

other common names

English: Brown hare; French: Lièvre brun; German: Feldhase; Spanish: Liebre común.

physical characteristics

Body length 20–27 in (52–68 cm), tail 3–5 in (8–12 cm), body weight 5.5–14 lb (2,500–6,000 g). A large hare that is brown throughout the year with long ears, relatively long tail, and long limbs.

distribution

Most of Europe, south to Iran, and into western Siberia. Introduced in eastern North America, southern South America, southeastern Australia, New Zealand, and several islands.

habitat

Open country as mixed farmland, moorland, steppe, but also open woodland. In introduced areas habitats can be variable, including pampas, sand dunes, marshes, and alpine grassland.

behavior

Solitary, but may aggregate in groups when feeding; males fight around females in estrous.

feeding ecology and diet

Diet varies with mostly grass in the summer and can include shrubs in the winter, but on agricultural land crops dominates.

reproductive biology

Males and females are promiscuous. Females give birth to two to four litters per year with a mean litter size of one to four.

conservation status

Common; not threatened.

significance to humans

Important game species that may damage crops.

Mountain hare

Lepus timidus

taxonomy

Lepus timidus Linnaeus, 1758, Uppsala, Sweden.

other common names

English: Blue hare, varying hare; French: Lièvre variable; German: Schneehase; Spanish: Liebre variable.

physical characteristics

Body length 18–24 in (46–61 cm); weight 4.4–10.4 lb (2,000–4,700 g). A medium-sized hare with long hind feet and short ears; summer fur is brown and turns mostly white in winter.

distribution

Circumpolar in tundra and taiga habitats from Britain to Japan.

habitat

Common in tundra, taiga habitats, and mixed forests. Prefer transition zones with open clearings, and in swamps and river valleys. In Scotland found on heather moorland, and in Ireland and Hokkaido, Japan also on agricultural land.

behavior

A solitary animal. During mating season, it can be seen in groups, sometimes fights upright on its hind legs.

feeding ecology and diet

Leaves and twigs of willow, rowan, birch, juniper, aspen, and heather; in tundra, alpine plants, especially dwarf willow, and blueberry are important. Palatable grasses and herbs are eaten in the summer when available: grasses, sedge, and dicotyledons.

reproductive biology

Promiscuous. In tundra areas, only one litter per year with a mean litter size of five to six young, but up to four litters per year depending on the length of the vegetation season with a mean litter size of two to four. At birth, the young are fully furred, their eyes are open, and they start suckling at once.

conservation status

Very widespread and abundant over most of its range; not threatened.

significance to humans

Important game species; may damage crops and tree plantations.

Annamite striped rabbit

Nesolagus limminsi

taxonomy

Nesolagus limminsi Averianov, Abramov, and Tikhonov, 2000, Huong Son District, Ha Tinh Province, Vietnam.

other common names

French: Lapin Annamite; German: Annamitekaninchen; Spanish: Conejo de Annamite.

physical characteristics

Body length 13.5–15.8 in (35240 cm). Coat buffy gray with brown stripes. Under parts are white.

distribution

Known only from the vicinity of the type locality in Vietnam.

habitat

Forest with thick undergrowth.

behavior

Probably nocturnal.

feeding ecology and diet

Nothing is known.

reproductive biology

Nothing is known.

conservation status

Presumed rare and potentially endangered.

significance to humans

Unknown, but used as food.

European rabbit

Oryctolagus cuniculus

taxonomy

Oryctolagus cuniculus (Linneaus, 1758), Germany.

other common names

French: Lapin de garenne; German: Wildkaninchen; Spanish: Conejo europeo.

physical characteristics

Body length 14–20 in (35–50 cm), weight 3.3–6.6 lb (1,500–3,000 g). The fur is brown with light and black tips, tail white below.

distribution

Original distribution in the Iberian Peninsula and northwest Africa, but introduced 2,000 years ago to the rest of Europe. Later introductions also to Australia, New Zealand, South America, and several islands all over the world.

habitat

In Europe, found in well-drained areas where it is easy to dig dens such as riverbanks. But in introduced areas, it shows great adaptability and inhabits anything from deserts to sub-alpine valleys, including gardens and parks.

behavior

It digs complex burrows for protection against predators and for reproduction. Scent marking is common in social communication, with dominant males being most active.

feeding ecology and diet

Grass, herbs, roots, bark, and cultivated plants.

reproductive biology

Gestation length is about 30 days and they produce five to seven litters per year with a mean size of five to six. It forms highly social groups with one to four males and one to nine females; promiscuous mating.

conservation status

Not threatened.

significance to humans

Many introduced populations have reached pest proportions. Important game animal in many areas, and domestic breeds are used in laboratory research and as pet animals.

Volcano rabbit

Romerolagus diazi

taxonomy

Romerolagus diazi (Ferrari-Pérez, 1893), San Martin Texmelusán, Mexico.

other common names

English: Zacatuche; French: Lapin des volcans; German: Vulkankaninchen; Spanish: Conejo zacatuche, teporingo.

physical characteristics

One of the smallest leporids; body length 10–14 in (27–36 cm), weight 14–18 oz (400–500 g). The ears are short and the hair is dark brown, and it lacks a visible external tail.

distribution

They occur only on the slopes of the Mexican volcanoes, Ajusco and Ixtacihuatl; Popocateptl at 9,190–13,120 ft (2,800–4,000 m).

habitat

Open pine forests with a unique grass habitat, zacaton.

behavior

Vocalizations resemble those of pikas. Mainly active during the days.

feeding ecology and diet

Feeding on the typical zacaton grasses.

reproductive biology

Assumed promiscuous. Average litters size one to three. Breeding season December–July.

conservation status

Listed as Endangered by the IUCN an U.S. Endangered Species Act. Also listed on CITES Appendix I. A very restricted distribution close to Mexico City.

significance to humans

Threatened by habitat destruction, forest fires, agriculture, and property developments. Hunting is now illegal.

Desert cottontail

Sylvilagus audubonii

taxonomy

Sylvilagus audubonii (Baird, 1858), San Francisco Co., California, United States. Twelve subspecies.

other common names

English: Audubon's cottontail; French: Lapin Audubon; German: Audubonkaninchen; Spanish: Conejo del desierto.

physical characteristics

Body length 14–15 in (37–40 cm); tail 1.6–2.4 in (4–6 cm); weight 26.5–44 oz (750–1,250 g). A relatively large cottontail with large ears.

distribution

From Montana south to central Mexico and west to the Californian coast.

habitat

Typically arid areas, but also at higher altitudes.

behavior

Not social, but with overlapping home ranges of up to 15 acres (6 ha).

feeding ecology and diet

Diet mostly grasses, but also some wood species such as Rosa and Rubus.

reproductive biology

Males and females promiscuous. Mean litter size two to three; breeding season from January to August; sexual maturity reached as early as 80 days after birth.

conservation status

Common; not threatened.

significance to humans

Important game species.

Eastern cottontail

Sylvilagus floridanus

taxonomy

Sylvilagus floridanus (J. A. Allen, 1890), Florida, United States. Thirty-five recognized subspecies.

other common names

French: Lapin à queue blanche; German: Florida-Waldkaninchen; Spanish: Conejo castellano.

physical characteristics

Body length 15–19 in (40–48 cm); tail 1–2.4 in (2.5–6 cm); weight 1.8–3.3 lb (800–1,500 g). A large cottontail with brown or grayish fur.

distribution

From southern Canada through central and eastern United States, Central America, and northern South America.

habitat

Widely distributed in many habitats such as woodlands, prairies, farmlands, deserts, and rainforests.

behavior

Do not dig burrows, but females dig nests in holes where the young are reared. Males fight one another to establish dominance hierarchies at mating.

feeding ecology and diet

Herbs and grasses are preferred during growing season, and woody species during winter.

reproductive biology

Promiscuous. Normally three to five litters per year, but could be up to seven. Mean litter size vary geographically from, typically, two to six. Up to 50% of juveniles breed their first year.

conservation status

Common; not threatened.

significance to humans

Important game species, but could sometimes damage crops and forest plantations.

Resources

Books

Chapman, J. A., and J. E. C. Flux, eds. Rabbits, Hares and Pikas, Status Survey and Conservation Action Plan. Gland, Switzerland: International Union for Conservation of Nature, 1990.

Smith, A. J., and D. Bell. "Rabbits and Hares." In The New Encyclopedia of Mammals, edited by D. W. Macdonald. Oxford: Oxford University Press, 2001.

Periodicals

Angerbjörn, A., and J. E. C. Flux. "Lepus timidus." Mammalian Species 495 (1995): 1–11.

Averianov, A. O., A. V. Abramov, and A. N. Tikhonov. "A New Species of Nesolagus (Lagomorpha, Leoporidae) from Vietnam with Osteological Description." Contributions from the Zoological Institute, St. Petersburg 3 (2000): 1–22.

Chapman, J. A., and G. R. Willner. "Sylvilagus audubonii." Mammalian Species 106 (1978): 1–4.

Chapman, J. A., J. G. Hockman, and M. M. Ojeda C. "Sylvilagus floridanus." Mammalian Species 136 (1980): 1–8.

Surridge, A. K., R. J. Timmins, G. M. Hewitt, and D. J. Bell. "Striped Rabbits in Southeast Asia." Nature 400 (1999): 726.

Organizations

IUCN—The World Conservation Union, Species Survival Commission, Lagomorph Specialist Group. Web site: <http://www.ualberta.ca/dhik/lsg/>

Anders Angerbjörn, PhD