Leporidae &mdash; Details

Overview

Comprehensive Description

Diversity

The family Leporidae, consisting primarily of rabbits and hares, includes 54 species from 11 different genera. Leporids range in mass from 300 grams (1.4 lbs) in pygmy rabbits to 5 kilograms (11 lbs) in arctic hares. Adult head and body length ranges from 250 to 700 mm. Unlike most mammals, females are usually larger than males. Color patterns vary between species and across seasons, and range from black to reddish brown to white. Leporids are widely distributed and have adapted to a broad range of habitat types. They can be found throughout the world with very few exceptions. Habitat type affects pelage color as well as litter size. Some leporids are extremely social, living in large communal dens, while others are solitary, coming together in groups or pairs for mating purposes only. The term 'true hares' includes hares and jackrabbits and consists of those species in the genus Lepus; all remaining species are referred to as rabbits. While hares are well adapted for running long distances, rabbits run in short bursts and have modified limbs adapted for digging. Hares have long muscle fibers in contrast to the short fibers found in rabbit muscle. Hares are often larger than rabbits, have black tipped ears, and have distinctly different skull morphologies.

• Gould, E., G. McKay. 1998. The Encyclopedia of Mammals. Sydney and San Francisco: Weldon Owen.
• Nowak, R. 1999. Order Lagomorpha. Pp. 1715-1738 in R Nowak, ed. Walker's Mammals of the World, Vol. 2, Sixth Edition. Baltimore and London: Johns Hopkins University Press.
• Schneider, E. 1990. Hares and Rabbits. Pp. 254-299 in S Parker, ed. Grzimek's Encyclopedia of Mammals, Vol. Volume 4, English Language Editioj Edition. New Jersey and New York: McGraw-Hill Publishing Company.

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Distribution

Geographic Range

Similar to its parent order, Lagomorpha, the family Leporidae has a wide geographic range. Leporids occupy most of the world’s land masses with the exception of southern South America, the West Indies, Madagascar, and most islands southeast of Asia. Although originally absent from South America, Australia, New Zealand, Java, leporids have been introduced to these locations during the last few centuries. The broad geographic range of leporids is largely due to introduction by humans.

Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Introduced ); australian (Introduced ); oceanic islands (Introduced , Native )

Other Geographic Terms: holarctic ; cosmopolitan ; island endemic

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Physical Description

Morphology

Physical Description

Leporids exhibit a great deal of physical diversity. European hares, one of the largest extant members of the family, reach a maximum size of 75 cm and 5 kg and pygmy rabbits, one of the smallest, reach a maximum size of 29.5 cm and 0.46 kg. Domestic leporids can be significantly larger, with an average weight of 7 kg. Female leporids are larger than males, an unusual condition among mammals. Leporids have long hind limbs and feet. Their ears, which are also relatively long, are proximally tubular with the lowest point of the external auditory meatus situated well above the skull. Pelage colors range from brown to black to white. Although spots are relatively common in domestic leporids, most wild species have relatively subdued coloration that helps them blend in with their surroundings. The Sumatran rabbit is one of two species with stripes. Neither albanism nor melanism are uncommon in leporids, and some species that inhabit higher latitudes have white coats during the winter, which are then molted during spring. Most leporids are counter colored, with dark-colored dorsal pelage and light-colored ventral pelage. Pelage texture can be thick and soft or coarse and woolly (e.g., hispid hares) and may become increasingly sparse along the length of the ears. Rabbits and hares have short bushy tales, which are sometimes conspicuously marked, and the soles of their hind feet are covered with hair. The toes terminate in long, slightly curved claws.

Leporid skulls are unmistakeable. They have an arched profile and are only slightly constricted between the orbits, unlike those of their close relatives the pikas. They have prominant post- and supraorbital processes and the parietal, occipital and maxillae are fenestrated. In some species, the squamosals are fenestrated as well. They have a moderately robust zygomatic arch, a relatively short jugal, and tubular external auditory meatuses that are vertically positioned. The dental formula of most leporids is 2/1, 0/0, 3/2, 3/3 = 28. The primary incisors are enlarged, and the secondary are small, peglike, and located immediately posterior to the primaries. The primary incisors resemble those of rodents, except that they are completely encased in enamel. Canines are absent, and a large diastema separates the incisors from the cheek teeth. Their cheekteeth (i.e., molars and premolars) have relatively simple cusp morphology, with the occlusal surface being made up of two transverse ridges (e.g., bilophodont). The cheekteeth are strongly hypsodont in most species.

Rabbits and hares are often differentiated from pikas by the length of their tails and ears. Tail length in leporids ranges from 1.5 cm to 12 cm. Rabbits and hares are characterized by their elongated hind limbs and feet and their ears, which can reach 17 cm in antelope jackrabbit. Pikas have short, rounded ears whereas the ears of leporids are significantly longer than they are wide.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: female larger

• Feldhamer, G., B. Thompson, J. Chapman. 2003. Wild Mammals of North America. Baltimore and London: Johns Hopkins University Press.

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Ecology

Habitat

Leporids can be found in a wide range of environments, from open deserts to boreal forests. Habitat preference and cursorial ability are tightly linked, and as a result, hares and rabbits have distinct habitat requirements. Hares are most often found in open habitat where they can use their speed to evade potential predators. They also rely on their well-camouflaged pelage to hide from predators among the shrubs and rocks. However, some hare species, such as snowshoe hares and Manchurian hares, are well-adapted forest dwellers. While hares are most often found in open habitats, rabbits are confined to habitats with dense cover where they can hide amongst the vegetation or in burrows. Some species of rabbit, such as swamp rabbits and marsh rabbits are excellent swimmers and are considered semi-aquatic. In short, cursorially adept leporids reside in open habitats, whereas cursorially challenged species reside in closed habitats.

Habitat Regions: temperate ; tropical ; polar ; terrestrial

Terrestrial Biomes: tundra ; taiga ; desert or dune ; savanna or grassland ; chaparral ; forest ; rainforest ; scrub forest ; mountains

Wetlands: marsh ; swamp ; bog

Other Habitat Features: suburban ; agricultural ; riparian

• Wilson, D., S. Ruff. 1999. The Smithsonian Book of North American Mammals. Washington and London: Smithsonian Institution Press.
• Hutchins, M. 2004. Lagomorpha. Pp. 417-516 in D Kleiman, V Geist, M McDade, eds. Grzimek's Animal Life Encyclopedia, Vol. 16, Second Edition. New York: Thomson & Gale.
• MacDonald, D. 2001. The Encyclopedia of Mammals. Oxford: Andromeda Oxford Limited.
• Vaughan, T., J. Ryan, N. Czaplewski. 2000. Mammalogy. Fort Worth, TX: Brooks/Cole-Thomson Learning.

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Trophic Strategy

Food Habits

Leporids are obligate herbivores, with diets consisting of grasses, clover, and limited amounts of cruciferous (e.g., plants from the Brassicaceae family such as broccoli and brussels sprouts) and composite plants. They are opportunistic feeders and also eat fruits, seeds, roots, buds, and the bark of trees. During periods of high resource abundance, leporids tend to select forage in pre-reproductive and early reproductive stages of development. In general, the leporid diet is deficient in essential vitamins and micro-nutrients. Plant forage is high in fiber and contains cellulose and lignin as well. Mammals do not possess the digestive enzymes needed to breakdown these compounds. To compensate for this, however, the leporid caecum is up to ten times longer than their stomach and contains a diverse microbial community that helps break down cellulose and lignin. In addition, gut flora passing from the cecum into the small intestine are a significant source of protein for leporids, which have a notoriously protein deficient diet. Leporids are also coprophagic, re-ingesting soft green fecal pellets produced by the cecum. In addition to offsetting their dietary deficiencies, is has been suggested that coprophagy in leporids developed as a predator defense mechanism, allowing them to subsist in the safety of their burrows.

Primary Diet: herbivore (Folivore , Frugivore , Granivore , Lignivore)

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Associations

Ecosystem Roles

Long thought of as pests, rabbits and hares are well known for the damage they inflict on agriculture. As generalist herbivores, leporids are known for their voracious appetite and high reproductive potential. Their role as pests often overshadows their important role in maintaining canivore biological diversity, as leporids are an integral piece of the carnivore food chain. Their importance as a food source for small to medium-sized carnivores is well-illustrated by the 10 year cycle in which Canada lynx abundance closely mimics that of Snowshoe hare.

Leporids are host to a diverse array of endo- and ectoparasites. Many species of parasitic flatworms (Cestoda and Trematoda) and roundworms spend at least part of their lifecycle in the tissues of leporid hosts. Leporids are also vulnerable to various forms of of parasitic arthropods including ticks, mites, fleas, mosquitoes, and flies. Leporids also host various forms of parasitic protozoa (e.g., coccidians). Myxomytosis and rabbit haemorrhagic disease, caused by members of the viral genus Lagovirus, have resulted in the death of millions of wild and domestic leporids.

Ecosystem Impact: keystone species

Commensal/Parasitic Species:

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Predation

Leporids are a major prey item for a large number of mammalian and avian predators including humans, owls, hawks and eagles, falcons, wild, domestic, and feral canids, wild, domestic and feral felids, a number of different mustelid species, and some species of ground squirrel. Predation has likely had a major impact on the evolution of leporids as they are clearly adapted for fast and efficient cursorial locomotion. Their hindlimbs are significantly longer than their forelimbs, which gives them the ability to run in a zig-zag fashion increasing their chances of evading predators. While hares prefer to outrun their pursuers, rabbits find safety in dense cover or in a nearby burrow. Their large ears help them detect approaching predators, and the lateral position of their eyes gives them a complete 360 degree field of vision. Some species, such as snowshoe hare, have large pads on their feet that act as gripping cushions as they run across deep snow to evade predators. Some leporids are especially well adept at hiding from predators. For example, European hares practice motionless “ducking”. Upon detecting an approaching predator, they decrease their heart rate by half, which allows them to remain exceptionally still. Ducking also reduces respiration rates and probably decreases sounds produced during respiration.

Many cold-adapted leporids molt before winter and summer, which helps camouflage them from predators regardless of season. Winter pelage, which is typically snowy-white, consists of longer and denser hair that increases the coat's insulative capabilities. The winter coat is then molted in the spring, as the the typical brown summer pelage returns. Young hares are born above ground and are able to see and evade predators a few hours after birth. Rabbits are often born in a fur-lined underground nest. After nursing, mothers exit this nest from a secure “brooding tube”, which they carefully conceal after each visit. Rabbits are born with their eyes closed, and must be nursed before they are able to evade predators.

Known Predators:

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Known predators

• A. C. Twomey, The bird population of an elm-maple forest with special reference to aspection, territorialism, and coactions, Ecol. Monogr. 15(2):175-205, from p. 202 (1945).
• K. Paviour-Smith, The biotic community of a salt meadow in New Zealand, Trans. R. Soc. N.Z. 83(3):525-554, from p. 542 (1956).
• R. D. Bird, Biotic communities of the Aspen Parkland of central Canada, Ecology, 11:356-442, from p. 410 (1930).
• I. K. Sharma, A study of ecosystems of the Indian desert, Trans. Indian Soc. Desert Technol. and Univ. Center Desert Stud. 5(2):51-55, from p. 52 and A study of agro-ecosystems in the Indian desert, ibid. 5:77-82, from p. 79 1980).
• R. F. Chapman and J. H. P. Sankey, 1955. The larger invertebrate fauna of three rabbit carcasses. J. Anim. Ecol. 24:395-402, from p. 400.

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Known prey organisms

Leporidae (rabbit carcass) preys on:
Pryola
Corylus
Cornus
Aralia
Salix longifolia
Salix petiolaris
leaves
wood
bark
roots
Stipagrostis
Monsonia
Eragrostis
Eleucine
Cyperus
Cenchrus
Zizyphus
Crotalaria

Based on studies in:
New Zealand (Grassland)
USA: Illinois (Forest)
Namibia, Namib Desert (Desert or dune)
India, Rajasthan Desert (Desert or dune)

This list may not be complete but is based on published studies.
• A. C. Twomey, The bird population of an elm-maple forest with special reference to aspection, territorialism, and coactions, Ecol. Monogr. 15(2):175-205, from p. 202 (1945).
• E. Holm and C. H. Scholtz, Structure and pattern of the Namib Desert dune ecosystem at Gobabeb, Madoqua 12(1):3-39, from p. 21 (1980).
• K. Paviour-Smith, The biotic community of a salt meadow in New Zealand, Trans. R. Soc. N.Z. 83(3):525-554, from p. 542 (1956).
• R. D. Bird, Biotic communities of the Aspen Parkland of central Canada, Ecology, 11:356-442, from p. 410 (1930).
• I. K. Sharma, A study of ecosystems of the Indian desert, Trans. Indian Soc. Desert Technol. and Univ. Center Desert Stud. 5(2):51-55, from p. 52 and A study of agro-ecosystems in the Indian desert, ibid. 5:77-82, from p. 79 1980).

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Life History and Behavior

Behavior

Communication and Perception

Very few species of leporids communicate through auditory methods, as most rely on their senses of sight and smell for intraspecific communication. However, certain species (e.g., volcano rabbits) rely heavily on vocalizations for intraspecific communication. Though leporids are typically silent, they still posses a highly developed and acute sense of hearing and emit high pitched distress calls when captured by a predator. For example, European rabbits, brush rabbits, and Audubon's cottontails are known to thump the ground with their hind feet to warn conspecifics of potential danger (e.g., approaching predators). Many leporids have white fur on the ventral surface of their tail, which they silently wave at conspecifics to warn of a predator's presence.

Leporids possess large, protruding eyes that are laterally positioned near the apex of the skull. The position and protrusion of the eyes help them detect predators over a wide visual arc and aid in overcoming the low light availability during crepuscular and nocturnal conditions, during which they are most active.

All Leporids have scent glands in the groin, cheeks, and under the chin that are used to rub pheromones on their coat during grooming. These glands and the pheromones they produce likely play an important role during mating. Glandular activity in male leporids, specifically the amount of pheromone produced and its degree of pungency, is correlated with testicle size. It has been suggested that pheromones serve as a status marker that identify one's position in the social hierarchy.

Communication Channels: visual ; acoustic ; chemical

Other Communication Modes: pheromones ; scent marks ; vibrations

Perception Channels: visual ; acoustic ; vibrations

• Whitaker, J. 1996. National Audobon Society Field Guide to North American Mammals. New York: Alfred A. Knopf.

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Life Expectancy

Lifespan/Longevity

Leporid’s face a number of factors that affect longevity, the most notable being heavy predation from a variety of mammalian, reptilian, and avian predators. In their natural environment, populations of certain species have been shown to have an average lifespan of less than a year. The oldest recorded age for European hares in the wild was 12.5 years with the maximum age estimated to be between 12 to 13 years.

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Reproduction

Most leporid species are polygynandrous. During mating season males and females form small groups in which males compete for access to estrus females and establish a social hierarchy. European Rabbits serve as an exception as they are highly social and have established hierarchies prior to mating season. Males find and attract mates by flagging their tail, involuntary urination, and rubbing against the female prior to copulation. Both sexes have multiple mates and females mate soon after giving birth or while carrying a litter. Gestation typically lasts longer in hares than in rabbits. For example, gestation lasts approximately 55 days in mountain hares and 30 days in European rabbits. Hares are born in a precocial state, fully furred with their eyes open, and are able to run a few hours after parturition. Rabbits are born in an altricial state and are able to see a few days after parturition.

Mating System: polygynandrous (promiscuous)

Some members of the family Leporidae do not have a specific breeding season while others breed during spring and summer. Female ovulation is induced during copulation, about twelve hours after insemination, and females can come into estrus at various times throughout the year. Many species mate immediately after or just before parturition, as females are able to carry two different litters at once (i.e., superfetation). Leporids have high reproductive potential and can produce several litters per breeding season, with several young per litter. Litters usually consist of 2 to 8 young with a maximum of 15 young rabbits (kittens) or hares (leverets) per litter. Resource abundance and quality play a major role in fecundity. For example, Alaskan hares and arctic hares are subjected to prolonged periods of resource scarcity during the winter and have only one litter per year. Black-tailed jackrabbits and antelope jackrabbits live in desert environments and produce several litters a year; however, the litters of these two species are relatively small, containing only 1 to 3 young.

Hares are born fully furred, with open eyes and are able to run a few hours after birth. Rabbits are born with no hair and closed eyes but often have full pelage and open eyes within a couple of days after birth. Sexual maturity and weaning can occur at a young age for both groups but varies according to species. Generally, sexual maturation can occur from 3 to 9 months after birth in rabbits and 1 to 2 years after birth for hares. Females are larger than males in most species, which is unusual in mammals, and are able to reproduce before males. Weaning age is also species specific, but females generally nurse young for at least 3 to 4 weeks, beginning the weaning process about 10 days after parturition.

Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); induced ovulation ; fertilization ; viviparous ; post-partum estrous

Leporids employ a reproductive strategy known as "absentee parentism". In hares, precocial leverets are born in forms, small depression in the ground or surrounding vegetation, while altricial rabbit kittens are born in well-formed, fur-lined nests, constructed in underground chambers or in dense vegetation. Maternal care in leporids is limited to one visit every twenty four hours, usually lasting no more than 5 minutes. Mothers nurse their young during this brief period, which usually occurs during the evening. In species that create subterranean nests for their young, the entrances to these chambers are re-covered after each visit. In form nesting hares, each leveret disperses about 3 days after birth to find their own hiding spot, but rejoin their litter-mates everyday around sunset for their daily nursing bout. Absentee parentism is thought to have evolved as a predator defense mechanism. Leporid milk is extremely rich in fat and protein and is rapidly pumped into offspring during nursing bouts. Paternal care is limited to protecting offspring from rival females.

Prior to the birth of the kittens, rabbit mothers prepare a fir-lined nest for her young. Some species create an underground nest that is either part of a communal den or a remote “brooding tube” dug by the mother for the specific purpose of raising her young. Other species give birth in forms, which consist of small surface depressions filled with chewed-up twigs and leaves, or small depressions among the shrubs. Hares give birth above ground in a nest heap or on a patch of exposed soil.

Hares are precocially born while rabbits are altricially born. Sexual maturity and weaning can occur at a young age for both groups but varies according to species. Weaning generally begins about 10 days after birth and can last anywhere from 17 to 23 days depending on the species. Sexual maturation can occur from 3 to 9 months after birth in rabbits and 1 to 2 years after birth for hares. In social leporids, a mother's position in the hierarchy can affect the social status of their young.

Parental Investment: altricial ; precocial ; male parental care ; female parental care ; pre-hatching/birth (Provisioning: Female); pre-weaning/fledging (Provisioning: Female); pre-independence (Provisioning: Female, Protecting: Male); maternal position in the dominance hierarchy affects status of young

• Feldhamer, G., B. Thompson, J. Chapman. 2003. Wild Mammals of North America. Baltimore and London: Johns Hopkins University Press.
• Gould, E., G. McKay. 1998. The Encyclopedia of Mammals. Sydney and San Francisco: Weldon Owen.
• Hall, E. 1981. Order Lagomorpha. Pp. 286-332 in E Hall, ed. The Mammals of North America, Vol. 1, Second Edition. New York: John Wiley & Sons.
• Hutchins, M. 2004. Mammals and humans: Mammalian invasives and pests. Pp. 182-193 in D Kleiman, V Geist, M McDade, eds. Grzimek's Animal Life Encyclopedia, Vol. 12, Second Edition. New York: Thomsan & Gale.
• MacDonald, D. 2001. The Encyclopedia of Mammals. Oxford: Andromeda Oxford Limited.
• Nowak, R. 1999. Order Lagomorpha. Pp. 1715-1738 in R Nowak, ed. Walker's Mammals of the World, Vol. 2, Sixth Edition. Baltimore and London: Johns Hopkins University Press.
• Schneider, E. 1990. Hares and Rabbits. Pp. 254-299 in S Parker, ed. Grzimek's Encyclopedia of Mammals, Vol. Volume 4, English Language Editioj Edition. New Jersey and New York: McGraw-Hill Publishing Company.

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Evolution and Systematics

Self-medicating prevents disease: rabbits

The ears of rabbits assist in Vitamin D acquisition because they have an oil on the surface that transforms to Vitamin D in sunlight, which is then ingested as the rabbits clean themselves.

"Even rabbits have a therapeutic trick or two - in their case, behind the ears. Mammals need vitamin D - which works with calcium to make healthy bones - in order to prevent such problems as fractures, as well as to keep diseases such as rickets at bay. It is well known that in mammals this vitamin is synthesized when the skin is exposed to sunlight. As noted by John Downer in SuperNatural (1999), rabbits put this principle to good medicinal use when they wash behind their ears with their paws. The oil on the outer surface of the rabbits' extra-long ears contains a chemical that transforms into vitamin D when there is enough sunlight. And when rabbits lick their paws after washing behind their ears, they transfer this vitamin supply to their mouths and, therefore, into their digestive system." (Shuker 2001:218)
• Shuker, KPN. 2001. The Hidden Powers of Animals: Uncovering the Secrets of Nature. London: Marshall Editions Ltd. 240 p.

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats

 Specimen Records: 361 Public Records: 16 Specimens with Sequences: 264 Public Species: 6 Specimens with Barcodes: 255 Public BINs: 6 Species: 17 Species With Barcodes: 17

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Barcode data

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Locations of barcode samples

Collection Sites: world map showing specimen collection locations for Leporidae

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Conservation

Conservation Status

Thirteen species within Leporidae are considered threatened or near-threatened by the International Union for the Conservation of Nature (IUCN), 7 of which are either endangered or critically endangered. Of the 62 species listed by the IUCN, those threatened with extinction are often the most primitive. As leporid habitat is being destroyed to create room for crops, irrigation, and ranch lands, many species of rabbits and hares are forced to persist on remnant habitat islands that result in significantly decreased genetic diversity and ultimately, genetic inbreeding. Many native species are also vulnerable to increased competition for resources with invasive rabbits, the introduction of new pathogens, and the introduction of new predators. While habitat destruction poses the biggest threat to many native leporids, they are also vulnerable to competition with livestock for food resources, over hunting, and poisoning by farmers. Suggested conservation measures include the eradication of exotic predators, reducing habitat destruction and fragmentation, creating strict hunting regulations and enforcing those already in place, the establishment of habitat reserves, and increasing public awareness about the importance of leporid conservation efforts.

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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

Leporids have had a long history of wreaking havoc on ecological systems and agriculture. Their high reproductive potential coupled with humankind’s desire raise them as a domestic animals has resulted in their nearly global distribution. In Australia, European rabbits have been credited with driving many marsupial species to extinction and on the Hawaiian Island of Laysan, rabbits have foraged 22 of 26 native plant species into extinction. Occasionally, leporids can damage crops and compete for forage with livestock.

Leporids can be vectors for many diseases that are transmittable to humans and domesticated animals. The most notable of these pathogens include tularemia or "rabbit fever", myxomatosis, coccidiosis, and pasteurellosis. Most diseases are contracted via the preparation and consumption of tainted meat. However, many diseases, like coccidiosis, are relatively species specific and only pose a threat to humans with significantly weakened immune systems.

Negative Impacts: injures humans (causes disease in humans , carries human disease); crop pest; causes or carries domestic animal disease

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Economic Importance for Humans: Positive

Beginning in the middle ages when Benedictine monks first domesticated them, leporids have had a long and beneficial impact on humans. For centuries rabbits have been an affordable source of protein to the general public, and their dense and soft pelts have provided materials for warm and insulative clothing. Today they are used as model organisms in biomedical research and are popular as game animals and as pets.

Positive Impacts: pet trade ; food ; body parts are source of valuable material; research and education

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Wikipedia

Rabbit

Rabbits are small mammals in the family Leporidae of the order Lagomorpha, found in several parts of the world. There are eight different genera in the family classified as rabbits, including the European rabbit (Oryctolagus cuniculus), cottontail rabbits (genus Sylvilagus; 13 species), and the Amami rabbit (Pentalagus furnessi, an endangered species on Amami Ōshima, Japan). There are many other species of rabbit, and these, along with pikas and hares, make up the order Lagomorpha. The male is called a buck and the female is a doe; a young rabbit is a kitten or kit.

Habitat and range

Outdoor entrance to a rabbit burrow

Rabbit habitats include meadows, woods, forests, grasslands, deserts and wetlands.[1] Rabbits live in groups, and the best known species, the European rabbit, lives in underground burrows, or rabbit holes. A group of burrows is called a warren.[1]

More than half the world's rabbit population resides in North America.[1] They are also native to southwestern Europe, Southeast Asia, Sumatra, some islands of Japan, and in parts of Africa and South America. They are not naturally found in most of Eurasia, where a number of species of hares are present. Rabbits first entered South America relatively recently, as part of the Great American Interchange. Much of the continent has just one species of rabbit, the tapeti, while most of South America's southern cone is without rabbits.

The European rabbit has been introduced to many places around the world.[2]

Biology

Evolution

Because the rabbit's epiglottis is engaged over the soft palate except when swallowing, the rabbit is an obligate nasal breather. Rabbits have two sets of incisor teeth, one behind the other. This way they can be distinguished from rodents, with which they are often confused.[3] Carl Linnaeus originally grouped rabbits and rodents under the class Glires; later, they were separated as the scientific consensus is that many of their similarities were a result of convergent evolution. However, recent DNA analysis and the discovery of a common ancestor has supported the view that they share a common lineage, and thus rabbits and rodents are now often referred to together as members of the superclass Glires.[4]

Morphology

Video of a European rabbit, showing ears twitching and a jump

The rabbit's long ears, which can be more than 10 cm (4 in) long, are probably an adaptation for detecting predators. They have large, powerful hind legs. The two front paws have 5 toes, the extra called the dewclaw. The hind feet have 4 toes.[5] They are plantigrade animals while at rest; however, they move around on their toes while running, assuming a more digitigrade form. Wild rabbits do not differ much in their body proportions or stance, with full, egg-shaped bodies. Their size can range anywhere from 20 cm (8 in) in length and 0.4 kg in weight to 50 cm (20 in) and more than 2 kg. The fur is most commonly long and soft, with colors such as shades of brown, gray, and buff. The tail is a little plume of brownish fur (white on top for cottontails).[2]

Ecology

Rabbits are hindgut digesters. This means that most of their digestion takes place in their large intestine and cecum. In rabbits the cecum is about 10 times bigger than the stomach and it along with the large intestine makes up roughly 40% of the rabbit's digestive tract.[6] The unique musculature of the cecum allows the intestinal tract of the rabbit to separate fibrous material from more digestible material; the fibrous material is passed as feces, while the more nutritious material is encased in a mucous lining as a cecotrope. Cecotropes, sometimes called "night feces", are high in minerals, vitamins and proteins that are necessary to the rabbit's health. Rabbits eat these to meet their nutritional requirements; the mucous coating allows the nutrients to pass through the acidic stomach for digestion in the intestines. This process allows rabbits to extract the necessary nutrients from their food.[7]

Rabbits are prey animals and are therefore constantly aware of their surroundings. For instances, in Mediterranean Europe, rabbits are the main prey of red foxes, badgers, and Iberian lynxes.[8] If confronted by a potential threat, a rabbit may freeze and observe then warn others in the warren with powerful thumps on the ground. Rabbits have a remarkably wide field of vision, and a good deal of it is devoted to overhead scanning.[9] They survive predation by burrowing, hopping away in a zig-zag motion, and, if captured, delivering powerful kicks with their hind legs. Their strong teeth allow them to eat and to bite in order to escape a struggle.[10]

Sleep

Rabbits are crepuscular, most active at dawn and dusk. The average sleep time of a captive rabbit is said to be 8.4 hours.[11]

Lifespan

A litter of rabbit kits (baby rabbits)
A nest containing baby rabbits

The expected rabbit lifespan is about 9–12 years;[12][13] the world's oldest rabbit on record lived 18 years.[14]

Diet and eating habits

A young rabbit looking through the grass.

Rabbits are herbivores that feed by grazing on grass, forbs, and leafy weeds. In consequence, their diet contains large amounts of cellulose, which is hard to digest. Rabbits solve this problem by passing two distinct types of feces: hard droppings and soft black viscous pellets, the latter of which are known as caecotrophs and are immediately eaten (a behaviour known as coprophagy). Rabbits reingest their own droppings (rather than chewing the cud as do cows and many other herbivores) to digest their food further and extract sufficient nutrients.[15]

Rabbits graze heavily and rapidly for roughly the first half hour of a grazing period (usually in the late afternoon), followed by about half an hour of more selective feeding. In this time, the rabbit will also excrete many hard fecal pellets, being waste pellets that will not be reingested. If the environment is relatively non-threatening, the rabbit will remain outdoors for many hours, grazing at intervals. While out of the burrow, the rabbit will occasionally reingest its soft, partially digested pellets; this is rarely observed, since the pellets are reingested as they are produced. Reingestion is most common within the burrow between 8 o'clock in the morning and 5 o'clock in the evening, being carried out intermittently within that period.

Hard pellets are made up of hay-like fragments of plant cuticle and stalk, being the final waste product after redigestion of soft pellets. These are only released outside the burrow and are not reingested. Soft pellets are usually produced several hours after grazing, after the hard pellets have all been excreted. They are made up of micro-organisms and undigested plant cell walls.

The chewed plant material collects in the large cecum, a secondary chamber between the large and small intestine containing large quantities of symbiotic bacteria that help with the digestion of cellulose and also produce certain B vitamins. The pellets are about 56% bacteria by dry weight, largely accounting for the pellets being 24.4% protein on average. These pellets remain intact for up to six hours in the stomach; the bacteria within continue to digest the plant carbohydrates. The soft feces form here and contain up to five times the vitamins of hard feces. After being excreted, they are eaten whole by the rabbit and redigested in a special part of the stomach. This double-digestion process enables rabbits to use nutrients that they may have missed during the first passage through the gut, as well as the nutrients formed by the microbial activity and thus ensures that maximum nutrition is derived from the food they eat.[2] This process serves the same purpose within the rabbit as rumination does in cattle and sheep.[16]

Rabbits are incapable of vomiting.[17]

Rabbit diseases

Rabbits may be affected by a number of diseases. These include pathogens that also affect other animals and/or humans, such as Bordetella bronchiseptica and Escherichia coli', as well as diseases unique to rabbits such as rabbit haemorrhagic disease and myxomatosis.

Rabbits and hares are almost never found to be infected with rabies and have not been known to transmit rabies to humans.[18]

Among the parasites that infect rabbits are tapeworms such as Taenia serialis, external parasites like fleas and mites, coccidia species, and Toxoplasma gondii.[19][20]

Differences from hares

The most obvious difference between rabbits and hares is how their kits are born. Rabbits are altricial, having young that are born blind and hairless. In contrast, hares are precocial, born with hair and good vision. All rabbits except cottontail rabbits live underground in burrows or warrens, while hares live in simple nests above the ground (as do cottontail rabbits), and usually do not live in groups. Hares are generally larger than rabbits, with longer ears, larger and longer hind legs and have black markings on their fur. Hares have not been domesticated, while European rabbits are both raised for meat and kept as pets.

As pets

Domestic rabbits can be kept as pets in a back yard hutch or indoors in a cage or house trained to have free roam. Rabbits kept indoors are often referred to as house rabbits. House rabbits typically have an indoor pen or cage and a rabbit-safe place to run and exercise, such as an exercise pen, living room or family room. Rabbits can be trained to use a litter box and some can learn to come when called. Domestic rabbits that do not live indoors can also serve as companions for their owners, typically living in a protected hutch outdoors. Some pet rabbits live in outside hutches during the day for the benefit of fresh air and natural daylight and are brought inside at night.

Whether indoor or outdoor, pet rabbits' pens are often equipped with enrichment activities such as shelves, tunnels, balls, and other toys. Pet rabbits are often provided additional space in which to get exercise, simulating the open space a rabbit would traverse in the wild. Exercise pens or lawn pens are often used to provide a safe place for rabbits to run.

A pet rabbit's diet typically consists of unlimited timothy-grass or other hay, a small amount of pellets, and a fair quantity of fresh vegetables and need unrestricted access to fresh clean water. Rabbits are social animals. Rabbits as pets can find their companionship with a variety of creatures, including humans, other rabbits, birds, chinchillas, guinea pigs, and sometimes even cats and dogs (however they require supervision when with dogs and cats, as they might be preyed upon or attacked by these animals). Rabbits can make good pets for younger children when proper parental supervision is provided. As prey animals, rabbits are alert, timid creatures that startle fairly easily. They have fragile bones, especially in their backs, that require support on the belly and bottom when picked up. Older children and teenagers usually have the maturity required to care for a rabbit.[21]

Aggression in rabbits

Rabbits may grunt, lunge and even bite. Usually they do not bite hard enough to break skin. Rabbits become aggressive when they feel threatened. This behavior can be corrected with the proper tools. House Rabbit Society[22] says that the owner of the pet needs to win its trust, with certain behavioral tools.

As food and clothing

Rabbit meat sold commercially
Tanned rabbit pelt; rabbit pelt is prized for its softness.
An Australian 'Rabbiter' circa 1900
An old wooden cart, piled with rabbit skins, in New South Wales, Australia

Leporids such as European rabbits and hares are a food meat in Europe, South America, North America, some parts of the Middle East.

Rabbit is still sold in UK butchers and markets, and some supermarkets sell frozen rabbit meat. Additionally, some have begun selling fresh rabbit meat alongside other types of game. At farmers markets and the famous Borough Market in London, rabbits will be displayed dead and hanging unbutchered in the traditional style next to braces of pheasant and other small game. Rabbit meat was once commonly sold in Sydney, Australia, the sellers of which giving the name to the rugby league team the South Sydney Rabbitohs, but quickly became unpopular after the disease myxomatosis was introduced in an attempt to wipe out the feral rabbit population (see also Rabbits in Australia). Rabbit meat is also commonly used in Moroccan cuisine, where it is cooked in a tajine with "raisins and grilled almonds added a few minutes before serving".[23] Rabbit meat is unpopular in the Asia-Pacific.

When used for food, rabbits are both hunted and bred for meat. Snares or guns are usually employed when catching wild rabbits for food. In many regions, rabbits are also bred for meat, a practice called cuniculture. Rabbits can then be killed by hitting the back of their heads, a practice from which the term rabbit punch is derived. Rabbit meat is a source of high quality protein.[24] It can be used in most ways chicken meat is used. In fact, well-known chef Mark Bittman says that domesticated rabbit tastes like chicken because both are blank palettes upon which any desired flavors can be layered.[25] Rabbit meat is leaner than beef, pork, and chicken meat. Rabbit products are generally labeled in three ways, the first being Fryer. This is a young rabbit between 4.5 and 5 pounds and up to 9 weeks in age.[26] This type of meat is tender and fine grained. The next product is a Roaster; they are usually over 5 pounds and up to 8 months in age. The flesh is firm and coarse grained and less tender than a fryer. Then there are giblets which include the liver and heart. One of the most common types of rabbit to be bred for meat is New Zealand white rabbit.

There are several health issues associated with the use of rabbits for meat, one of which is tularemia or rabbit fever.[27] Another is so-called rabbit starvation, due most likely to deficiency of essential fatty acids in rabbit meat. Rabbits are a common food item of large snakes, such as Burmese pythons and reticulated pythons, both in the wild and in captivity.

Rabbit pelts are sometimes used for clothing and accessories, such as scarves or hats. Angora rabbits are bred for their long, fine hair, which can be sheared and harvested like sheep wool. Rabbits are very good producers of manure; additionally, their urine, being high in nitrogen, makes lemon trees very productive. Their milk may also be of great medicinal or nutritional benefit due to its high protein content.[28]

Environmental problems

Rabbits have been a source of environmental problems when introduced into the wild by humans. As a result of their appetites, and the rate at which they breed, feral rabbit depredation can be problematic for agriculture. Gassing, barriers (fences), shooting, snaring, and ferreting have been used to control rabbit populations, but the most effective measures are diseases such as myxomatosis (myxo or mixi, colloquially) and calicivirus. In Europe, where rabbits are farmed on a large scale, they are protected against myxomatosis and calicivirus with a genetically modified virus. The virus was developed in Spain, and is beneficial to rabbit farmers. If it were to make its way into wild populations in areas such as Australia, it could create a population boom, as those diseases are the most serious threats to rabbit survival. Rabbits in Australia and New Zealand are considered to be such a pest that land owners are legally obliged to control them.[29][30]

When introduced into a new area, rabbits can overpopulate rapidly, becoming a nuisance, as on this university campus
European Rabbit in Shropshire, England, infected with myxomatosis, a disease caused by the Myxoma virus

In culture and literature

Rabbits are often used as a symbol of fertility or rebirth, and have long been associated with spring and Easter as the Easter Bunny. The species' role as a prey animal also lends itself as a symbol of innocence, another Easter connotation.

Additionally, rabbits are often used as symbols of playful sexuality, which also relates to the human perception of innocence, as well as its reputation as a prolific breeder.

Folklore and mythology

The rabbit often appears in folklore as the trickster archetype, as he uses his cunning to outwit his enemies.

• In Aztec mythology, a pantheon of four hundred rabbit gods known as Centzon Totochtin, led by Ometotchtli or Two Rabbit, represented fertility, parties, and drunkenness.
• In Central Africa, the common hare (Kalulu), is "inevitably described" as a trickster figure.[31]
• In Chinese folklore, rabbits accompany Chang'e on the Moon. Also associated with the Chinese New Year (or Lunar New Year), rabbits are also one of the twelve celestial animals in the Chinese Zodiac for the Chinese calendar. It is interesting to note that the Vietnamese lunar new year replaced the rabbit with a cat in their calendar, as rabbits did not inhabit Vietnam.
• A rabbit's foot is carried as an amulet believed to bring good luck. This is found in many parts of the world, and with the earliest use being in Europe around 600 B.C.[32]
• In Japanese tradition, rabbits live on the Moon where they make mochi, the popular snack of mashed sticky rice. This comes from interpreting the pattern of dark patches on the moon as a rabbit standing on tiptoes on the left pounding on an usu, a Japanese mortar (See also: Moon rabbit).
• In Jewish folklore, rabbits (shfanim שפנים) are associated with cowardice, a usage still current in contemporary Israeli spoken Hebrew (similar to English colloquial use of "chicken" to denote cowardice).
• In Korean mythology, like in Japanese, presents rabbits living on the moon making rice cakes (Tteok in Korean).
• In Anishinaabe traditional beliefs, held by the Ojibwe and some other Native American peoples, Nanabozho, or Great Rabbit, is an important deity related to the creation of the world.
• A Vietnamese mythological story portrays the rabbit of innocence and youthfulness. The Gods of the myth are shown to be hunting and killing rabbits to show off their power.

On the Isle of Portland in Dorset, UK, the rabbit is said to be unlucky and speaking its name can cause upset with older residents. This is thought to date back to early times in the quarrying industry, where piles of extracted stone (not fit for sale) were built into tall rough walls (to save space) directly behind the working quarry face; the rabbit's natural tendency to burrow would weaken these "walls" and cause collapse, often resulting in injuries or even death. The name rabbit is often substituted with words such as “long ears” or “underground mutton”, so as not to have to say the actual word and bring bad luck to oneself. It is said that a public house (on the island) can be cleared of people by calling out the word rabbit and while this was very true in the past, it has gradually become more fable than fact over the past 50 years. See also Three hares.

Other fictional rabbits

The rabbit as trickster appears in American popular culture; for example the Br'er Rabbit character from African-American folktales and Disney animation; and the Warner Bros. cartoon character Bugs Bunny.

Anthropomorphized rabbits have appeared in a host of works of film, literature, and technology, notably the White Rabbit and the March Hare in Lewis Carroll's Alice's Adventures in Wonderland; in the popular novels Watership Down, by Richard Adams (which has also been made into a movie), Rabbit Hill by Robert Lawson, as well as in Beatrix Potter's Peter Rabbit stories and Oswald the Lucky Rabbit from 1920s and 1930s cartoons.

Urban legends

It was commonly believed that pregnancy tests were based on the idea that a rabbit would die if injected with a pregnant woman's urine. This is not true. However, in the 1920s it was discovered that if the urine contained the hCG, a hormone found in the bodies of pregnant women, the rabbit would display ovarian changes. The rabbit would then be killed to have its ovaries inspected, but the death of the rabbit was not the indicator of the results. Later revisions of the test allowed technicians to inspect the ovaries without killing the animal. A similar test involved injecting Xenopus frogs to make them lay eggs, but animal tests for pregnancy have been made obsolete by faster, cheaper, and simpler modern methods.

Classifications

Eastern Cottontail (Sylvilagus floridanus)

Rabbits and hares were formerly classified in the order Rodentia (rodent) until 1912, when they were moved into a new order Lagomorpha. This order also includes pikas.

Order Lagomorpha

 Rabbits and hares portal

References

1. ^ a b c "Rabbit Habitats". Retrieved 2009-07-07.
2. ^ a b c "rabbit". Encyclopædia Britannica (Standard ed.). Chicago: Encyclopædia Britannica, Inc. 2007.
3. ^ Brown, Louise (2001). How to Care for Your Rabbit. Kingdom Books. p. 6. ISBN 978-1-85279-167-4.
4. ^ Katherine Quesenberry & James W. Carpenter, Ferrets, Rabbits, and Rodents: Clinical Medicine and Surgery (3rd ed. 2011).
5. ^ "Rabbits: Rabbit feet". Retrieved 2010-07-13.
6. ^ "Feeding the Pet Rabbit"
7. ^ Dr. Byron de la Navarre's "Care of Rabbits" Susan A. Brown, DVM's "Overview of Common Rabbit Diseases: Diseases Related to Diet"
8. ^ Fedriani, J.M., Palomares, F., Delibes, M (1999). 23/Fedriani.pdf "Niche relations among three sympatric Mediterranean carnivores". Oecologia 121: 138–148. doi:10.1007/s004420050915. JSTOR 4222449.
9. ^ Tynes, Valarie V. Behavior of Exotic Pets. Wiley Blackwell, 2010, p. 70
10. ^ Davis, Susan E. and DeMello, Margo Stories Rabbits Tell: A Natural And Cultural History of A Misunderstood Creature. Lantern Books, 2003, p. 27.
11. ^ "40 Winks?" Jennifer S. Holland, National Geographic Vol. 220, No. 1. July 2011.
12. ^ Animal Lifespans from Tesarta Online (Internet Archive)
13. ^ The Life Span of Animals from Dr Bob's All Creatures Site
14. ^ "What's the lifespan of a rabbit?". House Rabbit Society. Retrieved 2010-09-27.
15. ^ "Information for Rabbit Owners — Oak Tree Veterinary Centre". Oaktreevet.co.uk. Retrieved 2010-08-30.
16. ^ The Private Life of the Rabbit, R. M. Lockley, 1964. Chapter 10.
17. ^
18. ^ "Rabies: Other Wild Animals". Centers for Disease Control and Prevention. 15 November 2011. Retrieved 20 December 2012.
19. ^ Wood, Maggie. "Parasites of Rabbits". Chicago Exotics, PC. Retrieved 8 April 2013.
20. ^ Boschert, Ken. "Internal Parasites of Rabbits". Net Vet. Retrieved 8 April 2013.
21. ^ "Children and Rabbits". Rabbit.org. Retrieved 2010-08-30.
22. ^ House Rabbit Society, additional text.
24. ^
25. ^ "How to Cook Everything :: Braised Rabbit with Olives". 2008. Archived from the original on 2008-05-17. Retrieved 2008-07-17.
26. ^ Sell, Randy Rabbit. North Dakota Department of Agricultural Economics.
27. ^ "Tularemia (Rabbit fever)". Health.utah.gov. 2003-06-16. Retrieved 2010-08-30.
28. ^ Houdebine, Louis-Marie; Fan, Jianglin (1 June 2009). Rabbit Biotechnology: Rabbit Genomics, Transgenesis, Cloning and Models. シュプリンガー・ジャパン株式会社. pp. 68–72. ISBN 978-90-481-2226-4. Retrieved 8 October 2010.
29. ^ "Feral animals in Australia — Invasive species". Environment.gov.au. 2010-02-01. Retrieved 2010-08-30.
30. ^ "Rabbits — The role of government — Te Ara Encyclopedia of New Zealand". Teara.govt.nz. 2009-03-01. Retrieved 2010-08-30.
31. ^ Brian Morris, The Power of Animals: An Ethnography, p. 177 (2000).
32. ^ Ellis, Bill: Lucifer Ascending: The Occult in Folklore and Popular Culture (University of Kentucky, 2004) ISBN 0-8131-2289-9

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Leporidae

Leporids are over 60 species of rabbits and hares which form the family Leporidae. The leporids, together with the pikas, constitute the mammalian order Lagomorpha. Leporids differ from pikas in having short, furry tails, and elongated ears and hind legs. The name leporid is simply an abbreviation of the family name Leporidae, meaning animals resembling lepus, Latin for hare.

Members of all genera except Lepus are usually referred to as rabbits, while members of Lepus (which accounts for almost half the species) are usually called hares. However, the distinction between these two common names does not map completely into current taxonomy, since jackrabbits are members of Lepus, and members of the genera Pronolagus and Caprolagus are sometimes called hares.

Leporids are native across the world except Antarctica, and in Oceania, where their introduction is a significant threat for the native mammals in Australia.

Characteristics

Leporids are small to moderately sized mammals, adapted for rapid movement. They have long hind legs, with four toes on each foot, and shorter fore legs, with five toes each. The soles of their feet are hairy, to improve grip while running, and they have strong claws on all of their toes. Leporids also have distinctive, elongated and mobile ears, and they have an excellent sense of hearing. Their eyes are large, and their night vision is good, reflecting their primarily nocturnal or crepuscular mode of living.[2]

Leporids range in size from the pygmy rabbit (Brachylagus idahoensis), with a head and body length of 25–29 cm, and a weight of around 300 grams, to the European hare (Lepus europaeus), which is 50–76 cm in head-body length, and weighs from 2.5 to 5 kilograms.

Both rabbits and hares are almost exclusively herbivorous (with exceptions among the members of Lepus),[3][4] feeding primarily on grasses and herbs, although they also eat leaves, fruit, and seeds of various kinds. They are coprophagous, as they pass food through their digestive systems twice, first expelling it as soft green feces, which they then reingest, eventually producing hard, dark fecal pellets. Like rodents, they have powerful front incisor teeth, but they also have a smaller second pair of incisors to either side of the main teeth in the upper jaw, and the structure is different from that of rodent incisors. Also like rodents, leporids lack any canine teeth, but they do have more cheek teeth than rodents do. Their jaws also contain a large diastema. The dental formula of most, though not all, leporids is: $Upper: 2.0.3.3, lower: 1.0.2.3$

They have adapted to a remarkable range of habitats, from desert to tundra, forests, mountains, and swampland. Rabbits generally dig permanent burrows for shelter, the exact form of which varies between species. In contrast, hares rarely dig shelters of any kind, and their bodies are more suited to fast running than to burrowing.[2]

The gestation period in leporids varies from around 28 to 50 days, and is generally longer in the hares. This is in part because young hares, or leverets, are born fully developed, with fur and open eyes, while rabbit kits are naked and blind at birth, having the security of the burrow to protect them.[2] Leporids can have several litters a year, which can cause their population to expand dramatically in a short period of time when resources are plentiful.

Evolution

Serengetilagus praecapensis skull, Naturkundemuseum, Berlin

The oldest known leporid species date from the late Eocene, by which time the family was already present in both North America and Asia. Over the course of their evolution, this group has become increasingly adapted to lives of fast running and leaping. For example, Palaeolagus, an extinct rabbit from the Oligocene of North America, had shorter hind legs than modern forms (indicating it ran rather than hopped) though it was in most other respects quite rabbit-like.[5] Two as yet unnamed fossil finds—dated ~48 Ma (from China) and ~53 Ma (India)—while primitive, display the characteristic leporid ankle, thus pushing the divergence of Ochotonidae and Leporidae yet further into the past.[6] The genus Praotherium was once considered to be be part of this family,[7] but this is now in doubt.[8]

Classification

Family Leporidae:[1] rabbits and hares

 Rabbits and hares portal

References

1. ^ a b Hoffman, R. S.; Smith, A. T. (2005). "Order Lagomorpha". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. pp. 194–211. ISBN 978-0-8018-8221-0. OCLC 62265494.
2. ^ a b c Chapman, J. & Schneider, E. (1984). MacDonald, D.. ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 714–719. ISBN 0-87196-871-1.
3. ^ Best, Troy L.; Henry, Travis Hill (1994). "Lepus arcticus". Mammalian Species (American Society of Mammalogists) (457): 1–9. June 2, 1994. doi:10.2307/3504088. JSTOR 3504088. OCLC 46381503
4. ^ "Snowshoe Hare". eNature: FieldGuides. eNature.com. 2007. Retrieved 2008-03-23.
5. ^ Savage, RJG, & Long, MR (1986). Mammal Evolution: an illustrated guide. New York: Facts on File. pp. 128–129. ISBN 0-8160-1194-X.
6. ^ Handwerk, Brian (2008-03-21). "Easter Surprise: World's Oldest Rabbit Bones Found". National Geographic News (National Geographic Society). Retrieved 2008-03-23.
7. ^
8. ^ "Praotherium palatinum (nomen dubium)". The Paleobiology Database. Retrieved 16 January 2013.

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