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Lagomorpha

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Lagomorphs[1]
Temporal range: Late Paleocene – recent
Eastern cottontail (Sylvilagus floridanus)
Large-eared pika (Ochotona macrotis)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Grandorder: Glires
Mirorder: Duplicidentata
Order: Lagomorpha
Brandt, 1855
Families
Range of Lagomorpha including areas of human introduction
Fossil occurrences of leporids and ochotonids and global environmental change (climate change, C3/C4 plants distribution).[2]

The lagomorphs (/ˈlæɡəmɔːrf/) are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae (rabbits and hares) and the Ochotonidae (pikas). There are 110 recent species of lagomorph of which 109 are extant, including 10 genera of rabbits (42 species), 1 genus of hare (33 species) and 1 genus of pika (34 species). The name of the order is derived from the Ancient Greek lagos (λαγώς, "hare") + morphē (μορφή, "form").

Taxonomy and evolutionary history

Other names used for this order, now considered synonymous, include: Duplicidentata - Illiger, 1811; Leporida - Averianov, 1999; Neolagomorpha - Averianov, 1999; Ochotonida - Averianov, 1999; and Palarodentia - Haeckel, 1895, Lilian, 2016.[1]

The evolutionary history of the lagomorphs is still not well understood. Until recently, it was generally agreed that Eurymylus, which lived in eastern Asia and dates back to the late Paleocene or early Eocene, was an ancestor of the lagomorphs.[3] More recent examination of the fossil evidence suggests that the lagomorphs may have instead descended from Anagaloidea, also known as "mimotonids", while Eurymylus was more closely related to rodents (although not a direct ancestor).[4] The leporids first appeared in the late Eocene and rapidly spread throughout the Northern Hemisphere; they show a trend towards increasingly long hind limbs as the modern leaping gait developed. The pikas appeared somewhat later in the Oligocene of eastern Asia.[5]

Lagomorphs were certainly more diverse in the past than in the present, with around 75 genera and over 230 species represented in the fossil record and many more species in a single biome. This is evidence that lagomorph lineages are declining.[6]

Recent finds suggest an Indian origin for the order, having possibly evolved in isolation when India was an island continent in the Paleocene.[7]

Characteristics

Lagomorphs are similar to other mammals in that they all have hair, four limbs (i.e., they are tetrapods), and mammary glands and are endotherms. Lagomorphs possess a moderately fused postorbital process to the cranium, unlike other small mammals.[8] They differ in that they have a mixture of "basal" and "derived" physical traits.

Differences between lagomorphs and other mammals

Lagomorphs and rodents form the clade or grandorder Glires. Despite the evolutionary relationship between lagomorphs and rodents, the two orders have some major differences.

Lagomorphs have four incisors in the upper jaw (smaller peg teeth behind larger incisors), whereas rodents only have two. They are similar to rodents in that their incisors grow continuously, thus necessitating constant chewing on fibrous food to prevent the teeth from growing too long. [9][10] In addition, all lagomorph teeth grow continuously,[11] while for most rodents, only the incisors grow continuously. [12] Lagomorph & rodent incisors are structured differently. Lagomorphs have more cheek teeth than rodents. Both have a large diastema.

Lagomorphs are almost strictly herbivorous, unlike rodents, many of which will eat both meat and vegetable matter. Lagomorphs have no paw pads; instead, the bottoms of their paws are entirely covered with fur. [13] [14] Similar to the rodents, bats, and some mammalian insectivores, they have a smooth-surfaced cerebrum. [15] Lagomorphs are unusual among terrestrial mammals in that the females are larger than males. [16]

Differences between families of lagomorphs

Rabbits and hares move by jumping, pushing off with their strong hind legs and using their forelimbs to soften the impact on landing. Pikas lack certain skeletal modifications present in leporids, such as a highly arched skull, an upright posture of the head, strong hind limbs and pelvic girdle, and long limbs.[17] Also, pikas have a short nasal region and entirely lack a supraorbital foramen, while leporids have prominent supraorbital foramina and nasal regions.[18]

Pikas

American pika in Alberta

Pikas, also known as conies,[19] are entirely represented by the family Ochotonidae and are small mammals native to mountainous regions of western North America and Central Asia. They are mostly about 15 cm (6 in) long and have greyish-brown, silky fur, small rounded ears, and almost no tail. Their four legs are nearly equal in length. Some species live in scree, making their homes in the crevices between broken rocks, while others construct burrows in upland areas. The rock-dwelling species are typically long-lived and solitary, having one or two small litters each year contributing to stable populations. The burrowing species, in contrast, are short-lived, gregarious and have multiple large litters during the year. These species tend to have large swings in population size. The gestation period of the pika is around one month long, and the newborns are altricial (eyes and ears closed, no fur).[20] The social behaviour of the two groups also differs: the rock dwellers aggressively maintain scent-marked territories, while the burrowers live in family groups, interact vocally with each other and defend a mutual territory. Pikas are diurnal and are active early and late in the day during hot weather. They feed on all sorts of plant material. As they do not hibernate, they make "haypiles" of dried vegetation which they collect and carry back to their homes to store for use during winter.[17]

Hares

Scrub hare in South Africa

Hares, members of genus Lepus of family Leporidae, are medium size mammals native to Europe, Asia, Africa, and North America. North American jackrabbits are actually hares. Species vary in size from 40 to 70 cm (16 to 28 in) in length and have long powerful back legs, and ears up to 20 cm (8 in) in length. Although usually greyish-brown, some species turn white in the winter. They are solitary animals. Newborns are precocial (eyes and ears open, fully furred). Several litters are born during the year in a form (a nest above ground, usually under a bush). They are preyed upon by large mammalian carnivores and birds of prey.[21]

Rabbits

Rabbits, members of the Leporidae family (excluding Lepus (hares)) are generally much smaller than hares and include the rock hares and the hispid hare. They are native to Europe, parts of Africa, Central and Southern Asia, North America and much of South America. They inhabit both grassland and arid regions. They vary in size from 20 to 50 cm (8 to 20 in) and have long, powerful hind legs, shorter forelegs and a tiny tail. The colour is some shade of brown, buff or grey and there is one black species and two striped ones. Domestic rabbits come in a wider variety of colours. Newborn rabbits are altricial (eyes and ears closed, no fur). Although most species live in burrows, the cottontails and hispid hares have forms (nests above ground, usually under a bush). Most of the burrowing species are colonial, and feed together in small groups. Rabbits play an important part in the terrestrial food chain, eating a wide range of forbs, grasses, and herbs, and being part of the staple diet of many carnivorous species. Domestic rabbits can be litter box trained, and—assuming they are given sufficient room to run and a good diet—can live long lives as house pets.

Distribution

Lagomorphs are widespread around the world and inhabit every continent except Antarctica. However, they are not found in most of the southern cone of South America, in the West Indies, Indonesia or Madagascar, nor on many islands. Although they are not native to Australia, humans have introduced them there and they have successfully colonized many parts of the country and caused disruption to native species.[22]

Biology

Digestion

Skeleton of Alaskan hare (Museum of Osteology)

Like other herbivores, lagomorphs have to deal with a bulky diet in which the cell walls are composed of cellulose, a substance which mammalian digestive enzymes are unable to break down. Despite this, lagomorphs have developed a way of extracting maximum nourishment from their diet. First they bite off and shred plant tissues with their incisors and then they grind the material with their molars. Digestion continues in the stomach and small intestine where nutrients are absorbed. After that, certain food remains get diverted into the caecum, a blind-ended pouch. Here, they are mixed with bacteria, yeasts and other micro-organisms that are able to digest cellulose and turn it into sugar, a process known as hindgut fermentation. Other faecal matter passes along the colon and is excreted in the normal way as small, dry pellets. About four to eight hours after the meal, the contents of the caecum pass into the colon and are eliminated as soft, moist pellets known as cecotropes. These are immediately eaten by the lagomorph, which can thus extract all the remaining nutrients in the food.[23]

Birth and early life

Many lagomorphs breed several times a year and produce large litters. This is particularly the case in species that live in underground, protective environments, such as burrows. The young of rabbits and pikas (called kits) are born after a short gestation period and the mother can become pregnant again almost immediately after giving birth. The mothers are able to leave these young safely and go off to feed, returning at intervals to feed them with their unusually rich milk. In some species, the mother only visits and feeds the litter once a day but the young grow rapidly and are usually weaned within a month.

Hare young are called leverets. Adults have a strategy to prevent predators from tracking down their litter by following the adults' scent. They approach and depart from the nesting site in a series of immense bounds, sometimes moving at right angles to their previous direction.[24] Each litter of hares have a small number of young and are born after a longer gestation period. [10]

Sociality and safety

Many species of lagomorphs, particularly the rabbits and the pikas, are gregarious and live in colonies, whereas hares are generally solitary species, although many hares travel and forage in groups of two, three, or four. Many rabbits and pikas rely on their burrows as places of safety when danger threatens, but hares rely on their long legs, great speed and jinking gait to escape from predators.

Classification

Alloptox japonicus jaw
Palaeolagus skull

Recent genera

Fossil genera

References

  1. ^ a b c d Hoffman, R.S.; Smith, A.T. (2005). "Order Lagomorpha". In Wilson, D.E.; Reeder, D.M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 185–211. ISBN 978-0-8018-8221-0. OCLC 62265494.
  2. ^ Ge, Deyan; Wen, Zhixin; Xia, Lin; Zhang, Zhaoqun; Erbajeva, Margarita; Huang, Chengming; Yang, Qisen (April 3, 2013). "Evolutionary History of Lagomorphs in Response to Global Environmental Change". PLoS ONE. 8 (4:e59668): e59668. Bibcode:2013PLoSO...859668G. doi:10.1371/journal.pone.0059668. PMC 3616043. PMID 23573205.
  3. ^ Palmer, D., ed. (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals. London: Marshall Editions. p. 285. ISBN 1-84028-152-9.
  4. ^ Rose, Kenneth David (2006). The Beginning of the Age of Mammals. The Johns Hopkins University Press. p. 315. ISBN 0-8018-8472-1.
  5. ^ Savage, RJG (1986). Mammal Evolution: an illustrated guide. Illustrated by Long, MR. New York: Facts on File. pp. 128–129. ISBN 0-8160-1194-X.
  6. ^ Alves, Paulo C.; Ferrand, Nuno; Hackländer, Klaus, eds. (2008). "Lopez-Martinez - The Lagomorph Fossil Record". Lagomorph Biology: Evolution, Ecology, and Conservation. Berlin, Heidelberg: Springer. pp. 27–46. doi:10.1007/978-3-540-72446-9. ISBN 978-3-540-72445-2. OCLC 166358165. ...which denotes that lagomorph lineages are also declining in recent times.
  7. ^ Rose, K.D.; Deleon, V.B.; Mmissian, P.; Rana, R.S.; Sahni, A.; Singh, L.; Smith, T. (2008). "Early Eocene lagomorph (Mammalia) from western India and the early diversification of Lagomorpha". Proceedings of the Royal Society B. 275 (1639): 1203–1208. doi:10.1098/rspb.2007.1661. PMC 2602686. PMID 18285282.
  8. ^ Wible, John R. (January 2007). "On the Cranial Osteology of the Lagomorpha". Bulletin of Carnegie Museum of Natural History. 2007 (39): 213–234. doi:10.2992/0145-9058(2007)39[213:OTCOOT]2.0.CO;2. ISSN 0145-9058. S2CID 85766674.
  9. ^ Best, T. L.; Henry, T. H. (1994-06-02). "Lepus arcticus". Mammalian Species (457): 1–9. doi:10.2307/3504088. ISSN 0076-3519. JSTOR 3504088. S2CID 253989268.
  10. ^ a b Smith, Andrew T. "Lagomorph". Encyclopædia Britannica. Retrieved 2020-09-08.
  11. ^ Myers, Phil. "Lagomorpha; hares, pikas, and rabbits". Animal Diversity (ADW). Retrieved 2024-01-03.
  12. ^ "Structure and placement of individual teeth". Animal Diversity (ADW). Retrieved 2024-01-03.
  13. ^ "Pika; mammal". Encyclopedia Britannica. Retrieved 2021-06-26.
  14. ^ Macdonald, David W. (David Whyte) (1984). The Encyclopedia of mammals. Internet Archive. New York, NY : Facts on File. ISBN 978-0-87196-871-5.
  15. ^ Ferrer, I.; Fabregues, I.; Condom, E. (1986). "A Golgi study of the sixth layer of the cerebral cortex I: The lissencephalic brain of Rodentia, Lagomorpha, Chiroptera, and Insectivora" (PDF). Journal of Anatomy. 145: 217–234. PMC 1166506. PMID 3429306. Archived (PDF) from the original on 2021-05-11.[need quotation to verify]
  16. ^ Ralls, Katherine (June 1976). "Mammals in Which Females are Larger Than Males". The Quarterly Review of Biology. 51 (2): 245–276. doi:10.1086/409310. PMID 785524. S2CID 25927323.
  17. ^ a b Smith, Andrew T. "Pika". Encyclopædia Britannica. Retrieved 2020-09-08.
  18. ^ "IUCN - Lagomorph specialist group". www.iucn.org. Archived from the original on 2015-08-03. Retrieved 2015-08-18.
  19. ^ "Lagomorphs - EnchantedLearning.com". www.enchantedlearning.com. Retrieved 2015-08-15.
  20. ^ "American Pika". National Wildlife Federation. Retrieved 2018-11-19.
  21. ^ Smith, Andrew T. "Hare". Encyclopædia Britannica. Retrieved 2020-09-08.
  22. ^ Klappenbach, Laura. "Hares, Rabbits and Pikas". About.com. Archived from the original on 2013-10-20. Retrieved 2013-08-14.
  23. ^ "Exploring a Rabbit's Unique Digestive System". Rabbits for Dummies. Retrieved 2013-08-14.
  24. ^ Burton, Maurice (1971). The Observer's Book of British Wild Animals. Frederick Warne & Co. pp. 109–112. ISBN 9780723215035.
  25. ^ a b "Lagomorpha". Fossilworks. Gateway to the Paleobiology Database. Retrieved 13 May 2010.