Elapidae

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Elapidae
L14cobra.jpg
Egyptian cobra, Naja haje
Scientific classification
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Elapidae

F. Boie, 1827
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The Elapidae (Greek ἔλλοψ éllops, "sea-fish") [1] are a family of venomous snakes found in tropical and subtropical regions around the world, terrestrially in Asia, Australia, Africa, North America, and South America, and aquatically in the Pacific and Indian Oceans.

Elapid snakes exhibit a wide range of sizes, from 18-cm species of Drysdalia to the 5.6-m king cobra. All elapids are characterized by hollow, fixed fangs through which they inject venom. Currently, 325 species in 61 genera are recognized;[2] 58 genera and 251 species are in the Old World, against a minor diversity of three genera and 74 species in the New World.

Description

All elapids have a pair of proteroglyphous fangs used to inject venom from glands located towards the rear of the upper jaws. In outward appearance, terrestrial elapids look similar to the Colubridae: almost all have long and slender bodies with smooth scales, a head covered with large shields and not always distinct from the neck, and eyes with round pupils. In addition, their behavior is usually quite active, and most are oviparous. Exceptions to all these generalizations occur: e.g. the death adders (Acanthophis) include short and fat, rough-scaled, very broad-headed, cat-eyed, live-bearing, sluggish ambush predators with partly fragmented head shields.

Some elapids are strongly arboreal (African Pseudohaje and Dendroaspis, Australian Hoplocephalus), while many others are more or less specialised burrowers (e.g. Ogmodon, Parapistocalamus, Simoselaps, Toxicocalamus, and Vermicella) in either humid or arid environments. Some species have very generalised diets (euryophagy), but many taxa have narrow prey preferences (stenophagy) and correlated morphological specialisations, e.g. for feeding on other snakes, elongated burrowing lizards, squamate eggs, mammals, birds, frogs, fish, etc.

Sea snakes (the Hydrophiinae, sometimes considered to be a separate family) have adapted to a marine way of life in different ways and to various degrees. All have evolved paddle-like tails for swimming and the ability to excrete salt. Most also have laterally compressed bodies, their ventral scales are much reduced in size, their nostrils are located dorsally (no internasal scales), and they give birth to live young (ovoviviparous). In general, they have the ability to respire through their skin; experiments with the yellow-bellied sea snake, Pelamis platurus, have shown that this species can satisfy about 20% of its oxygen requirements in this manner, allowing for prolonged dives. The sea kraits (Laticauda spp.) are the sea snakes least adapted to aquatic life. They spend much of their time on land, where they lay their eggs. They have wide ventral scales, their tails are not as well-developed for swimming, and their nostrils are separated by internasal scales.

Dentition

The fangs, which are enlarged and hollow, are the first two teeth on each maxillary bone. Usually only one fang is in place on each side at any time. The maxilla is intermediate in both length and mobility between typical colubrids (long, less mobile) and viperids (very short, highly mobile). When the mouth is closed, the fangs fit into grooved slots in the buccal floor and usually below the front edge of the eye and are angled backwards; some elapids (Acanthophis, Oxyuranus, Dendroaspis, and Ophiophagus) have long fangs on quite mobile maxillae and can make fast strikes.

A few species are capable of spraying their venom from forward-facing holes at the tips of their fangs using pressure, and this can act as a means of defense.

Distribution

On land, these snakes are found worldwide in tropical and subtropical regions, except in Europe. Sea snakes occur mainly in the Indian Ocean and the southwest Pacific. However, the range of one species, Pelamis platura, extends across the Pacific to the coasts of Central and South America.[3]

Venom

Many elapids are potentially deadly venomous snakes. Their venom is mainly neurotoxic, although many of them also possess several other types of toxins, including cardiotoxins and cytotoxins. This family has some members considered to be the world's most venomous land snakes based on the murine LD50 of their venom, such as the inland taipan.[4] Additionally, some large-sized elapids, such as the Asiatic king cobra, African black mamba, forest cobra, and Australasian coastal taipan, can inject a large quantity of venom during envenomation.

Elapids use their venom both to immobilize their prey and in self-defense.

Genera

Genus[2] Taxon
author[2]
Species[2] Subspecies*[2] Common
name
Geographic
range[3]
Acalyptophis Boulenger, 1869 1 0 spiny-headed seasnake Gulf of Thailand, South China Sea, the Strait of Taiwan, and the coasts of Guangdong, Indonesia, the Philippines, New Guinea, New Caledonia, Australia (Northern Territory, Queensland, Western Australia)
Acanthophis Daudin, 1803 7 0 death adders Australia, New Guinea, Indonesia (Seram and Tanimbar)
Aipysurus Lacépède, 1804 7 1 olive sea snakes Timor Sea, South China Sea, Gulf of Thailand, and coasts of Australia (Northern Territory, Queensland, Western Australia), New Caledonia, Loyalty Islands, southern New Guinea, Indonesia, western Malaysia and Vietnam
Aspidelaps Fitzinger, 1843 2 4 shieldnose cobras South Africa (Cape Province, Transvaal), Namibia, southern Angola, Botswana, Zimbabwe, Mozambique
Aspidomorphus Fitzinger, 1843 3 3 collared adders New Guinea
Astrotia Fischer, 1855 1 0 Stokes' sea snake Coastal areas from west India and Sri Lanka through Gulf of Thailand to China Sea, west Malaysia, Indonesia east to New Guinea, north and east coasts of Australia, the Philippines
Austrelaps Worrell, 1963 3 0 copperheads Australia (South Australia, New South Wales, Victoria, Tasmania)
Boulengerina Dollo, 1886 2 1 water cobras Cameroon, Gabon, the Democratic Republic of the Congo, Congo, the Central African Republic, Tanzania, Equatorial Guinea, Rwanda, Burundi, Zambia
Bungarus Daudin, 1803 12 4 kraits India (incl. Andaman Island), Myanmar, Nepal, Vietnam, Afghanistan, Pakistan, Sri Lanka, Bangladesh, Cambodia, Indonesia (Java, Sumatra, Bali, Sulawesi), Peninsular Malaysia, Singapore, Taiwan, Thailand
Cacophis Günther, 1863 4 0 rainforest crowned snakes Australia (New South Wales, Queensland)
Calliophis Gray, 1834 8 11 Oriental coral snakes India, Bangladesh, Sri Lanka, Nepal, Indonesia, Cambodia, Malaysia, Singapore, Thailand, Burma, Brunei, the Philippines, Vietnam, Laos, southern China, Japan (Ryukyu Islands), Taiwan
Demansia Gray, 1842 9 2 whipsnakes New Guinea, continental Australia
Dendroaspis Schlegel, 1848 4 1 mambas Kenya, Tanzania, Mozambique, Malawi, Zimbabwe, South Africa, Ghana, Togo, Benin, Nigeria, Cameroon, Guinea, Gabon, Principe (Gulf of Guinea), Central African Republic, Democratic Republic of the Congo, Congo, Uganda, Rwanda, Burundi, Equatorial Guinea, Angola, Sudan, Botswana, Burkina Faso, Eritrea, Senegal, Mali, Ethiopia, Ivory Coast, Namibia, Somalia, Swaziland, Zambia, Gambia, Guinea Bissau, Liberia, Ivory Coast, Sierra Leone
Denisonia Krefft, 1869 2 0 ornamental snakes Central Queensland and central northern New South Wales, Australia
Drysdalia Worrell, 1961 3 0 southeastern grass snakes Southern Australia (Western Australia, South Australia, Victoria, Tasmania, New South Wales)
Echiopsis Fitzinger, 1843 1 0 bardick Southern Australia (Western Australia, South Australia, Victoria, New South Wales)
Elapognathus Boulenger, 1896 2 0 southwestern grass snakes Western Australia
Elapsoidea Bocage, 1866 10 7 African or venomous garter snakes (not related to North American garter snakes, which are nonvenomous) Senegal, South Africa, Mozambique, Namibia, Botswana, Zimbabwe, Swaziland, Gambia, Angola, Benin, Burkina Faso, Cameroon, Central African Republic, Chad, Ghana, Ivory Coast, Malawi, Mali, Mauritania, Niger, Nigeria, Uganda, Senegal, Sudan, Tanzania, Togo, the Democratic Republic of the Congo, Congo, Zambia, Kenya, north Burundi, Rwanda, Ethiopia, Uganda, Somalia
Emydocephalus Krefft, 1869 2 0 turtlehead sea snakes The coasts of Timor (Indonesian Sea), New Caledonia, Australia (Northern Territory, Queensland, Western Australia), and in the Southeast Asian Sea along the coasts of China, Taiwan, Japan, and the Ryukyu Islands
Enhydrina Gray, 1849 2 0 beaked sea snakes In the Persian Gulf (Oman, United Arab Emirates, etc.), south to the Seychelles and Madagascar,

Southeast Asian Sea (Pakistan, India, Bangladesh, Myanmar, Thailand, Vietnam), Australia (North Territory, Queensland), New Guinea and Papua New Guinea

Ephalophis M.A. Smith, 1931 1 0 Grey's mudsnake Northwestern Australia
Furina A.M.C. Duméril, 1853 5 0 pale-naped snakes Mainland Australia, southern New Guinea, Aru Islands
Hemachatus Fleming, 1822 1 0 spitting cobra South Africa, Zimbabwe, Lesotho, Swaziland
Hemiaspis Fitzinger, 1861 2 0 swamp snakes Eastern Australia (New South Wales, Queensland)
Hemibungarus W. Peters, 1862 1 2 Asian coral snakes Taiwan, Japan (Ryukyu Islands)
Homoroselaps Jan, 1858 2 0 harlequin snakes South Africa
Hoplocephalus Wagler, 1830 3 0 broad-headed snakes Eastern Australia (New South Wales, Queensland)
Hydrelaps Boulenger, 1896 1 0 Port Darwin mudsnake Northern Australia, southern New Guinea
Hydrophis Latreille In Sonnini & Latreille, 1801 34 3 sea snakes Indoaustralian and Southeast Asian waters.[5]
Kerilia Gray, 1849 1 0 Jerdon's sea snake Southeast Asian waters[5]
Kolpophis M.A. Smith, 1926 1 0 bighead sea snake Indian Ocean[5]
Lapemis Gray, 1835 1 1 Shaw's sea snake Persian Gulf to Indian Ocean, South China Sea, Indo-Australian archipelago and the western Pacific[5]
Laticauda Laurenti, 1768 5 0 sea kraits Southeast Asian and Indo-Australian waters
Leptomicrurus K.P. Schmidt, 1937 4 2 blackback coral snake Northern South America
Loveridgelaps McDowell, 1970 1 0 Solomons small-eyed snake Solomon Islands
Micropechis Boulenger, 1896 1 0 New Guinea small-eyed snake New Guinea
Micruroides K.P. Schmidt, 1928 1 2 Western coral snakes USA (Arizona, southwestern New Mexico), Mexico (Sonora, Sinaloa)
Micrurus Wagler, 1824 69 54 coral snakes Southern North America, South America
Naja Laurenti, 1768 23 3 cobras Africa, Asia
Notechis Boulenger, 1896 2 0 tiger snakes Southern Australia, including many offshore islands
Ogmodon W. Peters, 1864 1 0 bola Fiji
Ophiophagus Günther, 1864 1 0 king cobra Bangladesh, Myanmar, Cambodia, China, India, Andaman Islands, Indonesia, Laos, Thailand, Vietnam, western Malaysia, the Philippines, Sri Lanka
Oxyuranus Kinghorn, 1923 3 2 taipans Australia, New Guinea
Parahydrophis Burger & Natsuno, 1974 1 0 Northern mangrove sea snake Northern Australia, southern New Guinea
Paranaja Loveridge, 1944 1 2 many-banded snakes West/central Democratic Republic of the Congo, Congo, Cameroon
Parapistocalamus Roux, 1934 1 0 Hediger's snake Bougainville Island, Solomons
Paroplocephalus Keogh, Scott, and Scanlon, 2000 1 0 Lake Cronin snake Western Australia
Pelamis Daudin, 1803 1 0 yellow-bellied sea snake Indian and Pacific Oceans
Praescutata viperina Wall, 1921 1 0 viperine sea snake Persian Gulf, Indian Ocean, South Chinese Sea northeast to coastal region of Fujian and Strait of Taiwan
Pseudechis Wagler, 1830 7 0 black snakes (and king brown) Australia
Pseudohaje Günther, 1858 2 0 tree cobras Angola, Burundi, Cameroon, Central African Republic, Democratic Republic of the Congo, Congo, Gabon, Ghana, Kenya, Nigeria, Rwanda, Uganda, Sierra Leone, Liberia, Ivory Coast, Togo, Nigeria
Pseudonaja Günther, 1858 8 2 venomous brown snakes (and dugites) Australia
Rhinoplocephalus F. Müller, 1885 6 0 Australian small-eyed snakes Southern and eastern Australia, southern New Guinea
Salomonelaps McDowell, 1970 1 0 Solomons coral snake Solomon Islands
Simoselaps Jan, 1859 13 3 Australian coral snakes Mainland Australia
Suta Worrell, 1961 10 2 hooded snakes (and curl snake) Australia
Thalassophis P. Schmidt, 1852 1 0 anomalous sea snake South Chinese Sea (Malaysia, Gulf of Thailand), Indian Ocean (Sumatra, Java, Borneo)
Toxicocalamus Boulenger, 1896 11 0 New Guinea forest snakes New Guinea (and nearby islands)
Tropidechis Günther, 1863 2 0 rough-scaled snake Eastern Australia
Vermicella Gray in Günther, 1858 5 0 bandy-bandies Australia
Walterinnesia Lataste, 1887 2[6] 0 black desert cobra Egypt, Israel, Lebanon, Syria, Jordan, Iraq, Iran, Kuwait, Saudi Arabia, Turkey [7]

* Not including the nominate subspecies

Taxonomy

The table above lists all of the elapid genera and no subfamilies. In the past, many subfamilies were recognized, or have been suggested for the Elapidae, including the Elapinae, Hydrophiinae (sea snakes), Micrurinae (coral snakes), Acanthophiinae (Australian elapids) and the Laticaudinae (sea kraits). Currently, none are universally recognized. Good molecular evidence via karyotyping, protein electrophoretic analysis, immunological distance, DNA sequence analysis, and so on, suggests reciprocal monophyly of two groups:

  • African, Asian, and New World Elapinae
  • Australasian and marine Hydrophiinae

Thus, the Australian terrestrial elapids are technically 'hydrophiines', although they are not sea snakes. It is believed that the Laticauda and the 'true sea snakes' evolved separately from Australasian land snakes. Asian cobras, coral snakes, and American coral snakes also appear to be monophyletic, while African cobras do not.[8][9]

The type genus for the Elapidae was originally Elaps, but the group was moved to another family. In contrast to what is typical of botany, the Elapidae family was not renamed. In the meantime, Elaps was renamed Homoroselaps and moved back to the Elapidae. However, Nagy et al. (2005) regard it as a sister taxon to Atractaspis, which shouldhave been assigned to the Atractaspididae.

See also

References

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  2. 2.0 2.1 2.2 2.3 2.4 Lua error in package.lua at line 80: module 'strict' not found.
  3. 3.0 3.1 Elapidae at the Reptarium.cz Reptile Database. Accessed 3 November 2008.
  4. WCH Clinical Toxinology Resources: Oxyuranus microlepidotus
  5. 5.0 5.1 5.2 5.3 The Hydrophiidae at Cyberlizard's home pages. Accessed [12 August] [2007].
  6. Nilson, G. & N. Rastegar-Pouyani (2007) Walterinnesia aegyptia Lataste, 1887 (Ophidia: Elapidae) and the status of Naja morgani Mocquard 1905. Russian Journal of Herpetology, 14: 7-14.
  7. Ugurtas, I. H., T. J. Papenfuss and N. L. Orlov. 2001. New record of Walterinnesia aegyptia Lataste, 1887 (Ophidia: Elapidae: Bungarinae) in Turkey. Russian Journal of Herpetology. 8(3):239-245.
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