Elapidae

Elapidae
Egyptian cobra, Naja haje
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Suborder: Serpentes
Infraorder: Alethinophidia
Family: Elapidae
F. Boie, 1827

Elapidae (Greek éllops = sea-fish) [1] is a family of venomous snakes found in tropical and subtropical regions around the world, terrestrially in Asia and North America and aquatically in the Pacific and Indian Oceans. Elapid snakes exist in a wide range of sizes, from 18 cm species of Drysdalia to the 6 m king cobra, and are characterized by hollow, fixed fangs through which they inject venom. Currently, 61 genera that include 325 species are recognized.[2] Snakes of this family are known as elapine.

Contents

Description

All elapids have a pair of proteroglyphous fangs that are 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 that is 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. There are exceptions to all these generalizations: 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, Vermicella) in either humid or arid environments. Some species have very generalised diets, but many taxa have narrow prey preferences (stenophagy) and correlated morphological specialisations, e.g. for feeding on other snakes, elongate burrowing lizards, squamate eggs, mammals, birds, frogs, fish, etc.

Sea snakes, which are also elapids, 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, ventral scales are much reduced in size, their nostrils are located dorsally (no internasal scales) and 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 least well-adapted to an aquatic life. They spend much of their time on land, where they lay their eggs. They have wide ventral scales, the tail is not as well-developed for swimming, and their nostrils are separated by internasal scales.

Dentition

The fangs are the first two teeth on each maxillary bone, which are enlarged and hollow, and usually only one is in place on each side at any time. The maxilla is intermediate in 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; in the longest-fanged elapids (e.g. Acanthophis, Oxyuranus) it is common for the fangs to pierce right through the intermandibular skin, which does not seem to endanger the snake. The fangs are usually below the front edge of the eye and are angled backwards; due to this construction, most elapids must actually bite to envenomate. This action is therefore not as quick as with the viperids, that can envenomate with only a quick, stabbing motion. Some elapids (Acanthophis, Oxyuranus, and especially Dendroaspis) have long fangs on quite mobile maxillae (the prefrontal and ectopterygoid contacts are nearly as close together as in viperids), and can therefore make very fast stabbing strikes like viperids. A few species are capable of spraying their venom from forward-facing holes at the tips of their fangs as a means of defense. Elapids use their venom both to immobilize their prey and in self defense.

Geographic range

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

All elapids are venomous and many are potentially deadly. The venoms are mostly neurotoxic and are considered more dangerous than the mainly proteolytic venoms of vipers. Many elapids are also large in size and can deliver a large quantity of potent venom. For example, the Asiatic king cobra, the African black mamba and the Australian taipan.

Genera

Genus[2] Taxon author[2] Species[2] Subsp.*[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, 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 (North Territory, Queensland, West 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 lineaticollis Fitzinger, 1843 3 3 Collared adders New Guinea.
Astrotia Fischer, 1855 1 0 Stoke's 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, Philippines
Austrelaps Worrell, 1963 3 0 Copperheads Australia (South Australia, New South Wales, Victoria, Tasmania)
Boulengerina Dollo, 1886 2 1 Water cobras Cameroon, Gabon, Democratic Republic of the Congo, Congo, 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, 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)
Edichnopsis Young, 2009 2 0 Sebastian cobra (separate species) India, Sri Lanka, Nepal, Bhutan, South east China (including Tibet and Hong Kong), Northern Burma, Laos and Vietnam. Also found in Southern Cambodia.
Elapognathus Boulenger, 1896 2 0 Southwestern grass snakes Western Australia.
Elapsoidea Bocage, 1866 10 7 African or venomous garter snakes (unrelated to North American nonvenomous garter snakes) 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, 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 (North Territory, Queensland, West Australia), and in the Southeast Asian Sea along the coasts of China, Taiwan, Japan, and the Ryukyu Island
Enhydrina Gray, 1849 2 0 Beaked sea snakes In the Persian Gulf (Oman, United Arab Emirates, etc.), south to the Seychelles and Madagascar,

SE 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 Duméril, 1853 3 0 Pale-naped snakes Mainland Australia.
Glyphodon Günther, 1858 2 0 Brown-headed snakes Australia (Queensland), New Guinea.
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 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.[4]
Kerilia Gray, 1849 1 0 Jerdon's sea snake Southeast Asian waters[4]
Kolpophis M.A. Smith, 1926 1 0 Bighead sea snake Indian Ocean[4]
Lapemis Gray, 1835 1 1 Shaw's sea snake Persian Gulf to Indian Ocean, South China Sea, Indo-Australian archipelago and the western Pacific[4]
Laticauda Laurenti, 1768 5 0 Sea kraits Southeast Asian and Indoaustralian waters.
Leptomicrurus 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 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 Peters, 1864 1 0 Bola Fiji
Ophiophagus Günther, 1864 1 0 King cobra Bangladesh, Myanmar, Cambodia, China, India, Andaman Islands, Indonesia, Laos, Thailand, Vietnam, west Malaysia, Philippines
Oxyuranus Kinghorn, 1923 2 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 Forest 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 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
Sinomicrurus (Calliophis) macclellandi Slowinski et al., 2001 5 4 MacClelland’s (Asian) coral snake India, Myanmar, Vietnam, China, Taiwan, Japan
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 9 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[5] 0 Black desert cobra Egypt, Israel, Lebanon, Syria, Jordan, Iraq, Iran, Kuwait, Saudi Arabia, Turkey [6]

*) 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. There is now good molecular evidence for reciprocal monophyly of two groups: the African, Asian and New World Elapinae, and Australasian and marine Hydrophiinae. Thus, the Australian terrestrial elapids are 'hydrophiines', though not sea snakes, while it is believed that Laticauda and the 'true sea snakes' evolved separately from among the Australasian land-snakes. Asian cobras, coral snakes, and American coral snakes also appear to be monophyletic, while African cobras do not.[7]

The type genus for the Elapidae was originally Elaps, but that group was moved to another family. In contrast to what usually happens in 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 should therefore have been assigned to the Atractaspididae.

See also

References

  1. "Definition of 'elapid'.". dictionary.com. http://dictionary.reference.com/browse/elapid?r=75&src=ref&ch=dic. Retrieved 2009-07-13. 
  2. 2.0 2.1 2.2 2.3 2.4 "Elapidae". Integrated Taxonomic Information System. http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=174348. Retrieved 27 November 2006. 
  3. 3.0 3.1 Elapidae at the Reptarium.cz Reptile Database. Accessed 3 November 2008.
  4. 4.0 4.1 4.2 4.3 The Hydrophiidae at Cyberlizard's home pages. Accessed [12 August] [2007].
  5. 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.
  6. 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.
  7. Slowinski, J. B. and Keogh J. S. (April 2000). "Phylogenetic Relationships of Elapid Snakes Based on Cytochrome b mtDNA Sequences". Molecular Phylogenetics and Evolution 15 (1): 157–164. doi:10.1006/mpev.1999.0725. PMID 10764543. http://www.ingentaconnect.com/content/ap/fy/2000/00000015/00000001/art00725.