Naja

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Naja (Uraeus)
)
This article is about a genus of venomous snakes. For other uses, see Naja (disambiguation).

Naja
Temporal range: Miocene–Holocene
Indian cobra (Naja naja), species typica of the genus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Suborder: Serpentes
Family: Elapidae
Genus: Naja
Laurenti, 1768
Type species
Coluber naja

Linnaeus, 1758

Naja is a

Ophiophagus (the king cobra/hamadryad).[2][3]

Until recently, the genus Naja had 20 to 22

Paranaja with Naja. According to that revision, the genus Naja now includes 38 species.[7]

Indian cobra (Naja naja)
Dissected head of Naja melanoleuca showing (A) the fangs and (B) the venom gland

Etymology

The origin of this genus name is from the

cognate with English "snake", Germanic: *snēk-a-, Proto-IE: *(s)nēg-o-,[8] but Manfred Mayrhofer calls this etymology "not credible", and suggests a more plausible etymology connecting it with Sanskrit nagna, "hairless" or "naked".[9]

Description

Naja species vary in length and most are relatively slender-bodied snakes. Most species are capable of attaining lengths of 1.84 m (6.0 ft). Maximum lengths for some of the larger species of cobras are around 3.1 m (10 ft), with the

Naja oxiana) have some degree of adaptation to spitting.[11]

Venom

All species in the genus Naja are capable of delivering a fatal bite to a human. Most species have strongly

cytotoxic features which cause swelling and necrosis, and have a significant anticoagulant effect. Some also have cardiotoxic
components to their venom.

Several Naja species, referred to as

front fangs, instead of ejecting venom downward through an elongated discharge orifice (similar to a hypodermic needle
), have a shortened, rounded opening in the front surface, which ejects the venom forward, out of the mouth. While typically referred to as "spitting", the action is more like squirting. The range and accuracy with which they can shoot their venom varies from species to species, but it is used primarily as a defense mechanism. The venom has little or no effect on unbroken skin, but if it enters the eyes, it can cause a severe burning sensation and temporary or even permanent blindness if not washed out immediately and thoroughly.

A recent study[12] showed that all three spitting cobra lineages have evolved higher pain-inducing activity through increased phospholipase A2 levels, which potentiate the algesic action of the cytotoxins present in most cobra venoms. The timing of the origin of spitting in African and Asian Naja species corresponds to the separation of the human and chimpanzee evolutionary lineages in Africa and the arrival of Homo erectus in Asia. The authors therefore hypothesise that the arrival of bipedal, tool-using primates may have triggered the evolution of spitting in cobras.

The Caspian cobra (N. oxiana) of Central Asia is the most venomous Naja species. According to a 2019 study by Kazemi-Lomedasht et al, the murine LD50 via intravenous injection (IV) value for Naja oxiana (Iranian specimens) was estimated to be 0.14 mg/kg (0.067-0.21 mg/kg)[13] more potent than the sympatric Pakistani Naja naja karachiensis and Naja naja indusi found in far north and northwest India and adjacent Pakistani border areas (0.22 mg/kg), the Thai Naja kaouthia (0.2 mg/kg), and Naja philippinensis at 0.18 mg/kg (0.11-0.3 mg/kg).[14] Latifi (1984) listed a subcutaneous value of 0.2 mg/kg (0.16-0.47 mg/kg) for N. oxiana.[15] The crude venom of N. oxiana produced the lowest known lethal dose (LCLo) of 0.005 mg/kg, the lowest among all cobra species ever recorded, derived from an individual case of envenomation by intracerebroventricular injection.[16] The Banded water cobra's LD50 was estimated to be 0.17 mg/kg via IV according to Christensen (1968).[17][18] The Philippine cobra (N. philippinensis) has an average murine LD50 of 0.18 mg/kg IV (Tan et al, 2019).[14] Minton (1974) reported 0.14 mg/kg IV for the Philippine cobra.[19][20][21] The Samar cobra (Naja samarensis), another cobra species endemic to the southern islands of the Philippines, is reported to have a LD50 of 0.2 mg/kg,[22] similar in potency to the monocled cobras (Naja kaouthia) found only in Thailand and eastern Cambodia, which also have a LD50 of 0.2 mg/kg. The spectacled cobras that are sympatric with N. oxiana, in Pakistan and far northwest India, also have a high potency of 0.22 mg/kg.[14][23]

Other highly venomous species are the 'forest cobras' and/or 'water cobras' (Boulengerina subgenus) are also highly venomous. The murine intraperitoneal

subcutaneous injection (SC).[25] The LD50 of the Cape cobra (N. nivea) according to Minton, 1974 was 0.35 mg/kg (IV) and 0.4 mg/kg (SC).[19][26] The Senegalese cobra (N. senegalensis) has a murine LD50 of 0.39 mg/kg (Tan et al, 2021) via IV.[27] The Egyptian cobra (N. haje) of Ugandan locality had an IV LD50 of 0.43 mg/kg (0.35–0.52 mg/kg).[28]

The Naja species are a medically important group of snakes due to the number of bites and fatalities they cause across their geographical range. They range throughout

Dendroaspis (mambas) and Bungarus (kraits), with mambas tending to almost always envenomate and kraits tending to envenomate more often than they attempt 'sham strikes.[30]

Many factors influence the differences in cases of fatality among different species within the same genus. Among cobras, the cases of fatal outcome of bites in both treated and untreated victims can be quite large. For example, mortality rates among untreated cases of envenomation by the cobras as a whole group ranges from 6.5–10% for N kaouthia.

endotracheal intubation is required. As a result, mortality among those treated for N. oxiana envenomation is still relatively high (up to 30%) compared to all other species of cobra (<1%).[15]

Taxonomy

Naja
(Naja)

Naja (Naja) naja

Naja (Naja) kaouthia

Naja (Naja) atra

Naja (Naja) sagittifera

Naja (Naja) oxiana

Naja (Naja) sputatrix

Naja (Naja) samarensis

Naja (Naja) philippinensis

Naja (Naja) mandalayensis

Naja (Naja) sumatrana

Naja (Naja) siamensis

(
Afronaja
)

Naja (Afronaja) pallida

Naja (Afronaja) nubiae

Naja (Afronaja) katiensis

Naja (Afronaja) nigricollis

(Boulengerina)

Naja (Boulengerina) multifasciata

Naja (Boulengerina) savannula

Naja (Boulengerina) subfulva

Naja (Boulengerina) guineensis

Naja (Boulengerina) peroescobari

Naja (Boulengerina) melanoleuca

(
Uraeus
)

Naja (Uraeus) nivea

Naja (Uraeus) senegalensis

Naja (Uraeus) haje

Naja (Uraeus) arabica

Naja (Uraeus) annulifera

Naja (Uraeus) anchietae

The genus contains several species complexes of closely related and often similar-looking species, some of them only recently described or defined. Several recent taxonomic studies have revealed species not included in the current listing in ITIS:[5][37]

  • N. annulifera by Broadley (1995). It is regarded as a full species by Broadley and Wüster (2004).[38][39]
  • Naja arabica Scortecci, 1932, the Arabian cobra, has long been considered a subspecies of N. haje, but was recently raised to the status of species.[40]
  • Naja ashei Broadley and Wüster, 2007, Ashe's spitting cobra, is a newly described species found in Africa and also a highly aggressive snake; it can spit a large amount of venom.[41][42]
  • Naja nigricincta Bogert, 1940, was long regarded as a subspecies of N. nigricollis, but was recently found to be a full species (with N. n. woodi as a subspecies).[43][44]
  • Naja senegalensis Trape et al., 2009, is a new species encompassing what were previously considered to be the West African savanna populations of N. haje.[40]
  • Naja peroescobari Ceríaco et al. 2017, is a new species encompassing what was previously considered the São Tomé population of N. melanoleuca.[45]
  • Naja guineensis Broadley et al., 2018, is a new species encompassing what were previously considered to be the West African forest populations of N. melanoleuca.[7]
  • Naja savannula Broadley et al., 2018, is a new species encompassing what were previously considered to be the West African savanna populations of N. melanoleuca.[7]
  • Naja subfulva Laurent, 1955, previously regarded as a subspecies of N. melanoleuca, was recently recognized as a full species.[7]

Two recent molecular phylogenetic studies have also supported the incorporation of the species previously assigned to the genera

Boulengerina and Paranaja into Naja, as both are closely related to the forest cobra (Naja melanoleuca).[43][46] In the most comprehensive phylogenetic study to date, 5 putative new species were initially identified, of which 3 have since been named.[4]

The controversial amateur herpetologist Raymond Hoser proposed the genus Spracklandus for the African spitting cobras.[47] Wallach et al. suggested that this name was not published according to the Code and suggested instead the recognition of four subgenera within Naja: Naja for the Asiatic cobras, Boulengerina for the African forest, water and burrowing cobras, Uraeus for the Egyptian and Cape cobra group and Afronaja for the African spitting cobras.[6] International Commission on Zoological Nomenclature issued an opinion that it “finds no basis under the provisions of the Code for regarding the name Spracklandus as unavailable”.[48]

Asiatic cobras are believed to further be split into two groups of southeastern Asian cobras (N. siamensis, N. sumatrana, N. philippinensis, N. samarensis, N. sputatrix, and N. mandalayensis) and western and northern Asian cobras (N. oxiana, N. kaouthia, N. sagittifera, and N. atra) with Naja naja serving as a basal lineage to all species.[49]

Species

Image[5] Species[5] Authority[5] Subsp.*[5] Common name Geographic range
N. anchietae Bocage, 1879 0 Anchieta's cobra (Angolan Cobra) Angola, Botswana, Namibia, Zambia, eastern Zimbabwe
N. annulata (Buchholz and Peters, 1876) 1
Banded water cobra
 Cameroon, the Central African Republic, the Democratic Republic of the Congo (Zaire), the Republic of Congo, Equatorial Guinea, Gabon, Rwanda, and the province of Cabinda in Angola
N. annulifera Peters, 1854 0 Snouted cobra Botswana, Malawi, Mozambique, South Africa, Swaziland, Zambia, Zimbabwe
†N. antiqua Rage, 1976 0 Miocene-aged strata of Morocco
N. arabica Scortecci, 1932 0 Arabian cobra Oman, Saudi Arabia, Yemen
N. ashei Wüster and Broadley, 2007 0
Ashe's spitting cobra
(giant spitting cobra) 		southern Ethiopia, Kenya, Somalia, eastern Uganda
N. atra
Cantor
, 1842 		0 Chinese cobra southern China, northern Laos, Taiwan, northern Vietnam
N. christyi (Boulenger, 1904) 0
Congo water cobra
 the Democratic Republic of the Congo (Zaire), the Republic of Congo, and the province of Cabinda in Angola
N. fuxi

Shi, Vogel, Chen, & Ding, 2022

0 Brown banded cobra China, Myanmar, Laos, Thailand, Vietnam
N. guineensis Broadley, Trape, Chirio, Ineich &Wüster, 2018 0 Black forest cobra Ghana, Guinea, Guinea-Bissau, the Ivory Coast, Liberia, Sierra Leone, Togo
N. haje Linnaeus, 1758 0 Egyptian cobra Tanzania, Kenya, Somalia, Ethiopia, Uganda, South Sudan, Sudan, Cameroon, Nigeria, Niger, Burkina Faso, Mali, Senegal, Mauritania, Morocco, Algeria, Tunisia, Libya, and Egypt
†N. iberica Szyndlar, 1985 Miocene-aged strata of Spain
N. kaouthia
Lesson
, 1831 		0 Monocled cobra Bangladesh, Bhutan, Burma, Cambodia, southern China, eastern India, Laos, northwestern Malaysia, Nepal, Thailand, southeastern Tibet, Vietnam
N. katiensis Angel, 1922 0 Mali cobra (Katian spitting cobra) Benin, Burkina Faso, Cameroon, Ghana, Guinea, the Ivory Coast, Mali, Gambia, Mauritania, Niger, Nigeria, Senegal, Togo
N. mandalayensis Slowinski & Wüster, 2000 0 Mandalay spitting cobra (Burmese spitting cobra) Myanmar (Burma)
N. melanoleuca Hallowell, 1857 0 Central African forest cobra Angola, Benin, Cameroon, the Central African Republic, the Republic of Congo, the Democratic Republic of the Congo (Zaire), Equatorial Guinea, Gabon, Nigeria
N. mossambica Peters, 1854 0 Mozambique spitting cobra extreme southeastern Angola, Botswana, Malawi, Mozambique, Somalia, northeastern Namibia, South Africa, Swaziland, Tanzania (including Pemba Island), Zambia, Zimbabwe
N. multifasciata Werner, 1902 0
Many-banded cobra
 Cameroon, Congo, the Democratic Republic of the Congo (Zaire), Gabon
N. naja (Linnaeus, 1758) 0 Indian cobra (spectacled cobra) Bangladesh, Bhutan, India, Nepal, Pakistan, Sri Lanka
Naja nana, Congo dwarf water cobra N. nana Collet & Trape, 2020 0 Dwarf water cobra Democratic Republic of Congo
N. nigricincta Bogert, 1940 1
Zebra spitting cobra
 Angola, Namibia, South Africa
N. nigricollis Reinhardt, 1843 0 Black-necked spitting cobra Angola, Benin, Burkina Faso, Burundi, Cameroon, the Central African Republic, Chad, the Democratic Republic of the Congo (Zaire) (except in the central region), Congo, Ethiopia, Gabon, Gambia, Ghana, Guinea-Bissau, Guinea, the Ivory Coast, Kenya, Liberia, Mali, Mauritania, Namibia, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Sudan, Tanzania, Somalia, Togo, Uganda, Zambia
N. nivea (Linnaeus, 1758) 0 Cape cobra (yellow cobra) Botswana, Lesotho, Namibia, South Africa
N. nubiae Wüster & Broadley, 2003 0 Nubian spitting cobra Chad, Egypt, Eritrea, Niger, Sudan
N. oxiana (Eichwald, 1831) 0 Caspian cobra Afghanistan, northwestern India, Iran, Kyrgyzstan, Pakistan, Tajikistan, Turkmenistan, Uzbekistan
N. pallida Boulenger, 1896 0 Red spitting cobra Djibouti, Ethiopia, Kenya, Somalia, Tanzania
N. peroescobari Ceríaco, Marques, Schmitz & Bauer, 2017 0 São Tomé forest cobra, cobra preta São Tomé and Príncipe (São Tomé)
N. philippinensis Taylor, 1922 0 Philippine cobra the Philippines (Luzon, Mindoro)
†N. romani (Hofstetter, 1939) 0 † Miocene-aged strata of France, Germany, Austria, Russia, Hungary, Greece and Ukraine.[50]
N. sagittifera Wall, 1913 0 Andaman cobra India (the Andaman Islands)
N. samarensis Peters, 1861 0 Samar cobra the Philippines (Mindanao, Bohol, Leyte, Samar, Camiguin)
N. savannula Broadley, Trape, Chirio & Wüster, 2018 0 West African banded cobra Benin, Burkina Faso, Cameroon, Chad, Gambia, Ghana, Guinea, the Ivory Coast, Mali, Niger, Nigeria, Senegal, Togo
N. senegalensis Trape, Chirio & Wüster, 2009 0 Senegalese cobra Benin, Burkina Faso, Ghana, Guinea, Mali, Niger, Nigeria, Senegal
N. siamensis Laurenti, 1768 0 Indochinese spitting cobra Cambodia, Laos, Thailand, Vietnam
N. sputatrix F. Boie, 1827 0 Javan spitting cobra Indonesia (Java, the Lesser Sunda Islands, East Timor)
N. subfulva Laurent, 1955 0 Brown forest cobra Angola, Burundi, Cameroon, the Central African Republic, Chad, the Republic of Congo, the Democratic Republic of the Congo (Zaire), Ethiopia, Kenya, Malawi, Mozambique, Nigeria, Rwanda, Somalia, South Africa, South Sudan, Tanzania, Uganda, Zambia, Zimbabwe
N. sumatrana Müller, 1887 0 Equatorial spitting cobra Brunei, Indonesia (Sumatra, Borneo, Bangka, Belitung), Malaysia, the Philippines (Palawan), southern Thailand, Singapore
  • Not including the nominate subspecies

† Extinct
T Type species[2]

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