Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Cladus: Craniata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Lepidosauromorpha
Superordo: Lepidosauria
Ordo: Squamata
Subordo: Serpentes
Infraordo: Caenophidia
Superfamilia: Elapoidea
Familia: Elapidae
Subfamilia: Elapinae
Genus: Naja
Subgenera: Naja (Naja) - Naja (Afronaja) - Naja (Boulengerina) - Naja (Uraeus)
Species:
N. (Naja): N. atra – N. kaouthia – N. mandalayensis – N. naja – N. oxiana – N. philippinensis – N. sagittifera – N. samarensis – N. siamensis – N. sputatrix – N. sumatrana
N. (Afronaja): N. ashei – N. katiensis – N. mossambica – N. nigricincta – N. nigricollis – N. nubiae – N. pallida
N. (Boulengerina): N. annulata – N. christyi – N. guineensis – N. melanoleuca – N. nana – N. peroescobari – N. savannula – N. subfulva – N. multifasciata
N. (Uraeus): N. arabica – N. anchietae – N. annulifera – N. haje – N. nivea – N. senegalensis
Name
Naja Laurenti, 1768
Afronaja Wallach, Wüster & Broadley, 2009
Synonym: Spracklandus Hoser, 2009: 8 (rejected name)
Gender: [feminine]
Original status: valid genus
Type species: Naja lutescens Laurenti 1768
References
Kaiser, H., Crother, B.I., Kelly, C.M.R., Luiselli, L., O’Shea, M., Ota, H., Passos, P., Schleip, W.D. & Wüster, W. 2013. Best practices: in the 21st Century, taxonomic decisions in herpetology are acceptable only when supported by a body of evidence and published via peer-review. Herpetological Review 44(1): 8–23. PDFOpen access Reference page.
Trape, J.-F., Chirio, L., Broadley, D.G. & Wüster, W. 2009. Phylogeography and systematic revision of the Egyptian cobra (Serpentes: Elapidae: Naja haje) species complex, with the description of a new species from West Africa. Zootaxa, 2236: 1–25. Abstract & excerpt Reference page.
Wallach, V., Wüster, W. & Broadley, D.G. 2009. In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae). Zootaxa 2236: 26–36. Abstract & excerpt PDF Reference page.
Wüster, W., Crookes, S., Ineich, I., Mané, Y., Pook, C.E., Trape, J.-F. & Broadley, D.G. 2007. The phylogeny of cobras inferred from mitochondrial DNA sequences: evolution of venom spitting and the phylogeography of the African spitting cobras (Serpentes: Elapidae: Naja nigricollis complex). Molecular Phylogenetics and Evolution 45(2): 437–453. DOI: 10.1016/j.ympev.2007.07.021 Paywall Reference page.
Wüster, W., Chirio, L., Trape, J.-F., Ineich, I., Jackson, K., Greenbaum, E., Barron, C., Kusamba, C., Nagy, Z.T., Storey, R., Hall, C., Wüster, C.E., Barlow, A. & Broadley, D.G.† 2018. Integration of nuclear and mitochondrial gene sequences and morphology reveals unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae). Zootaxa 4455(1): 68–98. DOI: 10.11646/zootaxa.4455.1.3. Reference page.
Vernacular names
беларуская: Сапраўдныя кобры
Deutsch: Echte Cobras
English: Cobras
suomi: Kobrat
日本語: フードコブラ属
lietuvių: Kobros
Diné bizaad: Tłʼiish bikʼós niteelígíí
ไทย: งูเห่า
Naja is a genus of venomous elapid snakes known as cobras ("true cobras"). Members of the genus Naja are the most widespread and the most widely recognized as "true" cobras. Various species occur in regions throughout Africa, Southwest Asia, South Asia, and Southeast Asia. Several other elapid species are also called "cobras", such as the king cobra (Ophiophagus hannah) and the rinkhals (Hemachatus haemachatus), but neither are true cobras. They are not true cobras in that they do not belong to the genus Naja, but instead each belong to monotypic genera Hemachatus (the rinkhals)[1] and Ophiophagus (the king cobra/hamadryad).[2][3]
Until recently, the genus Naja had 20 to 22 species, but it has undergone several taxonomic revisions in recent years, so sources vary greatly.[4][5] Wide support exists, though, for a 2009 revision[6] that synonymised the genera Boulengerina and 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 Sanskrit nāga (with a hard "g") meaning "snake". Some hold that the Sanskrit word is cognate with English "snake", Germanic: *snēk-a-, Proto-IE: *(s)nēg-o-,[8] but this is unlikely. Mayrhofer calls this etymology "unglaubhaft ", "not credible", and suggests a more plausible etymology connecting it with Sanskrit nagna, "hairless, 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 forest cobra arguably being the longest species.[10] All have a characteristic ability to raise the front quarters of their bodies off the ground and flatten their necks to appear larger to a potential predator. Fang structure is variable, all species except the Indian cobra (Naja naja) and Caspian cobra (Naja oxiana) have some degree of adaptation to spitting.[11]
Venom
Most venomous Naja species Rank Species LD50 SC
1 N. oxiana 0.10 mg/kg[12]
2 N. philippinensis 0.14 mg/kg[13][14]
3 N. samarensis 0.21 mg/kg[15]
4 N. melanoleuca 0.225 mg/kg[12][16]
5 N. siamensis 0.25 mg/kg[12]
6 N. atra 0.28 mg/kg[12][17]
7 N. naja 0.29 mg/kg[12][17]
8 N. nivea 0.37 mg/kg[17]
9 N. kaouthia 0.47 mg/kg[12]
10 N. sumatrana 0.60 mg/kg[18]
All species in the genus Naja are capable of delivering a fatal bite to a human. Most species have strongly neurotoxic venom, which attacks the nervous system, causing paralysis, but many also have 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 spitting cobras, have a specialized venom delivery mechanism, in which their front fangs, instead of ejecting venom downward through an elongate 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[19] 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 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. The murine LD99-100 subcutaneous value according to Brown (1973) is 0.4 mg/kg,[14] while Ernst and Zug et al. list a LD50 range of 0.09 mg/kg - 0.21 mg/kg SC and 0.037 mg/kg IV.[16] Latifi (1984) listed a subcutaneous value of 0.2 mg/kg.[20] In another study, where venom was collected from a number of specimens in Iran, the Subcutis LD50 in lab mice was 0.078 mg/kg, the most potent Naja venom by this route of envenomation.[12] 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 poisoning by intracerebroventricular injection.[21] The Philippine cobra (N. philippinensis) has an average murine LD50 of 0.2 mg/kg subcutis.[14] The lowest value reported for the Philippine cobra is 0.14 mg/kg subcutis.[17][13][22] Other highly venomous species are the water cobras (Boulengerina clade) are also highly venomous. The murine intraperitoneal LD50 of Naja annulata and Naja christyi venoms were 0.143 mg/kg (range of 0.131 mg/kg to 0.156 mg/kg) and 0.120 mg/kg, respectively.[23] Christensen (1968) also listed an IV LD50 of 0.17 mg/kg for N. annulata.[24] The Chinese cobra (N. atra) is also highly venomous. Minton (1974) listed a value of LD50 0.28 mg/kg subcutis and 0.22 mg/kg intravenous (IV),[17] while Lee and Tseng list a value of 0.67 mg/kg SC.[25] Brown (1973) lists the LD99-100 at 0.2 mg/kg SC.[14] The forest cobra (N. melanoleuca) has a LD50 of 0.225 mg/kg subcutis,[15]
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 Africa (including some parts of the Sahara where Naja haje can be found), Southwest Asia, Central Asia, South Asia, East Asia, and Southeast Asia. Roughly 30% of bites by some cobra species are dry bites, thus do not cause envenomation (a dry bite is a bite by a venomous snake which does not inject venom).[26]
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.[14][27] to about 80% for N. oxiana.[28] Mortality rate for Naja atra is between 15 and 20%, 5–10% for N. nigricollis,[29] 50% for N. nivea,[14] 20–25% for N. naja,[30] In cases where victims of cobra bites are medically treated using normal treatment protocol for elapid type envenomation, differences in prognosis depend on the cobra species involved. The vast majority of envenomated patients treated make quick and complete recoveries, while other envenomated patients who receive similar treatment result in fatalities. The most important factors in the difference of mortality rates among victims envenomated by cobras is the severity of the bite and which cobra species caused the envenomation. The Caspian cobra (N. oxiana) and the Philippine cobra (N. philippinensis) are the two cobra species with the most toxic venom based on LD50 studies on mice. Both species cause prominent neurotoxicity and progression of life-threatening symptoms following envenomation. Death has been reported in as little as 30 minutes in cases of envenomation by both species. N. philippinensis purely neurotoxic venom causes prominent neurotoxicity with minimal local tissue damage and pain[31] and patients respond very well to antivenom therapy if treatment is administered rapidly after envenomation. Envenomation caused by N. oxiana is much more complicated. In addition to prominent neurotoxicity, very potent cytotoxic and cardiotoxic components are in this species' venom. Local effects are marked and manifest in all cases of envenomation - severe pain, severe swelling, bruising, blistering, and tissue necrosis. Renal damage and cardiotoxicity are also clinical manifestations of envenomation caused by N. oxiana, though they are rare and secondary.[32] The untreated mortality rate among those envenomed by N. oxiana approaches 80%, the highest among all species within the genus Naja.[28] Antivenom is not as effective for envenomation by this species as it is for other Asian cobras within the same region, like the Indian cobra (N. naja) and due to the dangerous toxicity of this species' venom, massive amounts of antivenom are often required for patients. As a result, a monovalent antivenom serum is being developed by the Razi Vaccine and Serum Research Institute in Iran. Response to treatment with antivenom is generally poor among patients, so mechanical ventilation and 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%).[20]
Taxonomy
| Naja |
|
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][33]
Naja anchietae (Bocage, 1879), Anchieta's cobra, is regarded as a subspecies of N. haje by Mertens (1937) and of N. annulifera by Broadley (1995). It is regarded as a full species by Broadley and Wüster (2004).[34][35]
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.[36]
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.[37][38]
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).[39][40]
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.[36]
Naja peroescobari Ceríaco et al. 2017, is a new species encompassing what was previously considered the São Tomé population of N. melanoleuca.[41]
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).[39][42] 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.[43] 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”.[44]
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.[45]
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 | Moroccan cobra | 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. 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 | Spanish cobra | 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 | 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 |
 |
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 | |
.jpg) |
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 | †European cobra | Miocene-aged strata of France, Germany, Austria, Romania, and Ukraine | |
 |
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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