Wikipedia

Naja

Naja is a genus of venomous elapid snakes commonly known as cobras (or "true cobras"). Various species occur throughout Africa, Southwest Asia, South Asia, and Southeast Asia. Several other elapid species are often called "cobras", such as the king cobra and the rinkhals, but they are not "true cobras", in that they do not belong to the genus Naja.[1][2][3]

Naja
Temporal range: Miocene–Holocene
Indian cobra (Naja naja), species typica of the genus
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Suborder: Serpentes
Family: Elapidae
Genus: Naja
Laurenti, 1768
Type species
Coluber naja
Linnaeus, 1758

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

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The origin of the generic name, Naja, is from the Sanskrit nāga (with a hard "g") meaning "snake". Some[who?] hold that the Sanskrit word is 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

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Naja species vary in length, and most are relatively slender-bodied snakes. Most species are capable of attaining a total length (tail included) 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 (Naja melanoleuca) arguably being the longest species.[10] All have a characteristic ability to raise the front quarter of the body off the ground and flatten the neck to appear larger to a potential predator. Fang structure is variable. All species except the Indian cobra (Naja naja), Egyptian cobra (Naja haje) and Caspian cobra (Naja oxiana) have some degree of adaptation to spitting.[11]

Venom

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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 that 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 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, and 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 pain-inducing 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 hypothesize 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, the murine LD50 via intravenous injection 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) 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). 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.[24] Christensen (1968) also listed an IV LD50 of 0.17 mg/kg for N. annulata.[17] The Chinese cobra (N. atra) is also highly venomous. Minton (1974) listed a value of LD50 0.3 mg/kg intravenous (IV),[19] while Lee and Tseng list a value of 0.67 mg/kg 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] The Senegalese cobra (N. senegalensis) has a murine LD50 of 0.39 mg/kg (Tan et al, 2021) via IV.[26] The Egyptian cobra (N. haje) of Ugandan locality had an IV LD50 of 0.43 mg/kg (0.35–0.52 mg/kg).[27]

Medical importance

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The Naja species is a medically important group of snakes, due to the number of bites and fatalities the species causes across its geographical 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 (or venomless) bites, thus do not cause envenomation.[28] Brown (1973) noted that cobras with a higher rates of 'sham strikes' tend to be more venomous, while those with a less toxic venom tend to envenomate more frequently when attempting to bite. This can vary even between specimens of the same species. This is unlike related elapids, such as those species belonging to Dendroaspis (mambas) and Bungarus (kraits), with mambas tending to almost always envenomate and kraits tending to envenomate more often than they attempt 'sham strikes'.[29]

Many factors influence the differences in cases of fatality between 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.[29][30] to about 80% for N. oxiana.[31] Mortality rate for N. atra is between 15 and 20%, 5–10% for N. nigricollis,[32] 50% for N. nivea,[29] 20–25% for N. naja,[33] 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 are 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[34] 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.[35] The untreated mortality rate among those envenomed by N. oxiana approaches 80%, the highest among all species within the genus Naja.[31] 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 are 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 (under 1%).[15]

Taxonomy

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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][36]

  • 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).[37][38]
  • 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.[39]
  • 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.[40][41]
  • 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).[42][43]
  • 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.[39]
  • Naja peroescobari Ceríaco et al. 2017, is a new species encompassing what was previously considered the São Tomé population of N. melanoleuca.[44]
  • 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 molecular phylogenetic studies in the early 2000s 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).[42][45] 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.[46] 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".[47]

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 falling as the sister lineage to all other species in the subgenus.[36]

Species

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Genus Naja – Laurenti, 1768 – 35 extant species
Common name Scientific name and subspecies Range Size and ecology IUCN status and estimated population
Anchieta's cobra

{{{image-alt}}}

 N. anchietae
Bocage, 1879		 Southern Africa
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[48]

Banded water cobra

{{{image-alt}}}

 N. annulata
Buchholz & Peters, 1876

Two subspecies
  • N. a. annulata Buchholz & Peters, 1876
  • N. a. stormsi (Dollo, 1886)
 West and Central Africa
Map of range		 Size: {{{size}}}

Habitat: Wetlands in forests and dense savannas[49]

Diet: Fish, possibly amphibians[49]		  LC 


Population steady[49]

Snouted cobra

{{{image-alt}}}

 N. annulifera
W. Peters, 1854		 Southern Africa
Map of range		 Size: {{{size}}}

Habitat: Savannas, bushveld, lowveld[50]

Diet: {{{hunting}}}		  LC 


Population steady[50]

Arabian cobra N. arabica
Scortecci, 1932		 Oman, Saudi Arabia, Yemen
Map of range		 Size: {{{size}}}

Habitat: Mesic habitats[51]

Diet: {{{hunting}}}		  LC 


Unknown[51]

Ashe's spitting cobra

{{{image-alt}}}

 N. ashei
Wüster & Broadley, 2007		 Southern Ethiopia, Kenya, Somalia, eastern Uganda Size: {{{size}}}

Habitat: Semi-deserts, savannas, grasslands, and woodlands[51]

Diet: {{{hunting}}}		  LC 


Population steady[52]

Chinese cobra

{{{image-alt}}}

 N. atra
Cantor, 1842		 Southern China, northern Laos, Taiwan, northern Vietnam
Map of range		 Size: {{{size}}}

Habitat: Shrublands, agricultural land, occasionally urban areas[53]

Diet: Rats, frogs, reptiles, birds, fish[53]		  VU 


Population declining[53]

Congo water cobra N. christyi
(Boulenger, 1904)		 Democratic Republic of the Congo, Republic of Congo, Angola (Cabinda)
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population declining[54]

Brown banded cobra

{{{image-alt}}}

 N. fuxi
Shi, G. Vogel, Chen & Ding, 2022		 China, Myanmar, Laos, Thailand, Vietnam Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}
Black forest cobra

{{{image-alt}}}

 N. guineensis
Broadley, Trape, Chirio, Ineich & Wüster, 2018		 Ghana, Guinea, Guinea-Bissau, the Ivory Coast, Liberia, Sierra Leone, Togo Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}
Egyptian cobra

{{{image-alt}}}

 N. haje
(Linnaeus, 1758)		 Scattered areas across Africa
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population declining[55]

Monocled cobra

{{{image-alt}}}

 N. kaouthia
Lesson, 1831		 South and Southeast Asia
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population declining[56]

Mali cobra

{{{image-alt}}}

 N. katiensis
Angel, 1922		 Benin, Burkina Faso, Cameroon, Ghana, Guinea, the Ivory Coast, Mali, Gambia, Mauritania, Niger, Nigeria, Senegal, Togo Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[57]

Mandalay spitting cobra

{{{image-alt}}}

 N. mandalayensis
Slowinski & Wüster, 2000		 Myanmar
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  VU 


Unknown Population declining[58]

Central African forest cobra

{{{image-alt}}}

 N. melanoleuca
Hallowell, 1857		 Sub-Saharan Africa
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population declining[59]

Mozambique spitting cobra

{{{image-alt}}}

 N. mossambica
W. Peters, 1854		 Southern Africa
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Unknown[60]

Many-banded cobra N. multifasciata
(F. Werner, 1902)		 Central Africa
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[61]

Indian cobra

{{{image-alt}}}

 N. naja
(Linnaeus, 1758)		 South Asia
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[62]

Dwarf water cobra

{{{image-alt}}}

 N. nana
Collet & Trape, 2020		 Democratic Republic of Congo Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}
Zebra spitting cobra

{{{image-alt}}}

 N. nigricincta
Bogert, 1940

Two subspecies
  • N. n. nigricincta Bogert, 1940
  • N. n. woodi Pringle, 1955
 Angola, Namibia, South Africa
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[63]

Black-necked spitting cobra

{{{image-alt}}}

 N. nigricollis
Reinhardt, 1843		 Sub-Saharan Africa
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Unknown[64]

Cape cobra

{{{image-alt}}}

 N. nivea
(Linnaeus, 1758)		 Southern Africa
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[65]

Nubian spitting cobra

{{{image-alt}}}

 N. nubiae
Wüster & Broadley, 2003		 Chad, Egypt, Eritrea, Niger, Sudan
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[66]

Obscure cobra N. obscura
Saleh & Trape, 2023		 Egypt Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}
Caspian cobra

{{{image-alt}}}

 N. oxiana
(Eichwald, 1831)		 Afghanistan, northwestern India, Iran, Kyrgyzstan, Pakistan, Tajikistan, Turkmenistan, Uzbekistan Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  NT 


Population declining[67]

Red spitting cobra

{{{image-alt}}}

 N. pallida
Boulenger, 1896		 Djibouti, Ethiopia, Kenya, Somalia, Tanzania Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[68]

São Tomé forest cobra

{{{image-alt}}}

 N. peroescobari
Ceríaco, Marques, Schmitz & Bauer, 2017		 São Tomé and Príncipe (São Tomé) Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  EN 


Population declining[69]

Philippine cobra

{{{image-alt}}}

 N. philippinensis
Taylor, 1922		 Philippines (Luzon, Mindoro)
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  NT 


Population declining[70]

Andaman cobra

{{{image-alt}}}

 N. sagittifera
Wall, 1913		 India (Andaman Islands) Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  EN 


Population declining[71]

Samar cobra

{{{image-alt}}}

 N. samarensis
W. Peters, 1861		 Philippines (Mindanao, Bohol, Leyte, Samar, Camiguin)
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population declining[72]

West African banded cobra

{{{image-alt}}}

 N. savannula
Broadley, Trape, Chirio & Wüster, 2018		 Benin, Burkina Faso, Cameroon, Chad, Gambia, Ghana, Guinea, the Ivory Coast, Mali, Niger, Nigeria, Senegal, Togo Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[73]

Senegalese cobra N. senegalensis
Trape, Chirio & Wüster, 2009		 Benin, Burkina Faso, Ghana, Guinea, Guinea-Bissau, the Ivory Coast, Mali, Niger, Nigeria, Senegal Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[74]

Indochinese spitting cobra

{{{image-alt}}}

 N. siamensis
Laurenti, 1768		 Cambodia, Laos, Thailand, Vietnam
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  VU 


Population declining[75]

Javan spitting cobra

{{{image-alt}}}

 N. sputatrix
F. Boie, 1827		 Indonesia (Java, Lesser Sunda Islands)
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Unknown[76]

Brown forest cobra

{{{image-alt}}}

 N. subfulva
Laurent, 1955		 Angola, Burundi, Cameroon, the Central African Republic, Chad, the Republic of Congo, the Democratic Republic of the Congo, Ethiopia, Kenya, Malawi, Mozambique, Nigeria, Rwanda, Somalia, South Africa, South Sudan, Tanzania, Uganda, Zambia, Zimbabwe Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population steady[77]

Equatorial spitting cobra

{{{image-alt}}}

 N. sumatrana
J. Müller, 1887		 Southeast Asia
Map of range		 Size: {{{size}}}

Habitat: {{{habitat}}}

Diet: {{{hunting}}}		  LC 


Population increasing[78]

Fossil species

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Genus Naja – Laurenti, 1768 – 3 extinct species
Species name Image Range Size and ecology
N. antiqua

Rage, 1976

 Miocene

Morocco

 Size:

Habitat:
N. iberica

Szyndlar, 1985

 Miocene

Spain

 Size:

Habitat:
N. romani

(Hofstetter, 1939)

 Miocene[79]

France, Germany, Austria, Russia, Hungary, Greece and Ukraine[79]

 Size:

Habitat:

References

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  1. ↑ Spawls, S.; Branch, B. (1995). The Dangerous Snakes of Africa (1st. ed.). Sanibel Island, Florida: Ralph Curtis Books. 192 pp. ISBN 9780883590294. Retrieved 9 April 2020.
  2. ↑ Zhao, E.; Adler, K. (1993). Herpetology of China (1st ed.). Oxford, Ohio: Society for the Study of Amphibians & Reptiles. 522 pp. ISBN 9780916984281. OCLC 716490697.
  3. ↑ Vogel, G. (31 March 2006). Terralog: Venomous Snakes of Asia, Vol. 14 (1 ed.). Frankfurt Am Main: Hollywood Import & Export. p. 148. ISBN 978-3936027938.
  4. 1 2 von Plettenberg Laing, Anthony (2018). "A multilocus phylogeny of the cobra clade elapids". MSC Thesis.
  5. 1 2 "Naja". Integrated Taxonomic Information System. Retrieved 13 April 2008.
  6. 1 2 Wallach, Van; Wüster, W; Broadley, Donald G. (2009). "In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae)" (PDF). Zootaxa. 2236 (1): 26–36. doi:10.11646/zootaxa.2236.1.2. S2CID 14702999.
  7. 1 2 3 4 Wüster, W.; et al. (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. PMID 30314221.
  8. ↑ "Proto-IE: *(s)nēg-o-, Meaning: snake, Old Indian: nāgá- m. 'snake', Germanic: *snēk-a- m., *snak-an- m., *snak-ō f.; *snak-a- vb". Starling.rinet.ru.
  9. ↑ Mayrhofer, Manfred (1996). Etymologisches Wörterbuch des Altindoarischen. Heidelberg: Universitätsverlag C. Winter. p. II.33. ISBN 978-3-8253-4550-1. (in German).
  10. ↑ "Naja melanoleuca – General Details, Taxonomy and Biology, Venom, Clinical Effects, Treatment, First Aid, Antivenoms". WCH Clinical Toxinology Resource. University of Queensland. Retrieved 17 December 2011.
  11. ↑ Wüster, W.; Thorpe, R.S. (1992b). "Dentitional phenomena in cobras revisited: fang structure and spitting in the Asiatic species of Naja (Serpentes: Elapidae)". Herpetologica. 48: 424–434. Retrieved 31 October 2021.
  12. ↑ Kazandjian, T.D.; et al. (January 2021). "Convergent evolution of pain-inducing defensive venom components in spitting cobras" (PDF). Science. 371 (6527): 386–390. Bibcode:2021Sci...371..386K. doi:10.1126/science.abb9303. PMC 7610493. PMID 1294479. S2CID 231666401.
  13. ↑ Kazemi-Lomedasht, Fatemeh; Yamabhai, Montarop; Sabatier, Jean-Marc; Behdani, Mahdi; Zareinejad, Mohammad Reza; Shahbazzadeh, Delavar (5 December 2019). "Development of a human scFv antibody targeting the lethal Iranian cobra (Naja oxiana) snake venom". Toxicon. 171: 78–85. Bibcode:2019Txcn..171...78K. doi:10.1016/j.toxicon.2019.10.006. PMID 31622638. S2CID 204772656.
  14. 1 2 3 Wong, Kin Ying; Tan, Choo Hock; Tan, Nget Hong (3 January 2019). "Venom and Purified Toxins of the Spectacled Cobra (Naja naja) from Pakistan: Insights into Toxicity and Antivenom Neutralization". The American Journal of Tropical Medicine and Hygiene. 94 (6): 1392–1399. doi:10.4269/ajtmh.15-0871. PMC 4889763. PMID 27022154.
  15. 1 2 Latifi, Mahmoud (1984). Snakes of Iran. Oxford, Ohio: Society for the Study of Amphibians & Reptiles. ISBN 978-0-916984-22-9. 156 pp.
  16. ↑ Khare, A.D.; Khole, V.; Gade, P.R. (December 1992). "Toxicities, LD50 prediction and in vivo neutralisation of some elapid and viperid venoms". Indian Journal of Experimental Biology. 30 (12): 1158–62. PMID 1294479.
  17. 1 2 Christensen, Poul Agerholm (1968). "Chapter 16 – The Venoms of Central and South African Snakes". Venomous Animals and their Venoms, Volume I, Venomous Vertebrates. Cambridge, Massachusetts: Academic Press. pp. 437–461. ISBN 978-1-4832-2949-2. Retrieved 24 May 2021.
  18. ↑ "Boulengerina annulata – General Details, Taxonomy and Biology, Venom, Clinical Effects, Treatment, First Aid, Antivenoms". Clinical Toxinology Resource. University of Adelaide. Retrieved 7 September 2022.
  19. 1 2 3 Minton, S.A., Jr. (1974). Venom Diseases. Springfield, Illinois: C.C. Thomas. ISBN 978-0-398-03051-3.{{cite book}}: CS1 maint: multiple names: authors list (link) 256 pp.
  20. ↑ Watt, G.; Theakston, R.D.; Hayes, C.G.; Yambao, M.L.; Sangalang, R.; Ranoa, C.P.; Alquizalas, E.; Warrell, D.A. (4 December 1986). "Positive response to edrophonium in patients with neurotoxic envenoming by cobras (Naja naja philippinensis). A placebo-controlled study". New England Journal of Medicine. 315 (23): 1444–8. doi:10.1056/NEJM198612043152303. PMID 3537783.
  21. ↑ Hauert, Jacques; Maire, Michel; Sussmann, Alexandre; Bargetzi, J. Pierre (July 1974). "The major lethal neurotoxin of the venom of Naja naja phillippinensis: Purification, physical and chemical properties, partial amino acid sequence". International Journal of Peptide and Protein Research. 6 (4): 201–222. doi:10.1111/j.1399-3011.1974.tb02380.x. PMID 4426734.
  22. ↑ Palasuberniam, Praneetha; Chan, Yi Wei; Tan, Kae Yi; Tan, Choo Hock (2021). "Snake Venom Proteomics of Samar Cobra (Naja samarensis) from the Southern Philippines: Short Alpha-Neurotoxins as the Dominant Lethal Component Weakly Cross-Neutralized by the Philippine Cobra Antivenom". Frontiers in Pharmacology. 12: 3641. doi:10.3389/fphar.2021.727756. ISSN 1663-9812. PMC 8740184. PMID 35002690.
  23. ↑ Lysz, Thomas W.; Rosenberg, Philip (May 1974). "Convulsant activity of Naja naja oxiana venom and its phospholipase A component". Toxicon. 12 (3): 253–265. doi:10.1016/0041-0101(74)90067-1. PMID 4458108.
  24. ↑ Weinstein, S.A.; Schmidt, J.J.; Smith, L.A. (1991). "Lethal toxins and cross neutralization of venoms from the African Water Cobras, Boulengerina annulata annulata and Boulengerina christyi ". Toxicon. 29 (11): 1315–27. Bibcode:1991Txcn...29.1315W. doi:10.1016/0041-0101(91)90118-b. PMID 1814007.
  25. ↑ Lee, C.Y.; Tseng, L.F. (September 1969). "Species differences in susceptibility to elapid venoms". Toxicon. 7 (2): 89–93. Bibcode:1969Txcn....7...89L. doi:10.1016/0041-0101(69)90069-5. ISSN 0041-0101. PMID 5823351. Retrieved 24 May 2021.
  26. ↑ Wong, K.Y.; Tan, K.Y.; Tan, N.H.; Tan, C.H. (2021). "A Neurotoxic Snake Venom without A2: Proteomics and Cross-Neutralization of the Venom from Senegalese Cobra, Naja senegalensis (Subgenus: Uraeus)". Toxins. 13 (1): 60. doi:10.3390/toxins13010060. PMC 7828783. PMID 33466660.
  27. ↑ Harrison, G.O.; Oluoch, R.A.; Ainsworth, S.; Alsolaiss, J.; Bolton, F.; Arias, A.S.; et al. (2017). "The murine venom lethal dose (LD50) of the East African snakes". PLOS Neglected Tropical Diseases. doi:10.1371/journal.pntd.0005969.t003. Retrieved 20 September 2022.
  28. ↑ Naik, B.S. (2017). ""Dry bite" in venomous snakes: A review" (PDF). Toxicon. 133: 63–67. Bibcode:2017Txcn..133...63N. doi:10.1016/j.toxicon.2017.04.015. PMID 28456535. S2CID 36838996. Retrieved 24 May 2021.
  29. 1 2 3 Brown, John Haynes (1973). Toxicology and Pharmacology of Venoms from Poisonous Snakes. Springfield, Illinois: Charles C. Thomas. p. 81. ISBN 978-0-398-02808-4. 184 pp.
  30. ↑ Warrell, D.A. (22 August 1995). "Clinical Toxicology of Snakebite in Asia". Handbook of: Clinical Toxicology of Animal Venoms and Poisons (1st ed.). CRC Press. pp. 493–594. doi:10.1201/9780203719442-27. ISBN 9780203719442. Retrieved 24 May 2021.
  31. 1 2 Gopalkrishnakone, P.; Chou, L.M. (1990). Snakes of Medical Importance (Asia-Pacific Region). Singapore: National University of Singapore. ISBN 978-9971-62-217-6.[page needed]
  32. ↑ Warrell, David A. "Snake bite" (PDF). Seminar. Lancet 2010 (volume 375, issue 1). Archived from the original (PDF) on 29 October 2013. Retrieved 20 December 2011.
  33. ↑ Norris, Robert L.; Minton, Sherman A. (10 September 2013). "Cobra Envenomation". Medscape. United States. Retrieved 8 December 2013.
  34. ↑ Watt, G.; Padre L; Tuazon L; Theakston RD; Laughlin L. (September 1988). "Bites by the Philippine cobra (Naja naja philippinensis): prominent neurotoxicity with minimal local signs". The American Journal of Tropical Medicine and Hygiene. 39 (3): 306–11. doi:10.4269/ajtmh.1988.39.306. PMID 3177741.
  35. ↑ "Naja oxiana". Clinical Toxinology Resource. University of Adelaide. Retrieved 8 December 2013.
  36. 1 2 Kazemi, Elmira; Nazarizadeh, Masoud; Fatemizadeh, Faezeh; Khani, Ali; Kaboli, Mohammad (2021). "The phylogeny, phylogeography, and diversification history of the westernmost Asian cobra (Serpentes: Elapidae: Naja oxiana) in the Trans-Caspian region". Ecology and Evolution. 11 (5): 2024–2039. Bibcode:2021EcoEv..11.2024K. doi:10.1002/ece3.7144. ISSN 2045-7758. PMC 7920780. PMID 33717439.
  37. ↑ Broadley, D.G.; Wüster, W. (2004). "A review of the southern African 'non-spitting' cobras (Serpentes: Elapidae: Naja)". African Journal of Herpetology. 53 (2): 101–122. Bibcode:2004AfJH...53..101B. doi:10.1080/21564574.2004.9635504. S2CID 84853318.
  38. ↑ Naja anchietae at the Reptarium.cz Reptile Database. Accessed 13 April 2007.
  39. 1 2 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. doi:10.11646/zootaxa.2236.1.1.
  40. ↑ Wüster, W.; Broadley, D.G. (2007). "Get an eyeful of this: a new species of giant spitting cobra from eastern and north-eastern Africa (Squamata: Serpentes: Elapidae: Naja)". Zootaxa. 1532: 51–68. doi:10.11646/zootaxa.1532.1.4.
  41. ↑ Naja ashei at the Reptarium.cz Reptile Database. Accessed 13 April 2007.
  42. 1 2 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. Bibcode:2007MolPE..45..437W. doi:10.1016/j.ympev.2007.07.021. PMID 17870616.
  43. ↑ Naja nigricincta at the Reptarium.cz Reptile Database. Accessed 29 December 2008.
  44. ↑ Ceríaco, L; et al. (2017). "The "Cobra-preta" of São Tomé Island, Gulf of Guinea, is a new species of Naja Laurenti, 1768 (Squamata: Elapidae)". Zootaxa. 4324 (1): 121–141. doi:10.11646/zootaxa.4324.1.7.
  45. ↑ Nagy, Z.T.; Vidal, N.; Vences, M.; Branch, W.R.; Pauwels, O.S.G.; Wink, M.; Joger, U. (2005). "Molecular systematics of African Colubroidea (Squamata: Serpentes)". pp. 221–228. In: Huber, B.A.; Sinclair, B.J.; Lampe, K.-H. (editors) (2005). African Biodiversity: Molecules, Organisms, Ecosystems. Proceedings of the 5th International Symposium on Tropical Biology. Bonn: Museum Koenig.
  46. ↑ Hoser, R. (2009). "Naja, Boulengerina and Paranaja". Australasian Journal of Herpetology 7: 1–15.
  47. ↑ "Opinion 2468 (Case 3601) – Spracklandus Hoser, 2009 (Reptilia, Serpentes, Elapidae) and Australasian Journal of Herpetology issues 1–24: confirmation of availability declined; Appendix A (Code of Ethics): not adopted as a formal criterion for ruling on Cases". The Bulletin of Zoological Nomenclature. 78 (1): 42–45. 2021. doi:10.21805/bzn.v78.a012. ISSN 0007-5167. S2CID 233448875.
  48. ↑ Alexander, G.J.; Tolley, K.A. (2021). "Naja anchietae". IUCN Red List of Threatened Species. 2021 e.T110168507A110168532. doi:10.2305/IUCN.UK.2021-2.RLTS.T110168507A110168532.en. Retrieved 24 April 2026.
  49. 1 2 3 Wagner, P.; Branch, W.R.; Safari, I.; Chenga, J.; Gonwouo, N.L.; Kusamba, C.; Zassi-Boulou, A.-G. (2021). "Naja annulata". IUCN Red List of Threatened Species. 2021 e.T44929978A44929995. doi:10.2305/IUCN.UK.2021-1.RLTS.T44929978A44929995.en. Retrieved 24 April 2026.
  50. 1 2 Alexander, G.J.; Tolley, K.A. (2021). "Naja annulifera". IUCN Red List of Threatened Species. 2021 e.T110168579A120635887. doi:10.2305/IUCN.UK.2021-2.RLTS.T110168579A120635887.en. Retrieved 24 April 2026.
  51. 1 2 3 Els, J.; Al Johany, A.M.H. (2012). "Naja arabica". IUCN Red List of Threatened Species. 2012 e.T199850A2613995. doi:10.2305/IUCN.UK.2012.RLTS.T199850A2613995.en. Retrieved 27 April 2026.
  52. ↑ Spawls, S.; Malonza, P.; Beraduccii, J. (2021). "Naja ashei". IUCN Red List of Threatened Species. 2021 e.T20878760A20878765. doi:10.2305/IUCN.UK.2021-1.RLTS.T20878760A20878765.en. Retrieved 27 April 2026.
  53. 1 2 3 Wogan, G.; Stuart, B. (2012). "Naja mandalayensis". IUCN Red List of Threatened Species. 2012 e.T192227A2058297. doi:10.2305/IUCN.UK.2012-1.RLTS.T192227A2058297.en. Retrieved 24 April 2026.
  54. ↑ Kusamba, C. (2021). "Naja christyi". IUCN Red List of Threatened Species. 2021 e.T110168644A110168680. doi:10.2305/IUCN.UK.2021-1.RLTS.T110168644A110168680.en. Retrieved 27 April 2026.
  55. ↑ Wilms, T.; Wagner, P.; Spawls, S.; Beraduccii, J.; Malonza, P.K. (2021). "Naja haje". IUCN Red List of Threatened Species. 2021 e.T184071A1748178. doi:10.2305/IUCN.UK.2021-2.RLTS.T184071A1748178.en. Retrieved 24 April 2026.
  56. ↑ Stuart, B.; Wogan, G. (2012). "Naja kaouthia". IUCN Red List of Threatened Species. 2012 e.T177487A1488122. doi:10.2305/IUCN.UK.2012-1.RLTS.T177487A1488122.en. Retrieved 27 April 2026.
  57. ↑ Luiselli, L.; Chirio, L.; Wagner, P.; Wilms, T.; Chippaux, J.-P. (2021). "Naja katiensis". IUCN Red List of Threatened Species. 2021 e.T13265887A126471645. doi:10.2305/IUCN.UK.2021-2.RLTS.T13265887A126471645.en. Retrieved 27 April 2026.
  58. ↑ Wogan, G.; Stuart, B. (2012). "Naja mandalayensis". IUCN Red List of Threatened Species. 2012 e.T192227A2058297. doi:10.2305/IUCN.UK.2012-1.RLTS.T192227A2058297.en. Retrieved 24 April 2026.
  59. ↑ Jallow, M.; Penner, J.; Rödel, M.-O.; Luiselli, L.; Chippaux, J.-P.; Gonwouo, N.L.; Kusamba, C.; Zassi-Boulou, A.-G. (2021). "Naja melanoleuca". IUCN Red List of Threatened Species. 2021 e.T13265900A13265907. doi:10.2305/IUCN.UK.2021-1.RLTS.T13265900A13265907.en. Retrieved 27 April 2026.
  60. ↑ Verburgt, L.; Pietersen, D.; Farooq, H.; Chapeta, Y.; Wagner, P.; Safari, I.; Chenga, J. (2022) [amended version of 2020 assessment]. "Naja mossambica". IUCN Red List of Threatened Species. 2022 e.T20878771A219349342. doi:10.2305/IUCN.UK.2022-2.RLTS.T20878771A219349342.en. Retrieved 27 April 2026.
  61. ↑ Kusamba, C.; Chirio, L.; Gonwouo, N.L.; Zassi-Boulou, A.-G. (2021). "Naja multifasciata". IUCN Red List of Threatened Species. 2021 e.T110168688A110168696. doi:10.2305/IUCN.UK.2021-1.RLTS.T110168688A110168696.en. Retrieved 27 April 2026.
  62. ↑ de Silva, A.; Ukuwela, K.; Shankar, G.; Srinivasulu, B.; Das, A.; Vyas, R.; Sawant, N.S.; Kulkarni, N.U.; Deepak, V.; Thakur, S.; Mohapatra, P.; Srinivasulu, C.; Achyuthan, N.S. (2021). "Naja naja". IUCN Red List of Threatened Species. 2021 e.T62241A3110222. doi:10.2305/IUCN.UK.2021-3.RLTS.T62241A3110222.en. Retrieved 27 April 2026.
  63. ↑ Alexander, G.J.; Tolley, K.A. (2021). "Naja nigricincta". IUCN Red List of Threatened Species. 2021 e.T110168705A139747789. doi:10.2305/IUCN.UK.2021-2.RLTS.T110168705A139747789.en. Retrieved 27 April 2026.
  64. ↑ Jallow, M.; Luiselli, L.; Penner, J.; Rödel, M.-O.; Segniagbeto, G.; Niagate, B.; Spawls, S.; Howell, K.; Msuya, C.A.; Ngalason, W. (2021). "Naja nigricollis". IUCN Red List of Threatened Species. 2021 e.T13265913A13265918. doi:10.2305/IUCN.UK.2021-2.RLTS.T13265913A13265918.en. Retrieved 27 April 2026.
  65. ↑ Alexander, G.J.; Maritz, B. (2021). "Naja nivea". IUCN Red List of Threatened Species. 2021 e.T110168750A139749003. doi:10.2305/IUCN.UK.2021-2.RLTS.T110168750A139749003.en. Retrieved 27 April 2026.
  66. ↑ Luiselli, L.; Chirio, L.; Wagner, P.; Baha El Din, S. (2021). "Naja nubiae". IUCN Red List of Threatened Species. 2021 e.T184072A1748449. doi:10.2305/IUCN.UK.2021-1.RLTS.T184072A1748449.en. Retrieved 27 April 2026.
  67. ↑ Ananjeva, N.B.; Orlov, N.L.; Nilson, G.; Papenfuss, T.; Borkin, L.; Milto, K.; Golynsky, E.; Rustamov, A, Nuridjanov, D.; Munkhbayar, K.; Murthy, B.H.C.; Mohapatra, P. (2021). "Naja oxiana". IUCN Red List of Threatened Species. 2021 e.T164642A1063259. doi:10.2305/IUCN.UK.2021-3.RLTS.T164642A1063259.en. Retrieved 27 April 2026.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  68. ↑ Beraduccii, J.; Spawls, S.; Malonza, P. (2021). "Naja pallida". IUCN Red List of Threatened Species. 2021 e.T20878782A20878789. doi:10.2305/IUCN.UK.2021-1.RLTS.T20878782A20878789.en. Retrieved 27 April 2026.
  69. ↑ Ceríaco, L.M.P. (2021). "Naja peroescobari". IUCN Red List of Threatened Species. 2021 e.T133837440A133837443. doi:10.2305/IUCN.UK.2021-2.RLTS.T133837440A133837443.en. Retrieved 27 April 2026.
  70. ↑ dela Cruz, C.; Lagat, R.; Salinas, R.; Pitogo, K.; Santos, G.; Sarmiento, M.; Sy, E. (2022). "Naja philippinensis". IUCN Red List of Threatened Species. 2022 e.T169768A180845902. Retrieved 27 April 2026.
  71. ↑ Mohapatra, P.; Achyuthan, N.S. (2021). "Naja sagittifera". IUCN Red List of Threatened Species. 2021 e.T202848A2757313. doi:10.2305/IUCN.UK.2021-3.RLTS.T202848A2757313.en. Retrieved 27 April 2026.
  72. ↑ Maglangit, E.P.; Santos, G.; Sy, E. (2022). "Naja samarensis". IUCN Red List of Threatened Species. 2022 e.T169763A180846510. Retrieved 27 April 2026.
  73. ↑ Chirio, L.; Gonwouo, N.L. (2021). "Naja savannula". IUCN Red List of Threatened Species. 2021 e.T133836145A133836160. doi:10.2305/IUCN.UK.2021-1.RLTS.T133836145A133836160.en. Retrieved 27 April 2026.
  74. ↑ Penner, J.; Wilms, T.; Wagner, P.; Niagate, B. (2013). "Naja senegalensis". IUCN Red List of Threatened Species. 2013 e.T13265924A13265927. doi:10.2305/IUCN.UK.2013-1.RLTS.T13265924A13265927.en. Retrieved 27 April 2026.
  75. ↑ Stuart, B.; Thy, N.; Chan-Ard, T.; Nguyen, T.Q.; Bain, R. (2012). "Naja siamensis". IUCN Red List of Threatened Species. 2012 e.T177488A1488437. doi:10.2305/IUCN.UK.2012-1.RLTS.T177488A1488437.en. Retrieved 27 April 2026.
  76. ↑ Iskandar, D.; Auliya, M.; Inger, R.F.; Lilley, R. (2012). "Naja sputatrix". IUCN Red List of Threatened Species. 2012 e.T192197A2054180. doi:10.2305/IUCN.UK.2012-1.RLTS.T192197A2054180.en. Retrieved 27 April 2026.
  77. ↑ Spawls, S.; Malonza, P.K.; Msuya, C.A.; Zassi-Boulou, A.-G.; Chippaux, J.-P.; Kusamba, C.; Gonwouo, N.L.; Chirio, L. (2021). "Naja subfulva". IUCN Red List of Threatened Species. 2021 e.T133837181A133837214. doi:10.2305/IUCN.UK.2021-2.RLTS.T133837181A133837214.en. Retrieved 27 April 2026.
  78. ↑ Grismer, L.; Chan-Ard, T.; Diesmos, A.C.; Sy, E. (2012). "Naja sumatrana". IUCN Red List of Threatened Species. 2012 e.T184073A1748598. doi:10.2305/IUCN.UK.2012-1.RLTS.T184073A1748598.en. Retrieved 27 April 2026.
  79. 1 2 Syromyatnikova, E.; Tesakov, A.; Titov, V. (2021). "Naja romani (Hoffstetter, 1939)(Serpentes: Elapidae) from the late Miocene of the Northern Caucasus: the last East European large cobra". Geodiversitas. 43 (19): 683–689. doi:10.5252/geodiversitas2021v43a19. S2CID 238231298.
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