A review on diversity, bio-geography, and updated checklist of Cobras (Naja, Laurenti, 1768) (Squamata: Serpentes: Elapidae): Present and prehistoric
DOI:
https://doi.org/10.5281/zenodo.17839995Keywords:
Africa, Asia, Biogeography, Diversity, Naja, ToxicityAbstract
The Naja Laurenti, 1768, is a genus of venomous snakes widely considered as "true" cobras and belongs to the family Elapidae. The living species of this genus are mainly found in Africa and Asia. This review article provides an updated checklist of 35 recognized living species of the genus Naja, including 22 (63%) African species, 13 (37%) Asian species, and four currently recognized extinct cobra species from Miocene and Pliocene strata discovered in Africa and Europe. The Naja genus is divided into four subgenera: 1. The typical subgenus Naja is restricted to Asian species, 2. The subgenus Uraeus, Wagler, 1830, is revived for a group of non-spitting cobras found in the savannas and open lands of Africa and the Arabian Peninsula, 3. Boulengerina, Dollo, 1886, is used for non-spitting African cobra species, including forest, aquatic, and semi-fossorial forms. The 4th subgenus, Afronaja, is designated for all African spitting cobras. This review also summarizes the diversity, distribution, conservation status, and toxicity levels of some Naja species. The searches were limited to peer-reviewed journals written in English, excluding books and other non-English sources.
References
Ali, W., Javid, A., Hussain, A., & Bukhari, S. M. (2018). “Diversity and habitat preferences of amphibians and reptiles in Pakistan”: A review. Journal of Asia-Pacific Biodiversity. 11(2), 173–187. https://doi.org/10.1016/j.japb.2018.01.009.
Alberdi, M.T., Morales, J., Moyà, S., & Sanchiz, B. (1981). Macrovertebrados (Reptilia & Mammalia) del yacimiento finimioceno de Librilla (Murcia). Estudios Geológicos, 37, 307–312.
Alférez Delgado, F., & Brea López, P. (1981). Estudio preliminar de los restos de peces, anfibios y reptiles delyacimiento mioceno de Córcoles (Guadalajara). Boletín de la Real Sociedad Española de Historia Natural, 79, 5–20.
Angel, F. (1922) Sur une collection de reptiles et de batraciens, recueillis au Soudan français par la mission du Dr. Millet
Bachmayer, F., & Szyndlar, Z. (1985). Ophidians (Reptilia: Serpentes) from the Kohfidisch fissures of Burgenland, Austria. Annalen des Naturhistorischen Museums in Wien, 87A, 79–100.
Bachmayer, F., & Szyndlar, Z. (1987). A second contribution to the ophidian fauna (Reptilia: Serpentes) of Kohfidisch, Austria. Annalen des Naturhistorischen Museums in Wien, 88A, 25–39.
Bailon, S. (1989). Les amphibiens et les reptiles du Pliocène supérieur de Balaruc II (Hérault, France). Palaeovertebrata, 19, 7–28.
Bailon, S. (1991). Amphibiens et reptiles du Pliocène et du Quaternaire de France et d'Espagne: Mise en place et évolution des faunes (Doctoral dissertation). Université Paris VII.
Bailon, S., Hossini, S., & Rage, J.-C. (2017). Les amphibiens et lépidosauriens du Cénozoïque du Maroc. In S. Zouhri (Ed.), Paléontologie des vertébrés du Maroc: État des connaissances (Mémoires de la Société Géologique de France, 180, pp. 453–484).
Bernor, R. L., Kordos, L., & Rook, L. (2004). Recent advances on multidisciplinary research at Rudabánya, Late Miocene (MN9), Hungary: A compendium. Palaeontographica Italica, 89, 3–36.
Biton, R., & Bailon, S. (2022). An African adder (Bitis arietans complex) at Qafzeh Cave, Israel, during the early Late Pleistocene (MIS 5). Journal of Quaternary Science, 37, 639–650.
Biton, R., & Bailon, S. (2023). Corrigendum: Correction to “An African adder (Bitis arietans complex) at Qafzeh Cave, Israel, during the early Late Pleistocene (MIS 5).” Journal of Quaternary Science 2023, 38, 608–611.
Blain, H.-A., Agustí, J., López-García, J.M., Haddoumi, H., Aouraghe, H., El Hammouti, K., Pérez-González, A., Chacón, M.G., & Sala, R. (2013). Amphibians and squamate reptiles from the late Miocene (Vallesian) of eastern Morocco (Guefaït-1, Jerada Province). Journal of Vertebrate Paleontology, 33, 804–816.
Bocage, J.V.B. Du (1879) Subsidios para a fauna das possessões portuguezas d’Africa occidental. Jornal de sciencias mathematicas, physicas, e naturaes da Academia Real das Sciencas de Lisboa, 7, 85–96.
Bogert, C. M. (1940). Herpetological results of the Vernay Angola Expedition, with notes on African reptiles in other collections. Part I: Snakes, including an arrangement of African Colubridae. Bulletin of the American Museum of Natural History, 77(1), 1–107.
Boie, F. (1827) Bemerkungen über Merrem’s Versuch eines Systems der Amphibien. Erste Lieferung: Ophidier. Isis von Oken, 20, col. 508–566.
Böhme, M., &Vasilyan, D. (2014). Ectothermic vertebrates from the late Middle Miocene of Gratkorn (Austria, Styria). Palaeobiodiversity and Palaeoenvironments, 94, 21–40.
Boulenger, G.A. (1896) Catalogue of the Snakes in the British Museum (Natural History) 3. British Museum (Natural History), London, 727 pp.
Broadley, D. G. (1995). The snouted cobra, Naja annulifera, a valid species in southern Africa. Journal of the Herpetological Association of Africa, 44, 26–32.
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.
Brown, J. H. (1973). Toxicology and Pharmacology of Venoms from Poisonous Snakes (p.81).
Carmona, R., Alba, D.M., Delfino, M., Robles, J.M., Rotgers, C., Bertó Mengual, J.V., Balaguer, J., Galindo, J., & Moyà-Solà, S. (2010). Snake fossil remains from the Middle Miocene stratigraphic series of Abocador de Can Mata (els Hostalets de Pierola, Catalonia, Spain). Cidaris, 30, 77– 84.
Čerňanský, A., Szyndlar, Z., & Mörs, T. (2017). Fossil squamae faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments, 97, 329–354.
Charles C. Thomas. 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.
Christensen, P.A. (1968). The Venoms of Central and South African Snakes. In venoms (Chapter 16, pp. 437-461). Academic Press.
Codrea, V. A., Bordeianu, M., & Venczel, M. (2022). Amphibians and squamate reptiles from the late Miocene of Fălciu (Eastern Romania), Palaeontologia Electronica, 25, a19.
Daxner-Höck, G., Miklas-Tempfer, P.M., Göhlich, U.B., Kuttunen, K., Kazar, E., Nagel, D., Roessner, G.E., Schultz, O., & Ziegler, R. (2004). Marine and terrestrial vertebrates from the Middle Miocene of Grund (Lower Austria). Geologica Carpathica, 55, 1–7.
Depéret, C. (1890–1897). Les animaux Pliocènes du Roussillon, Mémoires de la Société Géologique de France, Paléontologie, Série 3, pp. 1–194.
Eichwald, E. (1831) Zoologia specialis quam expositis animalibus tum vivis, tum fossilibus potissimum Rossiae in universum, et Poloniae in species, in usum lectionum publicarum in Universitate Caesareae Vilnensi habendarum. Part 3. Josephi Zawadski, Vilnius, 396 pp.
Georgalis, G.L., & Delfino, M. (2022). The fossil record of lizards and snakes (Reptilia: Squamata) in Greece. In E. Vlachos (Ed.), Fossil vertebrates of Greece (Vol. 1, pp. 205-235). Springer Nature.
Georgalis, G.L., Rage, J.-C., de Bonis, L., & Koufos, G.D. (2018). Lizards and snakes from the late Miocene hominoid locality of Ravin de la Pluie (Axios Valley, Greece), Swiss Journal of Geosciences, 111, 169–181.
Georgalis, G. L., Villa, A., Ivanov, M., & Delfino, M. (2019). Fossil amphibians and reptiles from the Neogene locality of Maramena (Greece), the most diverse European herpetofauna at the Miocene/Pliocene transition boundary. Palaeontologia Electronica, 22(3), 1–99.
Hallowell, E. (1857) Notice on a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences in Philadelphia by Dr. Henry A. Ford. Proceedings of the Academy of Natural Sciences of Philadelphia, 9, 48–72
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 East African snakes. PLOS Neglected Tropical Diseases. https://doi.org/10.1371/journal.pntd.0005969
Hashmi, M. U. A., Yousufzai, S., & Safi, A. (2024). Assessment of human–snake interaction in the industrial zone of Port Qasim, Malir, Karachi, Pakistan. International Journal of Biology and Biotechnology, 21(3), 369–376.
Hashmi, M. U. A., & Safi, A. (2025). Some aspects of ecology, systematics, threats, and conservation status of the monitor lizard, Varanus bengalensis, from Sindh, Pakistan. International Journal of Biology and Biotechnology, 22(3), 623–629.
Hoser, R. (2009). Naja, Boulengerina, and Paranaja. Australasian Journal of Herpetology, 7, 1–15.
Hoffstetter, R. (1938). Sur la présence d’Elapidae, représentés par un genre nouveau, dans le Néogène français. Comptes Rendus de l’Académie des Sciences, 206, 1916–1918.
Hoffstetter, R. (1939). Contribution à l’étude des Elapidae actuels et fossiles et de l’ostéologie des Ophidiens (Archives du Muséum d’Histoire Naturelle de Lyon, Vol. 15, pp. 1–78).
Ivanov, M. (2000). Snakes of the lower/middle Miocene transition at Vieux Collonges (Rhône, France), with comments on the colonisation of western Europe by colubroids. Geodiversitas, 22, 559–588.
Ivanov, M. (2001). Changes in the composition of the European snake fauna during the Early Miocene and at the Early/Middle Miocene transition. Paläontologische Zeitschrift, 74, 563–573.
Jaen, M. J., & Sanchíz, B. (1985). Fossil snakes from the Pliocene of Layna, central Spain. In Third Ordinary General Meeting of the Societas Herpetologica Europaea: Information, Programme, Abstracts (19–23 August 1985, Prague).
Karl, H. V., Safi, A., & Tichy, G. (2024). Review of fossil records of prehistoric reptiles, their distribution, and paleobiogeographic evolution in Pakistan. Scientific Reports in Life Sciences, 5(3), 1–23. https://doi.org/10.5281/zenodo.13771832
Khare, A. D., Khole, V., & Gade, P. R. (1992). Toxicities, LD₅₀ prediction and in vivo neutralisation of some elapid and viperid venoms. Indian Journal of Experimental Biology, 30(12), 1158–1162.
Kazandjian, T. D. (2021). Convergent evolution of pain-inducing defensive venom components in spitting cobras. Science, 371(6527), 386–390.
Kazemi, E., Nazarizadeh, M., Fatemizadeh, F., Khani, A., & Kaboli, M. (2020). Phylogeny, phylogeography and diversification history of the westernmost Asian cobra (Naja oxiana) in the Trans-Caspian region.
Kazemi-Lomedasht, F., Yamabhai, M., Sabatier, J., Behdani, M., Zareinejad, M. R., & Shahbazzadeh, D. (2019). Development of a human scFv antibody targeting the lethal Iranian cobra (Naja oxiana) snake venom. Toxicon, 171, 78–85.
Latifi, M. (1984). Snakes of Iran. Society for the Study of Amphibians and Reptiles.
Laurent, R. F. (1955). Exploration du Parc National de l’Upemba. Reptiles. Institut des Parcs Nationaux du Congo Belge, 32, 1–125.
Laurenti, J.N. (1768) Austriaci Viennensis Specimen Medicum, Exhibens Synopsin Reptilium Emendatam Cum Experimentis Circa Venena et Antidota Reptilium Austriacorum. J.T. De Trattern, Viennae, 214 pp.
Lee, C. Y., & Tseng, L. F. (1969). Species differences in susceptibility to elapid venoms. Toxicon, 7(2), 89–93. https://doi.org/10.1016/0041-0101(69)90069-5
Lesson, R.P. (1831) Catalogue des reptiles qui font partie d’une collection zoologique recueillie dans l’Inde continentale ou en Afrique et rapportée en France par M. Lamare-Piquot. Bulletin des Sciences Naturelles et de Géologie, Paris, 25, 119–123.
Linnaeus, C. von. (1758) Systema Naturae per Regna Tria Naturae secundum Classes, Ordines, Genera, Species cum Characteribus Differentiis, Synonymis, Locis. Stockholm, Laurentii Salvii, 824 pp.
Lydekker, R. (1886). Indian Tertiary and post-Tertiary vertebrata: The fauna of the Karnul-Caves (Palaeontologia Indica, Vol. 4, p. 23).
Marquina, R., Crespo, V. D., Mansino, S., Fagoaga, A., Ruiz-Sánchez, F. J., Bailon, S., & Montoya, P. (2016). Amphibians and squamates from Late Miocene (MN 13) of Venta del Moro, Spain. In E. Manzanares, H. G. Ferrón, M. Suñer, B. Holgado, V. D. Crespo, S. Mansino, A. Fagoaga, R. Marquina, I. García-Sanz, C. Martínez-Pérez, M. Joanes-Rosés, B. Cascales-Miñana, & M. D. Marín-Monfort (Eds.), 1st International Meeting of Early-stage Researchers in Palaeontology & XIV Encuentro de Jóvenes Investigadores en Paleontología – New perspectives on the evolution of Phanerozoic biotas and ecosystems: Conference proceedings (p. 107).
Mertens, R. (1937). Reptilien und Amphibien aus dem südlichen Inner-Afrika. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 435, 1–23.
Meylan, P. A. (1987). Fossil snakes from Laetoli. In M. D. Leakey & J. M. Harris (Eds.), Laetoli: A Pliocene site in northern Tanzania (pp. 78–82). Oxford University Press.
Miklas-Tempfer, P. M. (2003). The Miocene herpetofaunas of Grund (Caudata; Chelonii, Sauria, Serpentes) and Mühlbach am Manhartsberg (Chelonii, Sauria, Amphisbaenia, Serpentes), Lower Austria. Annalen des Naturhistorischen Museums in Wien, 104A, 195–235.
Minton, S.A. (1974). Venom Diseases. Thomas.
Minton, S.A. (1996). Par'Book. Smithsonian Institution Scholarly Press.
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). In B. A. Huber, B. J. Sinclair, & K.-H. Lampe (Eds.), African biodiversity: Molecules, organisms, ecosystems (pp. 221–228). Museum Koenig.
Naik, B.S. (2017). "Dry bite" in venomous snakes: A review. Toxicon, 133, 63–67.
Müller, F. (1887 [1890]) Fünfter Nachtrag zum Katalog der herpetologischen Sammlung des Basler Museums. Verhandlungen der naturforschenden Gesellschaft in Basel, 8, 249–296.
Orimi, J. R., Eskandarzadeh, N., Sharifabadi, M. A., Miri, V., Aghabeiglooei, Z., & Rezghi, M. (2023). Analyzing the biological traits of snakes in Avicenna's Canon of Medicine and making a comparison with contemporary serpentology. Toxicon, 231 (1), 107198.
Palasuberniam, P., Chan, Y. W., Tan, K. Y., & Tan, C. H. (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. https://doi.org/10.3389/fphar.2021.727756
Peters, W.K.H. (1854) Diagnosen neuer Batrachier, welche zusammen mit der früher (24. Juli und 17. August) gegebenen Übersicht der Schlangen und Eidechsen mitgetheilt werden. Bericht über zur Bekanntmachung geeignete Verhandlungen der Königlichen Preussischen Akademie der Wissenschaften zu Berlin, 1854, 614–628.
Peters, W.K.H. (1861) Eine zweite Übersicht (vergl. Monatsberichte 1859 p. 269) der von Hrn. F. Jagor auf Malacca, Java, Borneo und den Philippinen gesammelten und dem Kgl. Zoologischen Museum übersandten Schlangen. Monatsberichte der Königlichen Preussischen Akademie der Wissenschaften zu Berlin, 1861, 683–691.
Peters, W. C. H. (1876) Eine zweite Mittheilung uber die von Hrn. Professor Dr. R. Buchholz in Westafrica gesammelten Amphibien. Monatsberichte der Koniglich preussischen Akademie der Wissenschaften zu Berlin, 1876, 117 - 123.
Pokrant, F., Kindler, C., Vamberger, M., Smith, K. T., & Fritz, U. (2017): Grass snakes (Natrix natrix, N. astreptophora) mimicking cobras display a ´fossil behavior´. Vertebrate Zoology, 67(2), 261-269.
Rage, J.-C. (1973). Fossil snakes from Olduvai, Tanzania. In L. S. B. Leakey, R. J. G. Savage, & S. C. Coryndon (Eds.), Fossil vertebrates of Africa (Vol. 3, pp. 1–8). Academic Press.
Rage, J.-C. (1976). Les squamates du Miocène de Beni Mellal. Géologie Méditerranéenne, 3, 57–70.
Rage, J.-C. (1979). Les serpents de la Rift Valley: Un aperçu général. Bulletin de la Société Géologique de France, 21, 329–330.
Rage, J.-C. (1982). Amphibia and Squamata. In H. Thomas, S. Sen, M. Khan, B. Battail, & G. Ligabue (Eds.), The Lower Miocene fauna of Al-Sarrar (Eastern Province, Saudi Arabia) (Atlal: Journal of Saudi Arabian Archaeology, Vol. 5, p. 117).
Rage, J.-C. (1984). Serpentes (Handbuch der Paläoherpetologie, Vol. 11). Friedrich Pfeil.
Rage, J.-C. (2003). Squamate reptiles from the early Miocene of Arrisdrift (Namibia). In M. Pickford & B. Senut (Eds.), Geology and palaeobiology of the Central and Southern Namib Desert, Southwestern Africa: Vol. 2. Palaeontology of the Orange River Valley, Namibia (Memoirs of the Geological Survey of Namibia, Vol. 19, pp. 43–50).
Rage, J.-C., & Bailon, S. (2011). Amphibia and Squamata. In T. Harrison (Ed.), Paleontology and geology of Laetoli: Human evolution in context (Vol. 2, pp. 467–478). Springer.
Rage, J.-C., & Şen, Ş. (1976). Les amphibiens et les reptiles du Pliocène supérieur de Çalta (Turquie). Géologie Méditerranéenne, 3, 127–134.
Reinhardt, J.T. (1843) Beskrivelse af nogle nye Slangearter. Det Kongelige Danske Videnskabernes Selskabs Naturvidenskabelige og Mathematiske Afhandlinger, 10, 233–277.
Reis, K. R., & Garong, A. M. (2001). Late Quaternary terrestrial vertebrates from Palawan Island, Philippines. Palaeogeography, Palaeoclimatology, Palaeoecology, 171, 409–421.
Safi, A., Hashmi, M. U. A., Yousufzai, S., Shah, M., Attaullah, Gul, & Karl, H. V. (2024). Study of lacertilian (lizards) diversity of District Dir (Lower), KPK, Pakistan. International Journal of Biological Research, 11(1), 16–27.
Safi, A., & Karl, H.-V. (2024). A review of the updated checklists of the amphibians and reptiles of Pakistan: Present and prehistoric. Journal of Biodiversity and Biotechnology, 4(1), 22–37. https://doi.org/10.20961/jbb.v4i1.106964
Saleh and Trape, Jean-François (2023). Guide des serpents d'Afrique occidentale, centrale et d'Afrique du Nord (in French). Marseille: IRD Éditions. ISBN 978-2-7099-2976-9.
Scortecci, G. (1932) Rettili dello Yemen. Atti della Società Italiana di Scienze Naturali, 71, 39–49.
Cantor, T.E. (1842) General features of Chusan, with remarks on the flora and fauna of that island. Annals and Magazine of Natural History, 1, 265–277.
Sheng-Chao Shi, Gernot Vogel, Li Ding, Ding-Qi Rao, Shuo Liu, Liang Zhang, Zheng-Jun Wu and Ze-Ning Chen. 2022. Description of A New Cobra (Naja Laurenti, 1768; Squamata, Elapidae) from China with Designation of A Neotype for Naja atra. Animals. 12(24); 3481. DOI: 10.3390/ani12243481
Slowinski, J.B. & Wüster, W. (2000) A new cobra (Elapidae: Naja) from Myanmar (Burma). Herpetologica, 56, 257– 270.
Spawls, S., & Branch, B. (1995). The dangerous snakes of Africa (1st ed.). Ralph Curtis Books.
Syromyatnikova, E., Mayda, S., & Tesakov, A. (2022). Late Miocene amphibians and reptiles: New insight into the pre-Messinian herpetofaunas in Turkey. Historical Biology, 34, 1964–1971.
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, 683–689.
Szyndlar, Z. (1985). Ophidian fauna (Reptilia: Serpentes) from the uppermost Miocene of Algora (Spain). Estudios Geológicos, 41, 447–465.
Szyndlar, Z. (1988). Two new extinct species of the genera Malpolon and Vipera (Reptilia, Serpentes) from the Pliocene of Layna (Spain). Acta Zoologica Cracoviensia, 31, 687–706.
Szyndlar, Z. (1991b). A review of Neogene and Quaternary snakes of central and eastern Europe. Part II: Natricinae, Elapidae, Viperidae. Estudios Geológicos, 47, 237–266.
Szyndlar, Z. (1995). The vertebrate locality Maramena (Macedonia, Greece) at the Turolian–Ruscinian boundary (Neogene). 4. Serpentes (Squamata, Reptilia). Münchner Geowissenschaftliche Abhandlungen, Reihe A, 28, 39–42.
Szyndlar, Z. (2005). Snake fauna from the late Miocene of Rudabánya. Palaeontographia Italica, 90, 31–52.
Szyndlar, Z. (2009). Snake fauna (Reptilia: Serpentes) from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 (Germany). Paläontologische Zeitschrift, 83, 55–66.
Szyndlar, Z., & Alférez, F. (2005). Iberian snake fauna of the Early/Middle Miocene transition. Revista Española de Herpetología, 19, 57–70.
Szyndlar, Z., & Rage, J.-C. (1990). West Palearctic cobras of the genus Naja (Serpentes: Elapidae): Interrelationships among extinct and extant species. Amphibia-Reptilia, 11, 385–400.
Szyndlar, Z., & Schleich, H. H. (1993). Description of Miocene snakes from Petersbuch 2 with comments on the Lower and Middle Miocene ophidian faunas of southern Germany (Stuttgarter Beiträge zur Naturkunde, Serie B, No. 192, pp. 1–47).
Szyndlar, Z., & Zerova, G. A. (1990). Neogene cobras of the genus Naja (Serpentes: Elapidae) of East Europe. Annalen des Naturhistorischen Museums Wien, 91A, 53–61.
Taylor, E.H. (1922) The Snakes of the Philippine Islands. Bureau of Printing, Manila, 312 pp.
Tempfer, P. M. (2005). The herpetofauna (Amphibia: Caudata, Anura; Reptilia: Scleroglossa) of the Upper Miocene locality Kohfidisch (Burgenland, Austria). Beiträge zur Paläontologie, 29, 145–253.
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.
Trape, J.F; Collet, M (2020). "A new and remarkable species of semi-aquatic naja (Elapidae, subgenus Boulengerina Dollo, 1886) from the Democratic Republic of Congo". Bulletin de la Société Herpétologique de France. 173: 41–52.
Uetz, P., Freed, P., & Hošek, J. (Eds.). (2024). The Reptile Database. http://www.reptile-database.org
Vasilyan, D., Čerňanský, A., Szyndlar, Z., & Mörs, T. (2022). Amphibian and reptilian fauna from the early Miocene of Echzell, Germany. Fossil Record, 25, 99–145.
Villa, A., Quadros, A. B., Delfino, M., Luján, À. H., Bolet, A., Casanovas-Vilar, I., Robles, J. M., & Alba, D. M. (2024). The rise and fall of the Iberian cobras (Elapidae, Naja) in the context of their European and global fossil record. Papers in Palaeontology, 10(4). https://doi.org/10.1002/spp2.1575
Wall, F. (1913) A popular treatise on common Indian snakes. Naia tripudians (Merrem). The cobra. Journal of the Bombay Natural History Society, 22, 243–259.
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.
Warrell, D. A. (1995). Clinical toxicology of snakebite in Asia. In J. A. Tracey (Ed.), Handbook of clinical toxicology of animal venoms and poisons (1st ed., pp. 493–594). CRC Press. https://doi.org/10.1201/9780203719442-27
Warrell, D. A. (2010). Snakebite. The Lancet, 375(9708), 77–88.
Werner, F. (1902) Über westafrikanische Reptilien. Verhandlungen der Zoologischen und Botanischen Gesellschaft inWien, 52, 332–348.
Watt, G., Theakston, R. D., Hayes, C. G., Yambao, M. L., Sangalang, R., et al. (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), 1444–1448. https://doi.org/10.1056/NEJM198612043152303
Watt, G., Padre, L., Tuazon, L., Theakston, R. D., & Laughlin, L. (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), 306–311. https://doi.org/10.4269/ajtmh.1988.39.306
Wong, K. Y., Tan, C. H., & Tan, N. H. (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.
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. https://doi.org/10.3390/toxins13010060
Wüster, W., & Thorpe, R. S. (1992). Dentitional phenomena in cobras revisited: Fang structure and spitting in the Asiatic species of Naja (Serpentes: Elapidae). Herpetologica, 48, 424–434.
Wüster, W. & Broadley, D.G. (2003) A new species of spitting cobra from northeastern Africa (Serpentes: Elapidae: Naja). Journal of Zoology, London, 259, 345–359.
Wüster, W., Crookes, S., Ineich, I., Mane, 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 (Naja nigricollis complex). Molecular Phylogenetics and Evolution, 45(2), 437–453. https://doi.org/10.1016/j.ympev.2007.07.021
Wüster, Wolfgang; Chirio, Laurent; Trape, Jean-François; Ineich, Ivan; Jackson, Kate; Greenbaum, Eli; Barron, Cesar; Kusamba, Chifundera; Nagy, Zoltán T.; Storey, Richard; Hall, Clara; Wüster, Catharine E.; Barlow, Axel; Broadley, Donald 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). Magnolia Press: 68–98. doi:10.11646/zootaxa.4455.1.3
Wynn, J. G., Alemseged, Z., Bobe, R., Geraads, D., Reed, D., & Roman, D. C. (2006). Geological and palaeontological context of a Pliocene juvenile hominin at Dikika, Ethiopia. Nature, 443, 332–336.
Zhao, E., & Adler, K. (1993). Herpetology of China (1st ed.). Society for the Study of Amphibians and Reptiles.
Online source:
The Reptile Database. (n.d.). http://www.reptile-database.org/
RepFocus. (n.d.). Naja. https://repfocus.dk/Naja.html
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