West Palearctic cobras of the genus Naja (Serpentes: Elapidae): interrelationships among extinct and extant species

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Abstract
Some basic osteological cranial features of living and fossil members of the genus Naja are described. The extinct genus Palaeonaja Hoffstetter, 1939, is synonymized with the modern Naja Laurenti, 1768, and the extinct species Palaeonaja crassa Hoffstetter, 1939, is synonymized with Naja romani (Hoffstetter, 1939). Anatomically, the genus Naja can be divided into two main complexes, composed of: (1) living African species, N. antiqua from the Moroccan Miocene, and N. iberica from the Spanish Miocene; (2) living Asiatic species and N. romani from the Miocene of France, Austria, and Ukraine. Living members of the Asiatic complex make up a monophyletic group; they belong to at least three distinct lineages: N. oxiana, N. naja s.s. ( = N. naja naja), and the remaining taxa named here informally the 'East Asiatic Naja'. The African complex is thought to be most primitive and perhaps paraphyletic; Africa is presumed to be the centre of earliest radiation of the genus. The precise relationships of Walterinnesia, a close relative of Naja occupying the area between Asiatic and African ranges of Naja, remain unclear.

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