Broadley, D.G. & W. Wüster (2004) A review of the southern African ‘non-spitting’ cobras (Serpentes: Elapidae: Naja). African Journal of Herpetology 53:101-122. (original) (raw)
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Zootaxa, 2018
Cobras are among the most widely known venomous snakes, and yet their taxonomy remains incompletely understood, particularly in Africa. Here, we use a combination of mitochondrial and nuclear gene sequences and morphological data to diagnose species limits within the African forest cobra, Naja (Boulengerina) melanoleuca. Mitochondrial DNA sequences reveal deep divergences within this taxon. Congruent patterns of variation in mtDNA, nuclear genes and morphology support the recognition of five separate species, confirming the species status of N. subfulva and N. peroescobari, and revealing two previously unnamed West African species, which are described as new: Naja (Boulengerina) guineensis sp. nov. Broadley, Trape, Chirio, Ineich & Wüster, from the Upper Guinea forest of West Africa, and Naja (Boulengerina) savannula sp. nov. Broadley, Trape, Chirio & Wüster, a banded form from the savanna-forest mosaic of the Guinea and Sudanian savannas of West Africa. The discovery of cryptic diversity in this iconic group highlights our limited understanding of tropical African biodiversity, hindering our ability to conserve it effectively.
A new species of spitting cobra Naja nubiae sp. nov. is described from north-eastern Africa. The distinctiveness of the new species is confirmed by multivariate analysis of pattern and scalation data. Phylogenetic analysis of mitochondrial DNA sequences reveals the new species to be the sister taxon of N. pallida, but with considerable levels of sequence divergence relative to that species. The populations concerned had previously been assigned to N. pallida. The new species differs from N. pallida principally in having more than one dark band across the neck and under the throat, as well as a pair of spots under the throat. It occupies a disjunct range across Egypt, the Sudan, Chad, Niger and Eritrea, where it seems to occupy primarily relatively mesic habitats. Naja mossambica is more closely related to N. nigricollis than to N. pallida and the new species. A key to the African species of Naja is presented.
Molecular phylogenetics and evolution, 2007
We use phylogenetic analysis of 1333 bp of mitochondrial DNA sequence to investigate the phylogeny and historical biogeography of the cobra-like elapid snakes, with special reference to the evolution of spitting and the phylogeography of the African spitting cobras, a radiation widespread in open vegetational formations throughout sub-Saharan Africa. Our results suggest that spitting adaptations appear to have evolved three times in cobras, but alternative scenarios cannot be rejected. The Asiatic Naja are monophyletic and originate from a single colonization of Asia from Africa. The radiation of the African spitting Naja appears to date back to the early Miocene and many speciation events in the group predate the Pliocene expansion of grasslands and the radiation of large grazing mammals in Africa. The cladogenic events in this complex appear to have been triggered by both ecological changes and tectonic events associated with the formation and expansion of the African Rift Valley....
We use a combination of phylogenetic analysis of mtDNA sequences and multivariate morphometrics to investigate the phylogeography and systematics of the Egyptian cobra (Naja haje) species complex. Phylogenetic analysis of mitochondrial haplotypes reveals a highly distinct clade of haplotypes from the Sudano-Sahelian savanna belt of West Africa, and that the haplotypes of Naja haje arabica form the sister group of North and East African N. h. haje. Multivariate morphometrics confirm the distinctness of the Arabian populations, which are consequently recognised as a full species, Naja arabica Scortecci. The Sudano-Sahelian populations are also found to represent a morphologically distinct taxon, and thus a separate species, which we describe as Naja senegalensis sp. nov. The new species differs from all other members of the N. haje complex by a combination of colour pattern and scalation characteristics (especially higher numbers of scale rows around the neck), and the possession of a unique clade of mtDNA haplotypes. The distribution of the new species includes savanna areas of West Africa, from Senegal to western Niger and Nigeria.
A new species of spitting cobra (Naja) from north-eastern Africa (Serpentes: Elapidae)
Journal of Zoology, 2003
A new species of spitting cobra Naja nubiae sp. nov. is described from north-eastern Africa. The distinctiveness of the new species is confirmed by multivariate analysis of pattern and scalation data. Phylogenetic analysis of mitochondrial DNA sequences reveals the new species to be the sister taxon of N. pallida, but with considerable levels of sequence divergence relative to that species. The populations concerned had previously been assigned to N. pallida. The new species differs from N. pallida principally in having more than one dark band across the neck and under the throat, as well as a pair of spots under the throat. It occupies a disjunct range across Egypt, the Sudan, Chad, Niger and Eritrea, where it seems to occupy primarily relatively mesic habitats. Naja mossambica is more closely related to N. nigricollis than to N. pallida and the new species. A key to the African species of Naja is presented.
Ecology of cobras from southern Africa
Journal of Zoology, 2007
Large slender-bodied snakes that forage actively for a generalized array of small vertebrates are conspicuous elements of the terrestrial snake fauna of most continents; the venomous elapid species fill this role in much of Asia, Africa and Australia. Our dissections of eight species of cobras from southern Africa Aspidelaps, Hemachatus, Naja; Serpentes and Elapidae (total of 1290 specimens) provide extensive data on sexual dimorphism, reproductive biology and food habits. Females grow larger than males in Aspidelaps lubricus and Naja nigricincta, but (perhaps reflecting selection on male body size due to male-male combat) males grow as large as females in Naja anchietae, Naja melanoleuca, Naja mossambica, Naja nivea and Hemachatus haemachatus, and males grow larger than females in Naja annulifera. Overall, the degree of male size superiority is higher in species with a larger absolute mean adult body size. Male cobras typically have larger heads and longer tails than conspecific females. Fecundity increases with maternal body size, and is higher in the viviparous rhinkals H. haemachatus than in the oviparous Naja species studied. Diets are broad in all eight species, comprising a wide variety of amphibians, reptiles, mammals and (less often) birds. Ontogenetic (size-related) shifts in dietary composition (amphibian to reptile to mammal) are significant within some taxa (N. annulifera, N. nigricincta) but absent in others (notably N. nivea, the most arid-adapted species). Overall, despite substantial interspecific variation among the eight study species, strong parallels are evident between the cobras of southern Africa and their ecological counterparts in other continents.
The genus Naja Laurenti, 1768, is partitioned into four subgenera. The typical form is restricted to 11 Asian species. The name Uraeus Wagler, 1830, is revived for a group of four non-spitting cobras inhabiting savannas and open formations of Africa and Arabia, while Boulengerina Dollo, 1886, is applied to four non-spitting African species of forest cobras, including terrestrial, aquatic and semi-fossorial forms. A new subgenus is erected for seven species of African spitting cobras. We recommend the subgenus rank as a way of maximising the phylogenetic information content of classifications while retaining nomenclatural stability.
Vertebrate Zoology
The morphological variation, extended distribution, and sequence divergence of a recently described of cobra Naja fuxiShi et al., 2022 captured from mountainous areas in Thailand are evaluated by using molecular and morphological analyses. We investigated the genetic variation and affinities of 72 specimens in the genus Naja by using mitochondrial DNA (cytochrome b and control region) and the nuclear DNA gene, C-mos. Morphological examination was conducted for 33 cobra specimens obtained from the northern, western, and north-eastern regions, and data on their natural history were gathered during field surveys. A high degree of genetic differentiation was shown to exist between the cobras collected from lowlands and those from mountainous areas. N. fuxi occurs in uplands bordering Thailand’s Central Basin, whereas the similar looking N. kaouthia Lesson, 1831 is more or less restricted to the lowlands. All phylogenetic and network analyses supported a distinct clade of N. fuxi from no...