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Phylogenetic Analysis of Old World Viperid Snakes (Serpentes, Viperidae) Based on Skeletal Morphology

P. B. Snetkov, N. L. Orlov


The system of the family Viperidae remains controversial despite the fact that numerous studies had place during last years. The results of different analyses most of which are based on molecular data are conflicting. We conducted the phylogenetic analysis including 39 osteological characters and 31 species of Old World viperid snakes. According to these results, genera Azemiops and Causus do not belong to subfamily Viperinae, so their separation into distinct subfamilies is supported. The validity of subfamily Azemiopinae is confirmed by molecular data. The common primitive character of Azemiops and Causus is joining of the prefrontals to the lateral parts of the frontals, while in the true vipers (Viperinae) the prefrontals join to the anterior margins of the frontals and the contact between these bones nearly reaches the midline of the skull. This type of articulation and the particular shape of the prefrontal can be added to the diagnosis of subfamily Viperinae (without Azemiops and Causus). One more result of our analysis is that the most basal form among true vipers is genus Cerastes. Other true vipers are divided into two groups. Inside the first group, including genera Atheris, Bitis, Echis, and Vipera, the genera Atheris and Bitis form a clade what is consistent with results of some molecular studies. The second group includes genera Daboia, Macrovipera, Montivipera, and Pseudocerastes. In subfamily Crotalinae the clade including genera Calloselasma and Hypnale is strongly supported, that confirms the results of many molecular studies. Protobothrops mangshanensis is close to two other species of genus Protobothrops that is not in conflict with referring P. mangshanensis (originally Zhaoermia mangshanensis) to this genus.


Viperidae; true vipers; pitvipers; phylogeny; osteological phylogenetic analysis

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