Presence of the large aquatic snake Palaeophis africanus in the middle Eocene marine margin of the Congo Basin, Cabinda, Angola

Ten isolated snake vertebrae from Landana and Sassa-Zao, Cabinda Exclave, Angola, present a “primitive” grade morphology with a weak lateral compression and do not belong to Palaeophis aff. typhaeus as originally referred to. They well belong to a single taxon and are here attributed to Palaeophis africanus for which the intracolumnar variation is described and illustrated. This species is Lutetian (middle Eocene) in age and originates from a marine coastal environment confirming again the aquatic capabilities of palaeophiid snakes. It represents the third largest species of Palaeophis with P. colossaeus and P. maghrebianus to which it is closely related in our tentative phylogenetic analysis, indicating that these three taxa could belong to an African clade. This study also contributes to the debate on the existence of primitive and advanced grades among palaeophiid snakes. Palaeophis presents laterally compressed anterior trunk vertebrae that could have been often erroneously considered as representing advanced grade species and potential parataxonomy.