A new reconstruction of onychoselache traquairi, comments on early chondrichthyan pectoral girdles and hybodontiform phylogeny

A new, third, specimen of Onychoselache traquairi from the Visean (Holkerian) of Scotland allows a significant revision of the anatomy of this stem-group elasmobranch. This first report of material from the Mumbie Quarry exposure of the Glencartholm fish beds presents a new reconstruction of Onychoselache showing broad-based cephalic and nuchal spines, and exceptionally large pectoral fins. Details of the jaws, braincase and postcranial skeleton demonstrate that Onychoselache is a well-characterized member of the Hybodontiformes. Comparisons of the pectoral skeleton with other early chondrichthyan examples, including new material of Tristychius arcuatus and Plesioselachus macracanthus, highlight a range of early chondrichthyan conditions that are incorporated into a revised hybodontiform phylogeny. Close resemblance between Onychoselache and Mesozoic and late Palaeozoic hybodonts implies that these clades diverged within the Carboniferous and Permian. Major differences between Onychoselache and the coeval Tristychius (a modified reconstruction of which is included) indicate that the Neoselachii-Hybodontiformes split is probably Late Devonian, consistent with records of isolated teeth. The pectoral fins of Onychoselache, while unique among Palaeozoic forms, resemble those of Recent bamboo and epaulette sharks (Orectolobiformes). The functional corollary of this convergence is that Onychoselache represents an instance of a non-tetrapod early vertebrate with a near-walking gait.