New specimens of Bunaia woodwardi, Clarke 1919 (Euchelicerata): A new member of Offacolidae providing insight supporting the Arachnomorpha

The rapid early diversification of arthropods has made understanding internal relationships within the group fiendish. Particularly unresolved is the origin of Euchelicerata, a clade consisting of the Prosomapoda (comprising the extant Xiphosura and Arachnida and the extinct Chasmataspidida, Eurypterida and synziphosurines) and the extinct Offacolidae. Here we describe new material of the Silurian synziphosurine Bunaia woodwardi that reveals previously unknown features of its ventral anatomy: a pair of elongated chelicerae in the prosoma, followed posteriorly by five pairs of biramous appendages, a first pre-abdomen somite bearing a pair of paddle-like uniramous appendages (exopods), and a ventral pretelsonic process. Phylogenetic analyses retrieve B. woodwardi as an Offacolidae closely related to Setapedites abundantis from the early Ordovician Fezouata Biota. An anatomical comparison of the pretelsonic process of B. woodwardi, also present in Setapedites, with the posterior trunk morphologies of other Offacolidae, Habeliida and Vicissicaudata, suggests a possible homologous appendicular origin. This proposed apomorphic character supports a monophyletic Arachnomorpha, formed of Vicissicaudata, Habeliida and Euchelicerata. The establishment of this new homology could help to clarify the highly enigmatic phylogeny at the base of the euchelicerates as well as the sequence of character acquisition during their early evolution.