Reappraisal of the morphology and phylogenetic relationships of the alligatoroid crocodylian Diplocynodon hantoniensis from the late Eocene of the United Kingdom

Diplocynodon is a genus of basal alligatoroid comprising nine species, which spanned the late Palaeocene to middle Miocene of Europe. Despite recent revisions of most Diplocynodon species, one of the earliest named and most complete, Diplocynodon hantoniensis, has not been re-described for over 150 years. This species is known from the remains of numerous individuals from the Priabonian (late Eocene) Headon Hill Formation, which crops out at Hordle (Hordwell) Cliff in Hampshire, United Kingdom. Here we re-describe and diagnose Diplocynodon hantoniensis, providing the first detailed description of postcranial anatomy in Diplocynodon, and indeed any basal alligatoroid. Diplocynodon hantoniensis is diagnosed by four autapomorphies, including retention of the ectopterygoid-pterygoid flexure through ontogeny and a unique anterior process of the ectopterygoid adjacent to the posteriormost maxillary alveoli. A critical review of previously referred remains from elsewhere in Europe and the USA restricts Diplocynodon hantoniensis to the late Eocene of the UK. Through comparisons with extant crocodylians, the well- preserved postcranial skeleton enables the interpretation of numerous muscle attachments in the forelimbs and hindlimbs, providing a potentially rich source of character data for future phylogenetic analyses. Based on a comparison of humeral morphology between a large sample of crocodylian species, we outline two new morphological characters in the humerus. We include D. hantoniensis in a phylogenetic analysis, including all putative Diplocynodon species (103 taxa scored for 187 characters). We use four different character-weighting schemes: equal weighting, implied weighting (k value = 8) and extended implied weighting with k-values of 4 and 8. In general, these weighted analyses produce congruent results with the equal-weights analysis, and increase the resolution within Diplocynodon. We recover a monophyletic Diplocynodon in three of the four analyses. However, the fourth analysis, with the strongest downweighting of homoplastic characters and missing data (extended implied weighting with k = 4), recovers the Palaeocene Diplocynodon remensis outside Diplocynodon. Our comprehensive revision of one of the most completely known Diplocynodon species facilitates comparisons in the genus, as well as between other basal alligatoroids, and forms the basis for comparing postcranial anatomy in other fossil crocodylians.