Redescription of Yinostius major (Arthrodira: Heterostiidae) from the Lower Devonian of China, and the interrelationships of Brachythoraci

Yinosteus major is a heterostiid arthrodire (Placodermi) from the Lower Devonian Jiucheng Formation of Yunnan Province, south-western China. A detailed redescription of this taxon reveals the morphology of neurocranium and visceral side of skull roof. Yinosteus major shows typical heterostiid characters such as anterodorsally positioned small orbits and rod-like anterior lateral plates. Its neurocranium resembles those of advanced eubrachythoracids rather than basal brachythoracids, and provides new morphological aspects in heterostiids. Phylogenetic analysis based on parsimony was conducted using a revised and expanded data matrix. The analysis yields a novel scenario on the brachythoracid interrelationships, which assigns Heterostiidae (including Heterostius ingens and Yinosteus major) as the sister group of Dunkleosteus amblyodoratus. The resulting phylogenetic scenario suggests that eubrachythoracids underwent a rapid diversification during the Emsian, representing the placoderm response to the Devonian Nekton Revolution. The instability of the relationships between major eubrachythoracid clades might have a connection to their longer ghost lineages than previous scenarios have implied.