Structure, Growth and Histology of Gnathal Elements in Dunkleosteus (Arthrodira, Placodermi), with a Description of a New Species from the Famennian (Upper Devonian) of the Tver Region (North-Western Russia)

A new species of Dunkleosteus, D. tuderensis sp. nov., is named based on an infragnathal from the Famennian of the Tver Region, Russia. CT scanning of the holotype revealed two high-density bony constituents comparable in position and interrelations to components described for coccosteomorph arthrodires, supported by the presence of at least two clusters of large vascular canals marking separate arterial supplies. Coccosteomorph and dunkleosteid pachyosteomorphs exhibit similar growth patterns including labio-basal depositions of vascularized bone in the infragnathals and basally in the supragnathals. In contrast to coccosteomorphs, dunkleosteid reinforcement of the occlusal margins occurred via the formation of dense osteonal bone, in parallel with resorption forming extensive lingual fossae. Active bone remodeling proceeded without a complete reworking of the primary osteonal bone structure and the original arrangement of vascular canals. Due to inconsistent anatomical terminology in gnathal elements of dunkleosteid arthrodires, a revised terminology is suggested and new terms are introduced.