Squamation and scale morphology at the root of jawed vertebrates

Placoderms, as the earliest branching jawed vertebrates, are crucial to understanding how the characters of crown gnathostomes comprising Chondrichthyes and Osteichthyes evolved from their stem relatives. Despite the growing knowledge of the anatomy and diversity of placoderms over the past decade, the dermal scales of placoderms are predominantly known from isolated material, either morphologically or histologically, resulting in their squamation being poorly understood. Here we provide a comprehensive description of the squamation and scale morphology of a primitive taxon of Antiarcha (a clade at the root of jawed vertebrates), Parayunnanolepis xitunensis, based on the virtual restoration of an articulated specimen by using X-ray computed tomography. Thirteen morphotypes of scales are classified to exhibit how the morphology changes with their position on the body in primitive antiarchs, based on which nine areas of the post-thoracic body are distinguished to show their scale variations in the dorsal, flank, ventral, and caudal lobe regions. In this study, the histological structure of yunnanolepidoid scales is described for the first time based on disarticulated scales from the type locality and horizon of P. xitunensis. The results demonstrate that yunnanolepidoid scales are remarkably different from their dermal plates as well as euantiarch scales in lack of a well-developed middle layer. Together, our study reveals that the high regionalization of squamation and the bipartite histological structure of scales might be plesiomorphic for antiarchs, and jawed vertebrates in general.