Holocephali

Holocephali (sometimes spelled Holocephala; Greek for "complete head" in reference to the fusion of upper jaw with the skull) is a subclass of cartilaginous fish. The only living holocephalans are the three families of chimaeras, but the group also includes many extinct members and was more diverse during the Paleozoic and Mesozoic eras. The earliest known fossils of holocephalans date to the Middle Devonian, and the subclass likely reached its peak diversity during the following Carboniferous Period. Molecular clock studies suggest that holocephalans diverged from their closest relatives, elasmobranchs such as sharks and rays, during the Early Devonian or the Silurian Period.

Extinct holocephalans are typically divided into a number of orders, although the relationships between these groups are poorly understood. Several different definitions of Holocephali exist, with the group sometimes considered a less inclusive clade within the larger subclasses Euchondrocephali or Subterbranchialia and with its members spread into the now obsolete groups Paraselachimorpha or Bradyodonti. Per these classification schemes, the name Holocephali is used only for chimaeras and their closest relatives. Recent research has suggested that the orders Cladoselachiformes and Symmoriiformes, which were historically considered relatives or ancestors of sharks, should instead be considered holocephalans. Information on the evolution and relationships of extinct holocephalans is limited, however, because most are known only from isolated teeth or dorsal fin spines, which form much of the basis of their classification.