Placoderms (Armored Fish): Dominant Vertebrates of the Devonian Period

Placoderms, the most diverse group of Devonian fishes, were globally distributed in all habitable freshwater and marine environments, like teleost fishes in the modern fauna. Their known evolutionary history (Early Silurian–Late Devonian) spanned at least 70 million years. Known diversity (335 genera) will increase when diverse assemblages from new areas are described. Placoderms first occur in the Early Silurian of China, but their diversity remained low until their main evolutionary radiation in the Early Devonian, after which they became the dominant vertebrates of Devonian seas. Most current placoderm data are derived from the second half of the group's evolutionary history, and recent claims that they form a paraphyletic group are based on highly derived Late Devonian forms; 16 shared derived characters are proposed here to support placoderm monophyly. Interrelationships of seven placoderm orders are unresolved because Silurian forms from China are still poorly known. The relationship of placoderms to the two major extant groups of jawed fishes—osteichthyans (bony fishes) and chondrichthyans (cartilaginous sharks, rays, and chimaeras)—remains uncertain, but the detailed preservation of placoderm internal braincase structures provides insights into the ancestral gnathostome (jawed vertebrate) condition. Placoderms provide the most complex morphological and biogeographic data set for the Middle Paleozoic; marked discrepancies in stratigraphic occurrence between different continental regions indicate strongly endemic faunas that were probably constrained by marine barriers until changes in paleogeography permitted range enlargement into new areas. Placoderm distributions in time and space indicate major faunal interchange between Gondwana and Laurussia near the Frasnian-Famennian boundary; closure of the Devonian equatorial ocean is a possible explanation.