Early Jawless Vertebrates and Cyclostome Origins

Undoubted fossil lampreys are recorded since the Late Devonian (358 Ma), and probable fossil hagfishes since the Late Carboniferous (300 Ma), but molecular clock data suggest a much earlier divergence times for the two groups. In the early 20(th) century, hagfishes and lampreys were generally thought to have diverged much later from unknown ancestral cyclostomes, in turn derived through 'degeneracy' from some Paleozoic armored jawless vertebrates, or 'ostracoderms.' However, current vertebrate phylogenies suggest that most, if not all, 'ostracoderms' are in fact jawless stem gnathostomes, which retain certain features that were once regarded as unique to the cyclostomes, such as gill pouches or lack of horizontal semicircular canal. The dorsal, median, nasohypophysial complex of osteostracans has been regarded as identical and homologous to that of lampreys, but recent investigation (notably on the galeaspid braincase) now suggests that this resemblance is in fact a convergence. The anatomy and physiology of lampreys and hagfishes are so different that it is difficult to reconstruct an ancestral morphotype of the cyclostomes, assuming that they are a clade, and there is no clear evidence of any fossil taxon that is neither a fossil hagfish nor a fossil lamprey, but would be more closely related to the cyclostomes than to the gnathostomes. A possible exception is the Silurian-Devonian euphaneropids (or 'naked anaspids').