Fossils of early vertebrates and the evolution of the gnathostome face revealed by Synchrotron imaging

Cyclostomes and gnathostomes have distinct face patterns. Cyclostomes possess a median nasohypophysial duct, an anterior hypophysis and a short telencephalon, contra gnathostomes possessing a pair of nasal sacs opening externally, a separate posterior hypophysis opening onto the palate and a long telencephalon. Embryonic development also differs. In cyclostomes, premandibular crest cells migrate forwards either side of the nasohypophysial placode, forming an upper lip; in gnathostomes they migrate between the hypophysial and nasal placodes forming the trabecular region. Supraoptic neural crest remains posterior to the nasohypophysial duct in cyclostomes but moves forward to create the nasal capsules in gnathostomes. Fossil stem gnathostomes illustrate a transitional sequence between these two patterns: 1) The galeaspid Shuyu (jawless stem gnathostome): nasohypophysial duct, short telencephalon, and anteriorly oriented hypophysis as in a cyclostome, but paired nasal sacs and hypophysis separated by a rudimentary trabecula. 2) The primitive placoderm Romundina (jawed stem gnathostome): short telencephalon, anteriorly directed hypophysis, trabecular region long and wide, nasal capsule located far behind the tip of the snout but just in front of the orbits. These features are interpreted as uniquely primitive among gnathostomes. The trabeculae of Romundina form an extensive precerebral region resembling the upper lip of extant cyclostomes and Shuyu. The position of the nasal capsule suggests that the supraoptic crest had not migrated forwards. 3) The arthrodire Kujdanowiapsis (a more derived placoderm): short telencephalon and vertically oriented hypophysis. The trabecula has been shortened anteriorly, making the nasal capsule terminal. These positional relationships are maintained in crown gnathostomes.