Three-dimensional histology of tooth cushions of Lophosteus from the Upper Silurian of Estonia

• Lophosteus superbus from the Late Silurian of Estonia is one of the oldest and most plesiomorphic osteichthyans described to date. Unfortunately at present it is known only from fragmented dermal microremains. The affinities of Lophosteus are therefore controversial with the taxon placed as either basal to both actinopterygians and sarcopterygians, or ambiguously linked to either placoderms or acanthodians. To confound matters further, the character states diagnosing actinopterygians and sarcopterygians have recently been brought into question, and even monophyly of the traditional placoderm and acanthodian clades has been challenged. As a possible stem osteichthyan, Lophosteus could thus be central to our understanding of early gnathostome evolution and the origin of the osteichthyan body plan. Often the best-preserved, although incompletely documented, elements of Lophosteus are tooth cushions. These tooth-bearing arched ossicles could arguably be homologous with the parasymphysial tooth whorls in chondrichthyans, acanthodians, and sarcopterygians, or even the parasymphysial tooth plates in tetrapodomorphs. High-resolution synchrotron scans of 6 isolated tooth cushions from the Upper Silurian of Estonia has permitted a detailed reconstruction of their three-dimensional architecture. The absence of an enamel layer and the presence of large hollows (bigger than normal osteocytes) in the deepest lamellar layer confirmed assignment of the specimens to Lophosteus. The external surface displays irregularly distributed denticles and there are large parallel vessels running horizontally on the basal bone that feed the denticle rows internally. The odontodes have two distinct generations (with successive odontodes accreted between those of the preceding buried generation) and are organised in a similar manner to those found on Lophosteus scales. This new histological data on vascularization provides insight into early gnathostome tooth patterning and could contribute to future phylogenetic assessments.

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