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Summary: The myelin sheath was a transformative vertebrate acquisition, enabling great increases in impulse propagation velocity along axons. Not all vertebrates possess myelinated axons, however, and when myelin first appeared in the vertebrate lineage is an important open question. It has been suggested that the dual, apparently unrelated acquisitions of myelin and the hinged jaw were actually coupled in evolution . If so, it would be expected that myelin was first acquired during the Devonian period by the oldest jawed fish, the placoderms . Although myelin itself is not retained in the fossil record, within the skulls of fossilized Paleozoic vertebrate fish are exquisitely preserved imprints of cranial nerves and the foramina they traversed. Examination of these structures now suggests how the nerves functioned in vivo. In placoderms, the first hinge-jawed fish, oculomotor nerve diameters remained constant, but nerve lengths were ten times longer than in the jawless osteostraci. We infer that to accommodate this ten-fold increase in length, while maintaining a constant diameter, the oculomotor system in placoderms must have been myelinated to function as a rapidly conducting motor pathway. Placoderms were the first fish with hinged jaws and some can grow to formidable lengths, requiring a rapid conduction system, so it is highly likely that they were the first organisms with myelinated axons in the craniate lineage. [Copyright &y& Elsevier]