Delta wing design in earliest nektonic vertebrates.

The colonization of the pelagic realm by the vertebrates represents one of the major transitions in the evolutionary success of the group and in the establishment of modern complex marine ecosystem. It has been traditionally related with the Devonian rise of jawed vertebrates, but new evidences indicate that first active swimmers, invading the water column, occurred within earlier armoured jawless fishes ("ostracoderms"). These "primitive" fishes lacked conventional fish control surfaces and the precise mechanism used to generate lift and stabilizing forces still remains unclear. We show that, because of their shape, the rigid cephalic shield of Pteraspidiformes, a group of Silurian-Devonian "ostracoderms", generate significant forces for hydrodynamic lift. Particle Image Velocimetry and force measurements in a water channel shows that the flow over real-sized Pteraspidiformes models is similar to that over delta wings, dominated by the formation of leading-edge vortices resulting in enhanced vortex lift forces and delayed stall angles of attack. Additionally, experiments simulating ground effect show that Pteraspidiformes present better hydrodynamic performance under fully pelagic conditions than in a benthic scenario. This suggests that, lacking movable appendages other than the caudal fin, leading-edge vortices were exploited by earliest vertebrates to colonize the water column more than 400 Mya.