Primitive forms of dragonflies (Odonata) were the dominant aerial predators and probably dominated terrestrial insect predation as well. , and all are effectively semi-aquatic insects (aquatic immature stages, and terrestrial adults), as are all modern odonates. Their prototypes are the oldest winged fossils, dating back to the Devonian, and are different in several respects from the wings of other insects. Fossils suggest they may have possessed many modern attributes even by the late Carboniferous, and it is possible that they captured small vertebrates, for at least one species had a wing span of . Several other insect groups appeared or flourished during the Permian, including the Coleoptera (beetles), Hemiptera (true bugs) and Orthoptera.