Many species of the second major avialan lineage to diversify, the Euornithes (meaning "true birds", because they include the ancestors of modern birds), were semi-aquatic and specialized in eating fish and other small aquatic organisms. Unlike the enantiornithes, which dominated land-based and arboreal habitats, most early euornithes lacked perching adaptations and seem to have included shorebird-like species, waders, and swimming and diving species. The later included the superficially gull-like Ichthyornis, the Hesperornithiformes, which became so well adapted to hunting fish in marine environments that they lost the ability to fly and became primarily aquatic. The early euornithes also saw the development of many traits associated with modern birds, like strongly keeled breastbones, toothless, beaked portions of their jaws (though most non-avian euornithes retained teeth in other parts of the jaws). Euornithes also included the first avialans to develop true pygostyle and a fully mobile fan of tail feathers, which may have replaced the "hind wing" as the primary mode of aerial maneuverability and braking in flight.