A new confuciusornithid bird with a secondary epiphyseal ossification reveals phylogenetic changes in confuciusornithid flight mode

The confuciusornithids are the earliest known beaked birds, and constitute the only species-rich clade of Early Cretaceous pygostylian birds that existed prior to the cladogenesis of Ornithothoraces. Here, we report a new confuciusornithid species from the Lower Cretaceous of western Liaoning, northeastern China. Compared to other confuciusornithids, this new species and the recently reported Yangavis confucii both show evidence of stronger flight capability, although the wings of the two taxa differ from one another in many respects. Our aerodynamic analyses under phylogeny indicate that varying modes of flight adaptation emerged across the diversity of confuciusornithids, and to a lesser degree over the course of their ontogeny, and specifically suggest that both a trend towards improved flight capability and a change in flight strategy occurred in confuciusornithid evolution. The new confuciusornithid differs most saliently from other Mesozoic birds in having an extra cushion-like bone in the first digit of the wing, a highly unusual feature that may have helped to meet the functional demands of flight at a stage when skeletal growth was still incomplete. The new find strikingly exemplifies the morphological, developmental and functional diversity of the first beaked birds.