An unusual bird (Theropoda, Avialae) from the Early Cretaceous of Japan suggests complex evolutionary history of basal birds

The Early Cretaceous basal birds were known largely from just two-dimensionally preserved specimens from north-eastern China (Jehol Biota), which has hindered our understanding of the early evolution of birds. Here, we present a three-dimensionally-preserved skeleton (FPDM-V-9769) of a basal bird from the Early Cretaceous of Fukui, central Japan. Unique features in the pygostyle and humerus allow the assignment of FPDM-V-9769 to a new taxon, Fukuipteryx prima. FPDM-V-9769 exhibits a set of features comparable to that of other basalmost birds including Archaeopteryx. Osteohistological analyses indicate that FPDM-V-9769 is subadult. Phylogenetic analyses resolve F. prima as a non-ornithothoracine avialan basal to Jeholornis and outgroup of the Pygostylia. This phylogenetic result may imply a complex evolutionary history of basal birds. To our knowledge, FPDM-V-9769 represents the first record of the Early Cretaceous non-ornithothoracine avialan outside of the Jehol Biota and increases our understanding of their diversity and distribution during the time.