A new adult specimen of the basalmost ornithuromorph bird Archaeorhynchus spathula (Aves: Ornithuromorpha) and its implications for early avian ontogeny

Most living birds characteristically grow rapidly and reach adult size within a year. Nevertheless, little is known about how such an advanced developmental strategy evolved despite many discoveries of early fossil birds. Here we assess the long-bone histology from a new adult specimen of Archaeorhynchus spathula, the basalmost taxon of Ornithuromorpha. Ornithuromorpha is the most inclusive clade containing extant birds but not the Mesozoic Enantiornithes. Histological analysis reveals that the cortex is composed of parallel-fibred bone with three lines of arrested growth, indicative of slow and annually interrupted growth for this taxon. Such bone histology is significantly different from that of other known basal ornithuromorphs, but resembles that of enantiornithines, which leads us to suggest protracted slow growth in the common ancestor of Ornithuromorpha and Enantiornithes. The fusion sequence of the tarsometatarsus between Enantiornithes and Ornithuromorpha has long been hypothesized to be different and regarded as indicative that enantiornithines are not close relatives of ornithuromorphs. Due to a lack of fossils recording early ontogenetic stages, little is known about the development of the tarsometatarsus in basal ornithuromorphs, making this hypothesis impossible to test. Here we show that the fusion sequence of the tarsometatarsus in Archaeorhynchus is similar to that of enantiornithines, and that the proximal-early fusion in tarsometatarsus represents a plesiomorphic trait for basal birds. Our findings also shed light on ontogenetic variation of sternal morphology, which highlights the importance of ontogeny in the taxonomic and phylogenetic study of early birds.