Anatomy of the primitive bird Sapeornis chaoyangensis from the Early Cretaceous of Liaoning, China

Two new, nearly completely articulated skeletons of Sapeornis chaoyangensis provide much new information about the anatomy of this basal avian, particularly in the skull, pectoral girdle, forelimb, and hind limb. This new material shows that the hand of Sapeornis, with a phalangeal formula of 2-3-2, was more derived than previously reconstructed. The skeleton of Sapeornis has several unique features, such as a distinctively elongated fenestra. on the proximal end of the humerus, a robust furcula with a distinctive hypocleidum, and an elongated forelimb. Sapeornis exhibits a combination of derived and primitive features, including a short, robust non-strut-like coracoid and a fibula reaching the distal end of the tarsal joint (as in Archaeopteryx), a pygostyle, reduced manual digits, and a well-fused carpometacarpus (as in more advanced birds). These features further indicate the mosaic pattern in the early evolution of birds and confirm the basal position of Sapeornis near Archaeopteryx and Jeholornic in the phylogeny of early birds. The preservation of gastroliths in one of the new specimens also represents the first Chinese Mesozoic bird with such evidence, indicating a herbivorous feeding habit and providing further evidence for our understanding of the diet diversification in early avian evolution.