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The osteology and plumage of Confuciusornis sanctus and Changchengornis hengdaoziensis from the Chaomidianzi Formation (previously referred to as the lower section of the Yixian Formation) of western Liaoning Province (China) are described in detail. Confusiusornis sanctus and Changchengornis hengdaoziensis have toothless, beaked skulls (the tomial crest is straight in the former species and strongly curved in the latter) and retain the dorsal portion of the nasal process of the maxilla. The skull of Confuciusornis sanctus is of typical diapsid plan. It possesses a triradiate postorbital that with the squamosal forms a complete supratemporal arcade.
Furthermore, a robust jugal-postorbital contact completely separates the infratemporal fenestra from the orbit. Although the postorbital region is not preserved in Changchengornis hengdaoziensis, it likely resembles that of Confuciusornis sanctus. Both species have abbreviated tails with long pygostyles, not the long, bony tail originally reconstructed in Confuciusornis sanctus. Scapulae and coracoids are fused to form scapulocorcoids. The coracoids are strutlike but much shorter than the scapulae. The furculae are robust and boomerang shaped. The sterna are long and nearly flat. The complete basket of gastralia follows the sternum caudally. The wing elements are short, and proportions among individual bones are primitive in that the hand is longer than either the humerus or the ulna and the ulna is shorter than the humerus. The pelvis is opisthopubic. The postacetabular wing of the ilium is much shorter than the preacetabular wing. Differences in plumage, namely the presence or absence of two very long tail feathers, are observed among several well-preserved specimens of Confuciusornis sanctus. This difference is likely the expression of sexual dimorphism, although other biological attributes known for extant populations (e.g. differential molting, correlation between sexual maturity and ornamental plumage) indicate that alternative explanations may also account for the observed plumage variation among specimens of Confusiusornis sanctus.
Given the latest Jurassic-earliest Cretaceous age of the Chaomidianzi Formation, Confusiusornis sanctus and Changchengornis hengdaoziensis are surely among the oldest know birds after the Early Tithonian Archaeopteryx lithographica. Confusiusornis sanctus and Changchengornis hengdaoziensis thus furnish the earliest record of beaked birds. The fully diapsid skull of Confusiusornis sanctus, and presumably of Changchengornis hengdaoziensis, and the absence of a bending zone on the base of the snout suggest that earlier interpretations of the skull of Confusiusornis sanctus as prokinetic are incorrect. Confuciusornis sanctus probably had very limited cranial kinetic capabilities, if any. Optimization of the postorbital-jugal contact, a character intimately correlated with intracranial kinesis, in a phylogeny of basal avians indicates that the essentially akinetic condition of the skull of Confuciusornis sanctus is a reversal derived from forms possessing kinetic properties.
Recent recognition of two additional species of Confuciusornis – Confuciusornis chuonzhous and Confuciusornis suniae – are based on anatomical misinterpretations. Thus, Confuciusornis suniae and Confuciusornis chuonzhous are regarded as junior synonyms of Confuciusornis sanctus. Confuciusornis sanctus is the sister-taxon of Changchengornis hengdaoziensis, and both are placed within the Confuciusornithidae. Some of the synapomorphies supporting this grouping include the presence of edentolous jaws, a rostrally forked mandibular symphysis, a reduced claw of manual digit II, and a V-shaped caudal margin of the sternum. The Confuciusornithidae is considered to be the sister-group of a clade composed of the Enantiornithomorpha and the Ornithuromorpha. This phylogenetic interpretation is far more parsimonious than previous systematic hypotheses placing the Confuciusornithidae as either within the Enantiornithes or as its sister-group. Purported climbing specializations of Confuciusornis sanctus are evaluated in light of the available anatomical evidence. We conclude that both Confuciusornis sanctus and Changchengornis hengdaoziensis were ill suited for tree climbing.
The foot of Changchengornis hengdaoziensis, however, suggests a greater grasping ability than that of Confuciusornis sanctus. It is argued that both Confuciusornis sanctus and Changchengornis hengdaoziensis were able to fly and take off from the ground. The remarkable concentration of specimens of Confuciusornis sanctus from a relatively small quarry near the village of Sihetun (Liaoning Province) suggests several events of mass mortality and, perhaps, a gregarious behavior.
