The evolution of the wrist bones in the lineage leading to birds.
Incomplete coloring (striped) indicates uncertain identification. A separate ossification of the intermedium (orange) is rarely observed in dinosaurs, but when present, it is seen closely appressed or fused to the radiale (purple). In Maniraptora, a single ossification is present that is commonly referred to as the radiale. However, in birds it develops from a composite radiale+intermedium cartilage and is referred to as the scapholunare. Thus, we propose the use of the term scapholunare for this ossification in bird-like dinosaurs (purple–orange). The distal-anterior ossification of birds (yellow-green) is homologous to the composite semilunate of dinosaurs. In early dinosaurs and most basal theropods, distal carpal 1 (yellow) and 2 (green) were separate bones. The semilunate bone of maniraptoran dinosaurs such as Deinonychus antirrhopus covered the proximal ends of metacarpal 1 and 2, and is thus considered to be a composite of dc1+dc2. This is consistent with our new developmental evidence that this bone in modern birds develops from a composite cartilage (Figure 4). Dc1 of Guanlong (uncertain, incomplete yellow) could arguably be a semilunate (dc1+dc2). Birds re-evolved a large, ossified pisiform (red). The pisiform and the ulnare were present in early dinosaurs, but thereafter they are not preserved, suggesting that if not absent, they were very small or failed to ossify, consistently escaping preservation. In birds, developmental evidence conclusively demonstrates that the ulnare is lost, but the pisiform is present. A large pisiform is frequently preserved in articulated fossil specimens of birds. The distal–posterior ossification (blue) fuses to the carpometacarpus during the late ontogeny of modern birds. Despite claims it is a neomorphous replacement of the ulnare, its position and development corresponds to dc3, which is found as an independent bone in early dinosaurs, several theropods, and Mesozoic birds (dc3 in Falcarius has also been suggested to be an intermedium).