The asymmetry of the carpal joint and the evolution of wing folding in maniraptoran theropod dinosaurs

Corwin Sullivan*, David W. E. Hone, Xing Xu, Fucheng Zhang

*Corresponding author for this work

Research output: Contribution to journalArticle (Academic Journal)

27 Citations (Scopus)


In extant birds, the hand is permanently abducted towards the ulna, and the wrist joint can bend extensively in this direction to fold the wing when not in use. Anatomically, this asymmetric mobility of the wrist results from the wedge-like shape of one carpal bone, the radiale, and from the well-developed convexity of the trochlea at the proximal end of the carpometacarpus. Among the theropod precursors of birds, a strongly convex trochlea is characteristic of Coelurosauria, a clade including the highly derived Maniraptora in addition to tyrannosaurs and compsognathids. The shape of the radiale can be quantified using a 'radiale angle' between the proximal and distal articular surfaces. Measurement of the radiale angle and reconstruction of ancestral states using squared-change parsimony shows that the angle was small (15 degrees) in primitive coelurosaurs but considerably larger (25 degrees) in primitive maniraptorans, indicating that the radiale was more wedge-shaped and the carpal joint more asymmetric. The radiale angle progressively increased still further within Maniraptora, with concurrent elongation of the forelimb feathers and the forelimb itself. Carpal asymmetry would have permitted avian-like folding of the forelimb in order to protect the plumage, an early advantage of the flexible, asymmetric wrist inherited by birds.

Original languageEnglish
Pages (from-to)2027-2033
Number of pages7
JournalProceedings of the Royal Society B: Biological Sciences
Issue number1690
Publication statusPublished - 7 Jul 2010


  • Theropoda
  • Maniraptora
  • Aves
  • carpus
  • radiale
  • feathers


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