Evolutionary and developmental aspects of avian-specific traits in limb skeletal pattern

The two sets of paired appendages, called limbs, are locomotory organs in tetrapods that are used for various functions (e.g., walking, running, crawling, digging, climbing, diving, swimming, and flying). Unlike such organs as the eye, which contain specialized tissues such as the lens and photoreceptor, the limb does not have any specialized cells or tissues, but consists of common tissues, such as bone, cartilage, muscle, blood vessels, and dermis. However, limb morphology is highly specialized and varies to provide species-specific modes of locomotion. As do the vertebrae and skull, the limb skeleton varies in morphology among species. The diversity of limb skeletal morphology provides examples of material for studies on morphogenesis. Avian forelimbs have evolved into wings for flight. The skeletal pattern in the avian limb has many traits that are unique among extant species of vertebrates; some of such traits are avian-specific, others are shared with more basal members of Theropoda, to which Aves belongs. Since such avian traits generally form during ontogenic development, determining when and how they appear in the developing embryonic limbs or limb buds provides important insights into the mechanisms underlying the generation of vertebrate morphological diversity. Here, we present an overview of several features of the skeletal pattern in the avian limb and discuss the developmental mechanisms responsible for their unique and lineage-specific traits.