Homologies of digits in the avian hand have been debated for 150 years
. Cladistic analysis nests birds with theropod dinosaurs. Theropod han
ds retain only digits I-II-III, so digits of the modern bird hand are
often identified as I-II-III, Study of the developing manus and pes in
amniote embryos, including a variety of avian species, shows stereoty
ped patterns of cartilage condensations. A primary axis of cartilage c
ondensation is visible in all species that runs through the humerus in
to digit IV. Comparison to serially homologous elements of the hindlim
b indicates that the retained digits of the avian hand are II-III-IV.