Development and evolution of regionalization within the avian axial column

The origin of birds from their terrestrial antecedents was accompanied by a wholesale transformation of their skeleton as they transitioned from a terrestrial to aerial realm. Part of this dramatic transformation is the reduction of separate vertebral elements into regional fusions to limit axial flexibility. This is partially mirrored within the development of the axial column, with regions of the axial column experiencing increasing morphological modularity and the loss of skeletal elements through vertebral fusions. Using a detailed description of the morphological development of the axial column in the model domestic chicken, Gallus gallus domesticus, we present a map of axial ossification based on discrete characters. Delays in ossification are found to occur in conjunction with the formation of fusions. Our study shows that the pattern and sequence of fusion and ossification during development may reflect the presence of independent modules as subsets within the typical regions of the avian axial column. Interestingly, few of these fusion modules correspond to the initial axial Hox expression patterns, suggesting another patterning mechanism is driving axial fusion regionalization. Additionally, two regions of fusion are discovered in the synsacrum. The anterior region of seven fused synsacrals may correspond to the non-ornithuran pygostylian synsacrum of the same number of vertebrae.