Birds have peramorphic skulls, too: anatomical network analyses reveal oppositional heterochronies in avian skull evolution

In contrast to the vast majority of reptiles, the skulls of adult crown birds are characterized by a high degree of integration due to bone fusion, e.g., an ontogenetic event generating a net reduction in the number of bones. To understand this process in an evolutionary context, we investigate postnatal ontogenetic changes in the skulls of crown bird and non-avian theropods using anatomical network analysis (AnNA). Due to the greater number of bones and bone contacts, early juvenile crown birds have less integrated skulls, resembling their non-avian theropod ancestors, including Archaeopteryx lithographica and Ichthyornis dispars. Phylogenetic comparisons indicate that skull bone fusion and the resulting modular integration represent a peramorphosis (developmental exaggeration of the ancestral adult trait) that evolved late during avialan evolution, at the origin of crown-birds. Succeeding the general paedomorphic shape trend, the occurrence of an additional peramorphosis reflects the mosaic complexity of the avian skull evolution. Plateau and Foth use anatomical network analysis to study the evolution of avian skull anatomy. They report that the ontogenetic changes in the morphology and modularity of the avian skulls is comparable to evolutionary transformations from non-avian theropods to modern birds. Their work highlights the complexity of avian skull evolution.