Galloanseran cranial development highlights exceptions to von Baer's laws

The remarkable morphological disparity of the animal kingdom is underpinned by changes in embryonic development across the tree of life; as such, deciphering evolutionary patterns of developmental divergence depends on investigations of different species across a range of comparable developmental stages. Among the most influential ideas regarding such developmental divergences are von Baer's Laws of Development and Haeckel's Theory of Recapitulation. Here, we assess several predictions following from these ideas at the tissue-level by comparing skull osteogenesis in representatives of the bird clade Galloanserae. We investigated high-resolution µCT scans of embryonic series for four galloanseran species: chickens and quails, representing Galliformes (landfowl), and ducks and geese, representing Anseriformes (waterfowl). To compare skull osteogenesis across our taxon sample, we devised a skull-specific staging system based on ossification sequences to discretise the process into five stages. During skull osteogenesis, we found that the location of the onset of ossification within each element and the direction of ossification progression were the same in all species in our sample, implying a conserved developmental programme for induction and ossification progression across Galloanserae. Moreover, we found that the appearance of synapomorphies diagnostic of broader clades often overlapped with species-specific ones during osteogenesis. Indeed, many diagnostic features of deep clades, such as osteological synapomorphies of the phylogenetically inclusive clade Galloanserae, appear at surprisingly late stages of development. These observations fail to support several predictions of von Baer's Laws of Development and Haeckel's Theory of Recapitulation, instead suggesting what we term a 'braiding' pattern of developmental divergence in which degrees of interspecific morphological similarity wax and wane during development as a result of the interplay between developmental constraints and phyletic variation.