Comparative digital reconstruction of Pica pica and Struthio camelus and their cranial suture ontogenies.

To date, several studies describe post-hatching ontogenetic variation in birds; however, none of these studies document and compare ontogenetic variation of the entire skull in multiple avian species. Therefore, we studied ontogenetic skull variation of two bird species with very different ecologies, Pica pica, and Struthio camelus, using µCT based 3D reconstructions. For each specimen, we performed bone-by-bone segmentation in order to visualize and describe the morphological variation of each bone during ontogeny and estimated the average sutural closure of the skulls to identify different ontogenetic stages. Although bone fusion of P. pica occurs more rapidly than that of S. camelus the general sequence of bone fusion follows a similar trend from posterior to anterior, but a more detailed analysis reveals some interspecific variation in the fusion patterns. Although growth persists over a longer period in S. camelus than in P. pica and adults of the former species are significantly larger, the skull of the most mature S. camelus is still less fused than that of P. pica. Different growth and fusion patterns of the two species indicate that the interspecific ontogenetic variation could be related to heterochronic developments. Nevertheless, this hypothesis needs to be tested in a broader phylogenetic framework in order to detect the evolutionary direction of the potential heterochronic transformations.

Second-order division in sectors as a prepattern for sensory organs in vertebrate development.

We present in vivo observations of chicken embryo development which show that the early chicken embryo presents a principal structure made out of concentric rings and a secondary structure composed of radial sectors. During development, physical forces deform the main rings into axially directed, antero-posterior tubes, while the sectors roll up to form cylinders that are perpendicular to the antero-posterior axis. As a consequence, the basic structure of the chicken embryo is a series of encased antero-posterior tubes (gut, neural tube, body envelope, amnion, chorion) decorated with smaller orifices (ear duct, eye stalk, nasal duct, gills, mouth) forming at right angles to the main body axis. We argue that the second-order divisions reflect the early pattern of cell cleavage, and that the transformation of radial and orthoradial lines into a body with sensory organs is a generic biophysical mechanism more general than the chicken embryo.

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.

Correlated evolution of neck length and leg length in birds.

Despite a diversity of about 10 000 extant species, the sophisticated avian 'body plan' has not much changed once it was achieved around 160 Ma after the origin of powered flight. All birds are bipedal having wings, a rigid trunk, a short and ossified tail, a three-segmented leg and digitigrade feet. The avian neck, however, has always been regarded as a classic example of high variability ranging from short necks in songbirds to extremely long, serpentine necks in herons. Yet, the wide array of small to very large species makes it difficult to evaluate the actual neck length. Here, we investigate the evolution of the vertebral formulae in the neck of birds and the scaling relationships between skeletal dimensions and body size. Cervical count in birds is strongly related to phylogeny, with only some specialists having an exceptional number of vertebrae in the neck. In contrast with mammals, the length of the cervical vertebral column increases as body size increases and, thus, body size does not constrain neck length in birds. Indeed, neck length scales isometrically with total leg length suggesting a correlated evolution between both modules. The strong integration between the cervical and pelvic module in birds is in contrast with the decoupling of the fore- and hindlimb module and may be the result of the loss of a functionally versatile forelimb due to the evolution of powered flight.