Ichthyornithes

The earliest known ichthyornitheans appear in the fossil record about 95 million years ago, during the Cenomanian age of the Late Cretaceous. Based on fragmentary fossil remains, the two known species present in the Ashville Formation have not been given names, but overall were very similar to Ichthyornis dispar. I. dispar itself had a very long temporal range, and specimens referred to it or very similar species existed relatively unchanged (other than some fluctuations in average adult size) for nearly 10 million years, when species referable to I. dispar disappeared and were replaced by other, though still somewhat similar, species of ichthyornitheans.[1] The fact that ichthyornitheans appear to have existed virtually unchanged in their general size and anatomy for nearly the entire duration of the late Cretaceous period suggests that their evolutionary pace was in relative stasis compared to other, contemporary species of closely related avialans, like the hesperornitheans. Both the ichthyornitheans and hesperornitheans were strongly linked to their ecosystem in the Western Interior Seaway that bisected North America for much of the Cretaceous. It may be that while the flightless, aquatic hesperornitheans were much more sensitive to changes in shoreline extent and sea level during that time, driving their evolution and adaptation into more specialized forms, the flying, tern-like ichthyornitheans were not as dependent on specific coastal or sea-level conditions, and they were able to inhabit basically the same ecological niche for a very long period.[3] The icthyornithean lineage finally became extinct in the Cretaceous–Paleogene extinction event that caused the extinction of many major land and marine animal groups at the end of the Mesozoic era. The last ichthyornithean fossils are found in the Hell Creek Formation very close (within 300,000 years at least) of the K-Pg boundary, dated to 66 million years old.[4]

The study describing Protodontopteryx notes some "striking" similarities between the jaw structure of ichthyornitheans and pelagornithids. The study still classifies pelagornithids as neognaths based on a few post-cranial features and recovers them in a polytomy with Galloanserae and Neoaves, but does note that this link is worth investigating and that the pelagornithid palate is not known.[5]

A study on an Ichthyornis endocast reveals that it had a relatively "primitive" brain compared to modern birds, similar to that of Archaeopteryx and other non-avian theropods. Conversely, it had a palate remarkably convergent with that of modern neognaths.[6]

Ichthyornitheans were close to the ancestry of modern birds, the crown group Aves, but represents an independent lineage of toothed seabird-like animals. It was long believed that several similar Cretaceous species known from fragmentary remains, including Ambiortus, Apatornis, Iaceornis and Guildavis, were members of the Ichthyornithes in addition to the group's namesake, Ichthyornis. However, these seem to have been more closely related to modern birds than to Ichthyornis dispar. In Julia Clarke's 2004 review of Ichthyornis-like fossils, the former order Ichthyornithiformes and the family Ichthyornithidae were considered synonyms of the clade Ichthyornithes, which in that paper was defined according to phylogenetic taxonomy as all descendants of the most recent common ancestor of Ichthyornis dispar and modern birds.[1]