On the homology of crocodylian post-dentary bones and their macroevolution throughout Pseudosuchia

The lower jaw of early tetrapods is composed of several intramembranous ossifications. However, a tendency toward the independent reduction of the number of bones has been observed in the mandible of mammals, lepidosaurs, turtles, crocodiles, and birds. Regarding archosaurs, the coronoid and prearticular bones are interpreted to be lost during the evolution of stem-birds and stem-crocodiles, respectively, but the homology of the post-dentary bones retained in living pseudosuchians remains unclear. Here, we combine paleontological and embryological evidence to explore in detail the homology of the crocodylian post-dentary bones. We study the mandible embryogenesis on a sample of 71 embryos of Caiman and compare this pattern with the mandibular transformations observed across pseudosuchian evolution. In the pre-hatching ontogeny of caimans, at least five intramembranous ossification centers are formed along the margins of the internal mandibular fenestra (perifenestral centers) and, subsequently, merge to form the coronoid (three intramembranous centers), angular (one intramembranous center), and articular (one intramembranous and one chondral center). In the fossil record, an independent prearticular is lost around the base of Mesoeucrocodylia (optimized as reappearing in Thalattosuchia if they are placed within Neosuchia), and the coronoid is apomorphically lost in notosuchians. The integration of embryological and paleontological data indicates that most perifenestral centers are involved in the origin of the prearticular of non-mesoeucrocodylian pseudosuchians. These centers are rearranged during the evolution to contribute to different post-dentary bones in mesoeucrocodylians bolstering the idea that the coronoid and the articular of Crocodylia are not completely homologous to those of other diapsids.