Evolution of postcanine complexity in Gomphodontia (Therapsida: Cynodontia)

Gomphodonts form a Triassic radiation of small to medium-bodied (<0.5-2.5 m in length) quadrupedal cynodonts characterized by labiolingually expanded gomphodont postcanines. They were the dominant cynodont group in Middle and Late Triassic ecosystems from the Southern Hemisphere and the first predominantly herbivorous cynodonts to evolve. Gomphodonts were also the first therapsids to develop hypsodonty and a dentition with complex occlusal patterns, and their highly diagnostic upper and lower postcanines show many different morphologies. Here, we explored dental complexity in gomphodont cynodonts through time using geographic information system analysis and orientation patch count applied on 3D crown surfaces of upper and lower gomphodont postcanines belonging to 32 gomphodont taxa. This study reveals that the peak in postcanine complexity was reached early in the evolution of gomphodonts with the emergence in the Early Triassic of omnivorous or insectivorous forms with postcanines made of well-separated cusps and cingular cuspules. Traversodontids evolved simpler postcanines via coalescence of cusps into crests and the development of large occlusal basins, and the Middle Triassic radiation of traversodontids led to a sharp decrease in mean postcanine complexity. Simplification of the postcanines in traversodontids is interpreted as being related to a gradual increase in the consumption of plant material. Interestingly, the trend of insectivory/omnivory high postcanine complexity and herbivory low dental complexity in gomphodonts is opposite to the trend of dental complexity reported in some extant mammals, with omnivorous having low dental complexity and herbivorous higher. Postcanine complexity remained relatively stable throughout the evolution of traversodontids and only slightly diminished in the Late Triassic due to the presence of minute forms with particularly simple postcanines in the Rhaetian. The major phylogenetic diversity and taxonomic richness of Gomphodontia are represented in two periods of time: at the end of the Anisian, an age in which the postcanine complexity is simplifying, and at the early Carnian when the postcanine complexity in traversodontids, the only Gomphodontia represented, is stable.