Linked morphological changes during palate evolution in early tetrapods

We examined the shapes and sizes of dermal bones of the palate of selected Palaeozoic tetrapods in order to identify the ancestral states of palatal bone morphologies in the earliest tetrapods, to learn how the composition of the palate varies within and among early tetrapod radiations, and to recognize evolutionary correlations among the size and shapes of skeletal elements in this important group of animals. We find that whereas the palatal bones themselves and their arrangements are usually conserved, considerable correlated evolutionary change occurs in the shapes and sizes of the bones. Some of the changes in the bones are allometrically linked to overall palate size, which varies more than 100-fold among the taxa in our sample. Often, these allometries were only hinted at in traditional independent contrasts-based regressions of log transformed data, particularly because many allometries are subtle, their slopes may vary among subclades, and the scatter around some trendlines is high. Rather, the allometries showed up in analyses of size-standardized palatal bone dimensions investigated using independent contrasts, bivariate phylomorphospace plots, and mirrored character reconstructions on the phylogenetic tree. We find negative allometry for parasphenoid lengths and widths essentially across the entire tree of Palaeozoic tetrapods, but with different trajectories characterizing the two largest clades, the temnospondyls and the lepospondyls. The lengths of several other elements may show positive allometries, either across the entire tree or in just a subclade. One possible positive allometry exists for the ectopterygoid, which appears to shorten allometrically in temnospondyls that evolve small body and palate size, and, as in Doleserpeton can be lost altogether. Both shortening and loss could be by the same developmental change, paedomorphosis, a form of heterochrony. Paedomorphosis might also account for evolution of relatively large parasphenoids in both lepospondyls and diminutive temnospondyls, but does not seem to explain evolution of ectopterygoid loss in lepospondyls. A regularity observed across nearly all taxa in our study set is an inverse correlation between the lengths of the vomer and pterygoid, bones that lie adjacent to one another along the long palatal axis. Further work is needed to learn whether such correlated evolution might be due to adaptation and/or to developmental bias, and particularly to learn how correlations and allometries themselves evolve.