Distinct causes underlie double-peaked trilobite morphological disparity in cephalic shape

Trilobite cephalic shape disparity varied through geological time and was integral to their ecological niche diversity, and so is widely used for taxonomic assignments. To fully appreciate trilobite cephalic evolution, we must understand how this disparity varies and the factors responsible. We explore trilobite cephalic disparity using a dataset of 983 cephalon outlines of c. 520 species, analysing the associations between cephalic morphometry and taxonomic assignment and geological Period. Elliptical Fourier transformation visualised as a Principal Components Analysis suggests significant differences in morphospace occupation and in disparity measures between the groups. Cephalic shape disparity peaks in the Ordovician and Devonian. The Cambrian-Ordovician expansion of morphospace occupation reflects radiations to new niches, with all trilobite orders established by the late Ordovician. In comparison, the Silurian-Devonian expansion seems solely a result of within-niche diversification. Linear Discriminant Analyses cross-validation, average cephalon shapes, and centroid distances demonstrate that, except for Harpida and the Cambrian and Ordovician Periods, order and geological Period cannot be robustly predicted for an unknown trilobite. Further, k-means clustering analyses suggest the total dataset naturally subdivides into only seven groups that do not correspond with taxonomy, though k-means clusters do decrease in number through the Palaeozoic, aligning with findings of decreasing disparity.