Morphospace saturation in the stem-gnathostomes pteraspidiformes heterostracans: an early radiation of a 'bottom' heavy clade.

Ostracoderms (fossil armoured jawless fishes) shed light on early vertebrate evolution by revealing the step-wise acquisition of jawed vertebrate characters, and were important constituents of Middle Palaeozoic vertebrate faunas. A wide variety of head shield shapes are observed within and between the ostracoderm groups, but the timing of these diversifications and the consistency between different measures of their morphospace are unclear. Here, we present the first disparity (explored morphospace) versus diversity (number of taxa) analysis of Pteraspidiformes heterostracans using continuous and discrete characters. Patterns of taxic diversity and morphological disparity are in accordance: they both show a rise to a peak in the Lochkovian followed by a gradual decline in the Middle-Late Devonian. Patterns are largely consistent for disparity measures using sum of ranges or total variance, and when using continuous or discrete characters. Pteraspidiformes heterostracans can be classified as a "bottom-heavy clade", i.e., a group where a high initial disparity decreasing over time is detected. In fact, the group explored morphospace early in its evolutionary history, with much of the subsequent variation in dermal armour occurring as variation in the proportions of already evolved anatomical features. This Early Devonian radiation is also in agreement with the paleobiogeographic distribution of the group, with a maximum of dispersal and explored morphospace during the Lochkovian and Pragian time bins.