Influence of modularity and regularity on disparity of atelostomata sea urchins

A modularity approach is used to study disparity rates and evolvability of sea urchins belonging to the Atelostomata superorder. For this purpose, the pentameric sea urchin architecture is partitioned into modular spatial components and the interference between modules is quantified using areas and a measurement of the regularity of the spatial partitions. This information is used to account for the variability through time (disparity) and potential for morphological variation and evolution (evolvability) in holasteroid echinoids. We obtain that regular partitions of the space produce modules with high modular integrity, whereas irregular partitions produce low modular integrity; the former ones are related with high morphological disparity (facilitation hypothesis). Our analysis also suggests that a pentameric body plan with low regularity rates in Atelostomata reflects a stronger modular integration among modules than within modules, which could favors bilaterality against radial symmetry. Our approach constitutes a theoretical platform to define and quantify spatial organization in partitions of the space that can be related to modules in a morphological analysis.