Comparative architecture of the tessellated boxfish (Ostracioidea) carapace.

Tessellations (surface architectures of arrays of hard tiles) are common in natural and man-made designs. Boxfishes (Ostracioidea) are almost completely encased in a tessellated armor and have evolved a plethora of cross-sectional carapace shapes, yet whether the scutes constructing these exhibit comparable variation is unknown. Using high-resolution microCT and semi-automatic segmentation algorithms, we quantitatively examined thousands of scutes from 13 species of diverse body form. A cluster analysis revealed that certain scute types are associated with specific carapace regions independent of carapace shape. Scute types differentiate between carapace edges and flat regions, as well as between the head region with many carapace openings and the more consistently closed abdominal region, pointing at a constructional commonality or constraint shared by all boxfish species. However, the dimensions of edge scutes varied systematically with carapace shape (e.g., scute aspect ratio tended to increase with decreasing carapace height). This suggests that protection is maintained across body forms by managing scute- and carapace-level mechanisms for increasing bending resistance. Future studies on other taxa are necessary to understand whether these architectural principles are specific evolutionary solutions for building a boxfish carapace or whether they are shared by other biological systems that serve a similar protective function.