Grain boundaries and coincidence site lattices in the corneal nanonipple structure of the Mourning Cloak butterfly

In this study the highly ordered corneal nanonipple structure observed on the Mourning Cloak butterfly (Nymphalis antiopa) is analyzed with a particular emphasis on the high-angle grain-boundary-like defects that are observed between individual nanonipple crystals. It is shown that these grain boundaries are generated by rows of topological coordination defects, which create very specific misorientations between adjacent crystals. These specific orientations form coincidence site lattices, which (i) have unit cells larger than the unit cell in each individual crystal and (ii) extend from one crystal to the next, effectively creating order over areas larger than the individual crystals. A comparison to similar coincidence site lattices in engineering materials is made and the importance of such arrangements in terms of nipple packing density, corneal lens curvature and potential optical properties is discussed.