Conflicting optical and mechanical needs drive the evolution of wing membrane thickness in Ithomiini clearwing butterflies

Transparency provides benefits to prey animals as it makes them less detectable by predators. Transparent animals are often thin, which raises the question of their fragility. In clearwing Lepidoptera, the wing thickness is the evolutionary result of conflicting optical and mechanical needs. All else being equal, a thinner membrane lets light better go through, can still sustain the reduced scales it often bears, it has a low stiffness, which is advantageous for flight, but it resists less to fatigue and failure, a crucial point. An evolutionary way out of these conflicting needs can be spatial heterogeneity in stiffness, with thicker opaque patches compensating for thinner transparent ones, especially when transparency covers a great wing surface proportion. We tested these predictions in Ithomiine butterflies, a tribe comprising closely-related opaque and transparent unpalatable species. We found that species with partially transparent wings have a thinner membrane in the transparent zone than in the opaque one, which likely helps light getting through and agrees with the lighter weight wings have to support in the transparent zone. Despite this difference between opaque and transparent zones, among transparent species the more transparent ones surprisingly have a thicker membrane in their transparent zone. We find no relationship between membrane thickness and scale density, ruling out a predominant role of membrane thickness as a mechanical support for scales. Finally, species with a higher wing proportion occupied by transparency have thicker membranes on their transparent patch, and a greater ratio in thickness between opaque and transparent zones. These latter two results support the hypothesis that clearwing butterflies with larger transparent patches are potentially more fragile and that this frailty is offset by thicker surrounding opaque patches offering higher mechanical resistance, like tubules framing a kite sail. In clearwing butterflies, wing membrane thickness has likely evolved under optical and mechanical selective pressures and further research should experimentally measure the costs, if any, of thinner transparent membranes.