Imperfections in transparency and mimicry do not increase predation risk for clearwing butterflies with educated predators

Transparency is an intuitive form of concealment and, in certain butterflies, transparent patches on the wings can contribute to several distinct forms of camouflage. However, perhaps paradoxically, the largely transparent wings of many clearwing butterflies (Ithomiini, Nymphalidae) also feature opaque, and often colorful, elements which may reduce crypsis. In many instances, these elements may facilitate aposematic signaling, but little is known of how transparency and aposematism may interact. Here, we used field predation trials to ask two main questions regarding camouflage and signaling in Ithomiini clearwings. In Experiment 1, we focused on camouflage to ask where being transparent may have an advantage over being opaque. We predicted that, as a single opaque pattern can only match a limited range of backgrounds, transparent wings would offer more effective concealment, and experience lower predation risk, over a wider range of backgrounds colors (i.e., green vs. brown substrates) and behaviors (i.e., perched vs. flying) than opaque wings. In Experiment 2, we focused on the effect conspicuous opaque colors may have on clearwing survival. We predicted that although salient signals may increase detectability, those commonly associated with toxic Ithomiini clearwings would not increase predation risk. Both experiments were conducted among educated predators within the natural range of Ithomiini clearwings and we found predation rates to be very low. In Experiment 1, we found some marginal evidence to suggest that opaque, but not transparent, butterflies may suffer increased predation during flight, whereas in Experiment 2, we found equal survival across all model prey types regardless of coloration. Taken together we suggest that any loss of camouflage due to conspicuous coloration may be compensated by aversive signaling, and that educated predators may broadly generalize across a wide range of known and novel clearwing phenotypes.