Predation favours Bicyclus anynana butterflies with fewer forewing eyespots.

There are fewer eyespots on the forewings versus hindwings of nymphalids but the reasons for this uneven distribution remain unclear. One possibility is that, in many butterflies, the hindwing covers part of the ventral forewing at rest and there are fewer forewing sectors to display eyespots (covered eyespots are not continuously visible and are less likely to be under positive selection). A second explanation is that having fewer forewing eyespots confers a selective advantage against predators. We analysed wing overlap at rest in 275 nymphalid species with eyespots and found that many have exposed forewing sectors without eyespots: i.e. wing overlap does not constrain the forewing from having the same number or more eyespots than the hindwing. We performed two predation experiments with mantids to compare the relative fitness of and attack damage patterns on two forms of Bicyclus anynana butterflies, both with seven hindwing eyespots, but with two (in wild-type) or four (in Spotty) ventral forewing eyespots. Spotty experienced more intense predation on the forewings, were shorter-lived and laid fewer eggs. These results suggest that predation pressure limits forewing eyespot number in B. anynana. This may occur if attacks on forewing eyespots have more detrimental consequences for flight than attacks on hindwing eyespots.

Interacting Effects of Eyespot Number and Ultraviolet Reflectivity on Predation Risk in Bicyclus anynana (Lepidoptera: Nymphalidae).

Small marginal eyespots on lepidopteran wings are conspicuous elements that attract a predator's attention to deflect attacks away from the body, but the role of ultraviolet (UV) reflectivity at the center of these patterns and variation in eyespot number in altering the function of eyespots remains unclear. Here, we performed a field-based predation experiment with artificial prey items based on the appearance of squinting bush brown butterflies Bicyclus anynana (Butler, 1879). We tested how two visual properties of the wing pattern affect predation risk: i) the number of eyespots on the ventral forewing surface-two or four; and ii) the UV reflectivity of eyespot centers-normal (where the UV reflectivity of the centers contrasts strongly with that of the darker surrounding ring) or blocked (where this contrast is reduced). In total, 807 prey items were deployed at two sites. We found a significant interaction between the number of ventral forewing eyespots and UV reflectivity in the eyespot centers: in items with fewer eyespots, blocking UV resulted in increased predation risk whereas in items with more eyespots, blocking UV resulted in decreased predation risk. If higher predation of paper models can be equated with higher levels of wing margin/eyespot conspicuity, these results demonstrate that UV reflectivity is an important factor in making eyespots more conspicuous to predators and suggest that the fitness of particular butterfly eyespot number variants may depend on the presence or absence of UV in their centers and on the ability of local predator guilds to detect UV.

Pattern Matters in the Aposematic Colouration of Papilio polytes Butterflies.

Many toxic animals display bright colour patterns to warn predators about their toxicity. This sometimes leads other sympatric palatable organisms to evolve mimetic colour patterns to also evade predation. These mimics, however, are often imperfect, and it is unclear how much their colour patterns can vary away from the model before they become ineffective. In this study, we investigated how predation risk of the palatable Common Mormon butterfly (Papilio polytes) is affected by two alterations of its wing pattern that make it progressively more distinct from its model, the Common Rose (Pachliopta aristolochiae). We deployed butterfly paper models in the field, where all models displayed the same colours but had different patterns. In the first modification from the Wildtype pattern, we exchanged the position of the red and white colour patches but kept the overall pattern constant. In the second modification, we created an eyespot-like shape from the pre-existing pattern elements by moving their positions in the wing, altering the overall wing pattern. Both modifications increased attack risk from predators relative to Wildtype patterns, with the eyespot-like modification having the highest predation risk. Our results show that avian predators can distinguish between all three patterns tested, and that pattern is important in aposematic signals. Predators learn to avoid aposematic colours, not in isolation, but as part of specific patterns.