Swallowtail Butterflies Use Multiple Visual Cues to Select Oviposition Sites

SIMPLE SUMMARY: Butterflies must not only identify host plants on which to lay their eggs—which they achieve using chemical cues—but also select suitable leaves on that plant that will support the growth of their larval offspring. Here, we asked whether swallowtail butterflies lay eggs on particular leaves of a Citrus tree and, if so, which cues they use to select the leaves. We first observed that butterflies indeed select just a few leaves on which to lay eggs. These leaf preferences were observed across many individuals, implying that they were not idiosyncratic, and the butterflies descended directly towards the leaves from some distance, suggesting that they were using visual rather than chemical cues. We then investigated which visual cues are used by the butterflies, and found that the number of eggs laid upon a leaf was correlated with its height on the tree, flatness, green reflectance, brightness, and degree of polarization. These five features may be important both for selecting young leaves and those which are situated well for egg-laying. An outstanding question for future study is how visual and chemical cues interact in this context. ABSTRACT: Flower-foraging Japanese yellow swallowtail butterflies, Papilio xuthus, exhibit sophisticated visual abilities. When ovipositing, females presumably attempt to select suitable leaves to support the growth of their larval offspring. We first established that butterflies indeed select particular leaves on which to lay eggs; when presented with a single Citrus tree, butterflies significantly favored two out of 102 leaves for oviposition. These preferences were observed across many individuals, implying that they were not merely idiosyncratic, but rather based on properties of the leaves in question. Because the butterflies descended towards the leaves rather directly from a distance, we hypothesized that they base their selection on visual cues. We measured five morphological properties (height, orientation, flatness, roundness, and size) and four reflective features (green reflectance, brightness, and degree and angle of linear polarization). We found that the number of eggs laid upon a leaf was positively correlated with its height, flatness, green reflectance, and brightness, and negatively correlated with its degree of polarization, indicating that these features may serve as cues for leaf selection. Considering that other studies report ovipositing butterflies’ preference for green color and horizontally polarized light, butterflies likely use multiple visual features to select egg-laying sites on the host plant.