False holes as camouflage

Long noted by naturalists, leaf mimicry provides some of the most impressive examples of camouflage through masquerade. Many species of leaf-mimicking Lepidoptera also sport wing markings that closely resemble irregularly shaped holes caused by decay or insect damage. Despite proposals that such markings can either enhance resemblance to damaged leaves or act to disrupt surface appearance through false depth cues, to our knowledge, no attempt has been made to establish exactly how these markings function, or even whether they confer a survival benefit to prey. Here, in two field experiments using artificial butterfly-like targets, we show that false hole markings provide significant survival benefits against avian predation. Furthermore, in a computer-based visual search experiment, we demonstrate that detection of such targets by humans is impeded in a similar fashion. Equally contrasting light marks do not have the same effect; indeed, they lead to increased detection. We conclude that the mechanism is the disruption of the otherwise homogeneous wing surface (surface disruptive camouflage) and that, by resembling the holes sometimes found in real leaves, the disruptive benefits are not offset by conspicuousness costs.