State-dependent Anti-predator Response and Escape Refuge Choice of a Dead-leaf Butterfly (Kallima inachus chinensis)

Studies on animal mimicry have long focused on the morphological similarity between the mimic and the model, and validation of the anti-predator effect, with less attention being paid to the behavioral response following predator detection. Therefore, this study observed the anti-predator behavior of the dead-leaf butterfly (Kallima inachus chinensis) during basking, perching, feeding, and mating with a predatory threat from a highly realistic model great tit and analyzed the microhabitat selection preferences of adults after escape. The results showed, firstly, that the oakleaf butterfly could effectively distinguish between predatory threats and non-threatening disturbances, where 86% of tested individuals responded only to the model great tit stimulus and showed no response to the control stimulus, indicating a precise threat recognition ability. Secondly, its anti-predator behaviors primarily manifested as either a stationary response or an escape response, with the latter being dominant (61.5%). Behavioral state significantly influenced the response rate, and the majority of individuals responded during basking and perching states (response rates of 70.0% and 65.0%, respectively), while the response rate decreased to 45.0% during feeding. All individuals engaged in mating showed no observable response, reflecting a clear “reproduction-priority” strategy. Furthermore, after escaping, adults showed a significant preference for perching on the upper surface of green leaves, rather than near dead leaves, as described in previous studies. This provides new behavioral evidence for understanding its defensive strategy (morphological deception vs. background matching). There were also no significant differences in the aforementioned behavioral responses between sexes. This study is the first to empirically reveal the behavioral plasticity and state-dependence of the oakleaf butterfly under predation risk, demonstrating that even species relying on highly specialized morphological camouflage possess dynamic and adjustable behavioral defense strategies, thereby deepening the understanding of anti-predator adaptation mechanisms in invertebrates with deceptive appearances.