Unlike large animals, insects and other very small animals are so unsusceptible to impact-related injuries that they can use falling for dispersal and predator evasion. Reorienting to land upright can mitigate lost access to resources and predation risk. Such behaviours are critical for the spotted lanternfly (SLF) ( Lycorma delicatula ), an invasive, destructive insect pest spreading rapidly in the USA. High-speed video of SLF nymphs released under different conditions showed that these insects self-right using both active midair righting motions previously reported for other insects and novel post-impact mechanisms that take advantage of their ability to experience near-total energy loss on impact. Unlike during terrestrial self-righting, in which an animal initially at rest on its back uses appendage motions to flip over, SLF nymphs impacted the surface at varying angles and then self-righted during the rebound using coordinated body rotations, foot–substrate adhesion and active leg motions. These previously unreported strategies were found to promote disproportionately upright, secure landings on both hard, flat surfaces and tilted, compliant host plant leaves. Our results highlight the importance of examining biomechanical phenomena in ecologically relevant contexts, and show that, for small animals, the post-impact bounce period can be critical for achieving an upright landing.

Many small animals use aerial righting to mitigate the risks associated with falling, such as predation, starvation, and desiccation. Spotted lanternflies (Lycorma delicatula) (SLFs) are invasive insect pests that often fall from host plants in response to predators or abiotic factors (e.g., wind). We used high-speed video to study whether immature SLFs (nymphs) impact surfaces, and subsequently land upright, more often than expected by chance, and, if so, whether they do so via active or passive mechanisms. SLF nymphs were found to adopt a stereotypical falling posture proposed to promote passive righting, and similar to those assumed by falling insects, spiders, geckos, frogs and skydivers. Live SLF nymphs landed upright in more trials when releasing voluntarily (100%) or when dropped from tweezers (56%) than did dead specimens (33-35%), with differences being highly statistically significant, even when the dead specimens were posed in the falling posture. These results support a role for active aerial righting. The fraction landing upright also did not depend significantly either on orientation during release or at first impact. We found that significantly more live SLFs reoriented to upright after impacting in a non-upright orientation via bouncing in combination with adhering to the substrate with one or more feet. Videos of nymphs landing on leaves confirmed that these insects can use similar tactics to land upright on host plants. These findings indicate the important role post-impact righting plays in determining final orientation, and highlight the importance of studies that include ecologically-relevant substrates and naturalistic conditions.