Hawkmoth flight in the unsteady wakes of flowers

Flying animals maneuver and hover through environments where wind gusts and flower wakes produce unsteady flow. Although both flight maneuvers and aerodynamic mechanisms have been studied independently, little is known about how these interact in an environment where flow is already unsteady. Moths forage from flowers by hovering in the flower's wake. We investigated hawkmoths tracking a 3D-printed robotic flower in a wind tunnel. We visualized the flow in the wake and around the wings and compared tracking performance with previous experiments in a still-air flight chamber. As in still air, moths flying in the flower wake exhibit near-perfect tracking at the low frequencies at which natural flowers move. However, tracking in the flower wake results in a larger overshoot between 2 and 5 Hz. System identification of flower tracking reveals that moths also display reduced-order dynamics in wind compared with still air. Smoke visualization of the flower wake shows that the dominant vortex shedding corresponds to the same frequency band as the increased overshoot. Despite these large effects on tracking dynamics in wind, the leading edge vortex (LEV) remains bound to the wing throughout the wingstroke and does not burst. The LEV also maintains the same qualitative structure seen in steady air. Persistence of a stable LEV during decreased flower tracking demonstrates the interplay between hovering and maneuvering.