SUMMARY. Oscillations in body orientation were also substantially smaller in the small cylinder treatment when compared with the large cylinder, regardless of temporal or non-dimensional spatial scale. Moths flying in steady conditions reached a higher air speed than those flying into cylinder wakes. In general, flight effects produced by the cylinder wakes were qualitatively similar among the recirculating and vortex-dominated wake regions; the magnitude of those effects, however, declined gradually with downstream distance.Supplementary material available online at http://jeb.biologists.org/lookup/suppl

It is generally accepted among biology and engineering communities that insects are unstable at hover. However, existing approaches that rely on direct averaging do not fully capture the dynamical features and stability characteristics of insect flight. Here, we reveal a passive stabilization mechanism that insects exploit through their natural wing oscillations: vibrational stabilization. This stabilization technique cannot be captured using the averaging approach commonly used in literature. In contrast, it is elucidated using a special type of calculus: the chronological calculus. Our result is supported through experiments on a real hawkmoth subjected to pitch disturbance from hovering. This finding could be particularly useful to biologists because the vibrational stabilization mechanism may also be exploited by many other creatures. Moreover, our results may inspire more optimal designs for bioinspired flying robots by relaxing the feedback control requirements of flight.

We used videography to investigate direct lateral maneuvers, i.e. ‘sideslips’, of the hawkmoth Manduca sexta. M. sexta sideslip by rolling their entire body and wings to reorient their net force vector. During sideslip they increase net aerodynamic force by flapping with greater amplitude, (in both wing elevation and sweep), allowing them to continue to support body weight while rolled. To execute the roll maneuver we observed in sideslips, they use an asymmetric wing stroke; increasing the pitch of the roll-contralateral wing pair, while decreasing that of the roll-ipsilateral pair. They also increase the wing sweep amplitude of, and decrease the elevation amplitude of, the contralateral wing pair relative to the ipsilateral pair. The roll maneuver unfolds in a stairstep manner, with orientation changing more during downstroke than upstroke. This is due to smaller upstroke wing pitch angle asymmetries as well as increased upstroke flapping counter-torque from left-right differences in global reference frame wing velocity about the moth's roll axis. Rolls are also opposed by stabilizing aerodynamic moments from lateral motion, such that rightward roll velocity will be opposed by rightward motion. Computational modeling using blade-element approaches confirm the plausibility of a causal linkage between the previously mentioned wing kinematics and roll/sideslip. Model results also predict high degrees of axial and lateral damping. On the time scale of whole and half wing strokes, left-right wing pair asymmetries directly relate to the first, but not second, derivative of roll. Collectively, these results strongly support a roll-based sideslip with a high degree of roll damping in M. sexta.

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