Elastic wing deformations mitigate flapping asymmetry during manoeuvres in rose chafers (Protaetia cuprea).

To manoeuvre in air, flying animals produce asymmetric flapping between contralateral wings. Unlike the adjustable vertebrate wings, insect wings lack intrinsic musculature, preventing active control over wing shape during flight. However, the wings elastically deform as a result of aerodynamic and inertial forces generated by the flapping motions. How these elastic deformations vary with flapping kinematics and flight performance in free-flying insects is poorly understood. Using high-speed videography, we measured how contralateral wings elastically deform during free-flight manoeuvring in rose chafer beetles (Protaetia cuprea). We found that asymmetric flapping during aerial turns was associated with contralateral differences in chord-wise wing deformations. The highest instantaneous difference in deformation occurred during stroke reversals, resulting from differences in wing rotation timing. Elastic deformation asymmetry was also evident during mid-strokes, where wing compliance increased the angle of attack of both wings, but reduced the asymmetry in the angle of attack between contralateral wings. A biomechanical model revealed that wing compliance can increase the torques generated by each wing, providing higher potential for manoeuvrability, while concomitantly contributing to flight stability by attenuating steering asymmetry. Such stability may be adaptive for insects such as flower chafers that need to perform delicate low-speed landing manoeuvres among vegetation.

Allometry of wing twist and camber in a flower chafer during free flight: How do wing deformations scale with body size?

Intraspecific variation in adult body mass can be particularly high in some insect species, mandating adjustment of the wing's structural properties to support the weight of the larger body mass in air. Insect wings elastically deform during flapping, dynamically changing the twist and camber of the relatively thin and flat aerofoil. We examined how wing deformations during free flight scale with body mass within a species of rose chafers (Coleoptera: Protaetia cuprea) in which individuals varied more than threefold in body mass (0.38-1.29 g). Beetles taking off voluntarily were filmed using three high-speed cameras and the instantaneous deformation of their wings during the flapping cycle was analysed. Flapping frequency decreased in larger beetles but, otherwise, flapping kinematics remained similar in both small and large beetles. Deflection of the wing chord-wise varied along the span, with average deflections at the proximal trailing edge higher by 0.2 and 0.197 wing lengths compared to the distal trailing edge in the downstroke and the upstroke, respectively. These deflections scaled with wing chord to the power of 1.0, implying a constant twist and camber despite the variations in wing and body size. This suggests that the allometric growth in wing size includes adjustment of the flexural stiffness of the wing structure to preserve wing twist and camber during flapping.

Turning in mid-air allows aphids that flee the plant to avoid reaching the risky ground.

When forced to drop from the plant, flightless arboreal insects can avoid reaching the risky ground by maneuvering their body through the air. When wingless pea aphids (Acyrthosiphon pisum) are threatened by natural enemies, they often drop off their host plant while assuming a stereotypic posture that rotates them in mid-air, aligning them with their feet pointing downwards. This position may increase their chances of re-clinging onto lower plant parts and avoid facing the dangers on the ground, although its effectiveness in realistic field conditions has not been tested. We performed both laboratory and outdoor experiments, in which we dropped aphids upon host plants to quantify clinging success in plants with different characteristics such as height and leaf size. Live aphids had twofold higher clinging rates than dead ones, indicating that clinging success is indeed affected by the active aerial-righting of dropping aphids. The ability to cling was positively dependent on the plants' foliage cover as viewed in vertical direction from above. Therefore, we released aphids in commercial alfalfa (Medicago sativa) fields with varying plant heights and foliage cover and induced them to drop. Most (up to 75%) of the aphids avoided reaching the ground in taller plants (65 cm), and 17% in shorter plants (21 cm), demonstrating the efficiency of the aphids' response in averting risks: both those of an approaching enemy on the plant and the plethora of new risks on the ground. Evidently, even in complex field environment, the aerial-righting mechanism can substantially reduce the possible risks following escape from a predator.

The stimuli evoking the aerial-righting posture of falling pea aphids.

Some wingless insects possess aerial righting reflexes, suggesting that adaptation for controlling body orientation while falling through air could have preceded flight. When threatened by natural enemies, wingless pea aphids (Acyrthosiphon pisum) may drop off their host plant and assume a stereotypic posture that rotates them in midair to land on their feet. The sensory information triggering aphids to assume this posture has so far been unknown. We subjected aphids to a series of tests, isolating the sensory cues experienced during free-fall. Falling aphids assumed the righting posture and landed upright irrespective of whether the experiments were carried out in the light or in complete darkness. Detachment of the tarsi from the substrate triggered the aphids to assume the posture rapidly, but only for a brief period. Rotation (mainly roll and yaw) of the body in air, in the light, caused aphids to assume the posture and remain in it throughout rotation. In contrast, aphids rotated in the dark did not respond. Acceleration associated with falling or airflow over the body per se did not trigger the posture. However, sensing motion relative to air heightened the aphids' responsiveness to rotation in the light. These results suggest that the righting posture of aphids is triggered by a tarsal reflex, but, once the aphid is airborne, vision and a sense of motion relative to air can augment the response. Hence, aerial righting in a wingless insect could have emerged as a basic tarsal response and developed further to include secondary sensory cues typical of falling.