Superb autorotator: rapid decelerations in impulsively launched samaras.

Autorotation of botanical samaras, with a consequent reduction in their rate of descent, increases dispersal range in the presence of horizontal winds. Samaras in initial free fall from rest pass through a brief transitional phase prior to reaching their minimum rate of descent and stable autorotation. By contrast, intense wind gusts and elastic recoil of tree branches can produce impulsive samara detachment and accelerate them rapidly through the air. Here, we investigate the autorotation of maple samaras when launched with a high initial impulse. Norway maple seeds catapulted either vertically or horizontally at approximately 9 m s-1 exhibited remarkably high and rapid decelerations (10-15 g) and reached a near-zero translational speed in less than 150 ms. The initial rotational frequency of catapulted seeds was up to four times greater than that ultimately reached during steady-state autorotation. These helicopter seeds thus transiently produce very high lift forces (at Reynolds numbers near approximately 104) that act to enhance aerial transport. These findings are relevant to the modelling of long-distance seed dispersal in unsteady flows, as well as to the design of deceleration mechanisms based on lift generation, rather than drag-based devices such as parachutes.

Ascending flight and decelerating vertical glides in Anna's hummingbirds.

Hummingbirds are observationally well known for their capacity to vertically ascend whilst hovering, but the underlying mechanics and possible energetic limits to ascent rates are unclear. Decelerations during vertical ascent to a fixed target may also be associated with specific visual responses to regulate the body's trajectory. Here, we studied climbing flight and subsequent deceleration in male Anna's hummingbirds (Calypte anna) over an approximately 2 m vertical distance. Birds reached vertical speeds and accelerations up to ∼4 m s-1 and 10 m s-2, respectively, through the use of flapping frequencies as high as 56 Hz and stroke amplitudes slightly greater than 180 deg. Total mass-specific power at maximal ascent speed was up to 92 W kg-1 body mass. Near the end of the ascending trajectory, all individuals decelerated ballistically via cessation of flapping and folding of wings over the body without losing control, a behavior termed here a vertical glide. Visual modulation of the deceleration trajectory during ascent was indicated by a constant value (∼0.75) for the first derivative of the time-to-contact to target. Our results indicate that hummingbirds in rapid vertical ascent expended near-maximal power output during flight, but also tightly controlled their subsequent deceleration during the vertical glide.

Living in a trash can: turbulent convective flows impair Drosophila flight performance.

Turbulent flows associated with thermal convection are common in areas where the ground is heated by solar radiation, fermentation or other processes. However, it is unknown how these flow instabilities affect the locomotion of small insects, like fruit flies, that inhabit deserts and urban landscapes where surface temperatures can reach extreme values. We quantified flight performance of fruit flies (Drosophila melanogaster) traversing a chamber through still air and turbulent Rayleigh-Bénard convection cells produced by a vertical temperature gradient. A total of 34% of individuals were unable to reach the end of the chamber in convection, although peak flow speeds were modest relative to typical outdoor airflow. Individuals that were successful in convection were faster fliers and had larger wing areas than those that failed. All flies displayed higher pitch angles and lower mean flight speeds in convection. Successful individuals took longer to cross the chamber in convection, due to lower flight speeds and greater path sinuosity. All individuals displayed higher flapping frequencies in convection, and successful individuals also reduced stroke amplitude. Our results suggest that thermal convection poses a significant challenge for small fliers, resulting in increased travel times and energetic costs, or in some cases precluding insects from traversing these environments entirely.

Escape jumping by three age-classes of water striders from smooth, wavy and bubbling water surfaces.

Surface roughness is a ubiquitous phenomenon in both oceanic and terrestrial waters. For insects that live at the air-water interface, such as water striders, non-linear and multi-scale perturbations produce dynamic surface deformations which may impair locomotion. We studied escape jumps of adults, juveniles and first-instar larvae of the water strider Aquarius remigis on smooth, wave-dominated and bubble-dominated water surfaces. Effects of substrate on takeoff jumps were substantial, with significant reductions in takeoff angles, peak translational speeds, attained heights and power expenditure on more perturbed water surfaces. Age effects were similarly pronounced, with the first-instar larvae experiencing the greatest degradation in performance; age-by-treatment effects were also significant for many kinematic variables. Although commonplace in nature, perturbed water surfaces thus have significant and age-dependent effects on water strider locomotion, and on behavior more generally of surface-dwelling insects.

On the autorotation of animal wings.

Botanical samaras spin about their centre of mass and create vertical aerodynamic forces which slow their rate of descent. Descending autorotation of animal wings, however, has never been documented. We report here that isolated wings from Anna's hummingbirds, and also from 10 species of insects, can stably autorotate and achieve descent speeds and aerodynamic performance comparable to those of samaras. A hummingbird wing loaded at its base with the equivalent of 50% of the bird's body mass descended only twice as fast as an unloaded wing, and rotated at frequencies similar to those of the wings in flapping flight. We found that even entire dead insects could stably autorotate depending on their wing postures. Feather removal trials showed no effect on descent velocity when the secondary feathers were removed from hummingbird wings. By contrast, partial removal of wing primaries substantially improved performance, except when only the outer primary was present. A scaling law for the aerodynamic performance of autorotating wings is well supported if the wing aspect ratio and the relative position of the spinning axis from the wing base are included. Autorotation is a useful and practical method that can be used to explore the aerodynamics of wing design.

Meniscus ascent by thrips (Thysanoptera).

Meniscus climbing using a fixed body posture has been well documented for various aquatic and neustonic insects, but is not known from small flying insects that inadvertently become trapped on water surfaces. Here, we show that thrips (order Thysanoptera) can ascend a meniscus by arching their non-wetting bodies to translate head-first and upward along a water surface; if initially oriented backwards, they can turn by 180° to ascend head-first, and climb upward on a surrounding boundary. Using variable-concentration sucrose solutions, we show that translational and climbing speeds during meniscus ascent vary inversely with fluid viscosity. Becoming trapped in water is a frequent event for flying insects, and given that most of them are very small, dedicated behaviours to escape water may be commonplace among pterygotes.

Electrostatic Charge on Flying Hummingbirds and Its Potential Role in Pollination.

Electrostatic phenomena are known to enhance both wind- and insect-mediated pollination, but have not yet been described for nectar-feeding vertebrates. Here we demonstrate that wild Anna's Hummingbirds (Calypte anna) can carry positive charges up to 800 pC while in flight (mean ± s.d.: 66 ± 129 pC). Triboelectric charging obtained by rubbing an isolated hummingbird wing against various plant structures generated charges up to 700 pC. A metal hummingbird model charged to 400 pC induced bending of floral stamens in four plants (Nicotiana, Hemerocallis, Penstemon, and Aloe spp.), and also attracted falling Lycopodium spores at distances of < 2 mm. Electrostatic forces may therefore influence pollen transfer onto nectar-feeding birds.

Hovering performance of Anna's hummingbirds (Calypte anna) in ground effect.

Aerodynamic performance and energetic savings for flight in ground effect are theoretically maximized during hovering, but have never been directly measured for flying animals. We evaluated flight kinematics, metabolic rates and induced flow velocities for Anna's hummingbirds hovering at heights (relative to wing length R = 5.5 cm) of 0.7R, 0.9R, 1.1R, 1.7R, 2.2R and 8R above a solid surface. Flight at heights less than or equal to 1.1R resulted in significant reductions in the body angle, tail angle, anatomical stroke plane angle, wake-induced velocity, and mechanical and metabolic power expenditures when compared with flight at the control height of 8R. By contrast, stroke plane angle relative to horizontal, wingbeat amplitude and wingbeat frequency were unexpectedly independent of height from ground. Qualitative smoke visualizations suggest that each wing generates a vortex ring during both down- and upstroke. These rings expand upon reaching the ground and present a complex turbulent interaction below the bird's body. Nonetheless, hovering near surfaces results in substantial energetic benefits for hummingbirds, and by inference for all volant taxa that either feed at flowers or otherwise fly close to plant or other surfaces.

Hawkmoth flight performance in tornado-like whirlwind vortices.

Vertical vortex systems such as tornadoes dramatically affect the flight control and stability of aircraft. However, the control implications of smaller scale vertically oriented vortex systems for small fliers such as animals or micro-air vehicles are unknown. Here we examined the flapping kinematics and body dynamics of hawkmoths performing hovering flights (controls) and maintaining position in three different whirlwind intensities with transverse horizontal velocities of 0.7, 0.9 and 1.2 m s(-1), respectively, generated in a vortex chamber. The average and standard deviation of yaw and pitch were respectively increased and reduced in comparison with hovering flights. Average roll orientation was unchanged in whirlwind flights but was more variable from wingbeat to wingbeat than in hovering. Flapping frequency remained unchanged. Wingbeat amplitude was lower and the average stroke plane angle was higher. Asymmetry was found in the angle of attack between right and left wings during both downstroke and upstroke at medium and high vortex intensities. Thus, hawkmoth flight control in tornado-like vortices is achieved by a suite of asymmetric and symmetric changes to wingbeat amplitude, stroke plane angle and principally angle of attack.

Hawkmoth flight stability in turbulent vortex streets.

Shedding of vortices is a common phenomenon in the atmosphere over a wide range of spatial and temporal scales. However, it is unclear how these vortices of varying scales affect the flight performance of flying animals. In order to examine these interactions, we trained seven hawkmoths (Manduca sexta) (wingspan ~9 cm) to fly and feed in a wind tunnel under steady flow (controls) and in the von Kármán vortex street of vertically oriented cylinders (two different cylinders with diameters of 10 and 5 cm) at speeds of 0.5, 1 and 2 m s(-1). Cylinders were placed at distances of 5, 25 and 100 cm upstream of the moths. Moths exhibited large amplitude yaw oscillations coupled with modest oscillations in roll and pitch, and slight increases in wingbeat frequency when flying in both the near (recirculating) and middle (vortex dominated) wake regions. Wingbeat amplitude did not vary among treatments, except at 1 m s(-1) for the large cylinder. Yaw and roll oscillations were synchronized with the vortex shedding frequencies in moths flying in the wake of the large cylinder at all speeds. In contrast, yaw and pitch were synchronized with the shedding frequency of small vortices at speeds ≤1 m s(-1). 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.

Spiderweb deformation induced by electrostatically charged insects.

Capture success of spider webs has been associated with their microstructure, ornamentation, and wind-induced vibrations. Indirect evidence suggests that statically charged objects can attract silk thread, but web deformations induced by charged insects have not yet been described. Here, we show under laboratory conditions that electrostatically charged honeybees, green bottle flies, fruit flies, aphids, and also water drops falling near webs of cross-spiders (Araneus diadematus) induce rapid thread deformation that enhances the likelihood of physical contact, and thus of prey capture.

Flying in the rain: hovering performance of Anna's hummingbirds under varied precipitation.

Flight in rain represents a greater challenge for smaller animals because the relative effects of water loading and drop impact are greater at reduced scales given the increased ratios of surface area to mass. Nevertheless, it is well known that small volant taxa such as hummingbirds can continue foraging even in extreme precipitation. Here, we evaluated the effect of four rain intensities (i.e. zero, light, moderate and heavy) on the hovering performance of Anna's hummingbirds (Calypte anna) under laboratory conditions. Light-to-moderate rain had only a marginal effect on flight kinematics; wingbeat frequency of individuals in moderate rain was reduced by 7 per cent relative to control conditions. By contrast, birds hovering in heavy rain adopted more horizontal body and tail positions, and also increased wingbeat frequency substantially, while reducing stroke amplitude when compared with control conditions. The ratio between peak forces produced by single drops on a wing and on a solid surface suggests that feathers can absorb associated impact forces by up to approximately 50 per cent. Remarkably, hummingbirds hovered well even under heavy precipitation (i.e. 270 mm h(-1)) with no apparent loss of control, although mechanical power output assuming perfect and zero storage of elastic energy was estimated to be about 9 and 57 per cent higher, respectively, compared with normal hovering.

Aerial shaking performance of wet Anna's hummingbirds.

External wetting poses problems of immediate heat loss and long-term pathogen growth for vertebrates. Beyond these risks, the locomotor ability of smaller animals, and particularly of fliers, may be impaired by water adhering to the body. Here, we report on the remarkable ability of hummingbirds to perform rapid shakes in order to expel water from their plumage even while in flight. Kinematic performance of aerial versus non-aerial shakes (i.e. those performed while perching) was compared. Oscillation frequencies of the head, body and tail were lower in aerial shakes. Tangential speeds and accelerations of the trunk and tail were roughly similar in aerial and non-aerial shakes, but values for head motions while perching were twice as high when compared with aerial shakes [corrected] . Azimuthal angular amplitudes for both aerial and non-aerial shakes reached values greater than 180° for the head, greater than 45° for the body trunk and slightly greater than 90° for the tail and wings. Using a feather on an oscillating disc to mimic shaking motions, we found that bending increased average speeds by up to 36 per cent and accelerations of the feather tip up to fourfold relative to a hypothetical rigid feather. Feather flexibility may help to enhance shedding of water and reduce body oscillations during shaking.