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.

Into turbulent air: size-dependent effects of von Kármán vortex streets on hummingbird flight kinematics and energetics.

Animal fliers frequently move through a variety of perturbed flows during their daily aerial routines. However, the extent to which these perturbations influence flight control and energetic expenditure is essentially unknown. Here, we evaluate the kinematic and metabolic consequences of flight within variably sized vortex shedding flows using five Anna's hummingbirds feeding from an artificial flower in steady control flow and within vortex wakes produced behind vertical cylinders. Tests were conducted at three horizontal airspeeds (3, 6 and 9 m s(-1)) and using three different wake-generating cylinders (with diameters equal to 38, 77 and 173% of birds' wing length). Only minimal effects on wing and body kinematics were demonstrated for flight behind the smallest cylinder, whereas flight behind the medium-sized cylinder resulted in significant increases in the variances of wingbeat frequency, and variances of body orientation, especially at higher airspeeds. Metabolic rate was, however, unchanged relative to that of unperturbed flight. Hummingbirds flying within the vortex street behind the largest cylinder exhibited highest increases in variances of wingbeat frequency, and of body roll, pitch and yaw amplitudes at all measured airspeeds. Impressively, metabolic rate under this last condition increased by up to 25% compared with control flights. Cylinder wakes sufficiently large to interact with both wings can thus strongly affect stability in flight, eliciting compensatory kinematic changes with a consequent increase in flight metabolic costs. Our findings suggest that vortical flows frequently encountered by aerial taxa in diverse environments may impose substantial energetic costs.

Kinematics of flight and the relationship to the vortex wake of a Pallas' long tongued bat (Glossophaga soricina).

To obtain a full understanding of the aerodynamics of animal flight, the movement of the wings, the kinematics, needs to be connected to the wake left behind the animal. Here the detailed 3D wingbeat kinematics of bats, Glossophaga soricina, flying in a wind tunnel over a range of flight speeds (1-7 m s(-1)) was determined from high-speed video. The results were compared with the wake geometry and quantitative wake measurements obtained simultaneously to the kinematics. The wingbeat kinematics varied gradually with flight speed and reflected the changes observed in the wake of the bats. In particular, several of the kinematic parameters reflected the differences in the function of the upstroke at low and high flight speeds. At lower flight speeds the bats use a pitch-up rotation to produce a backward flick which creates thrust and some weight support. At higher speeds this mechanism disappears and the upstroke generates weight support but no thrust. This is reflected by the changes in e.g. angle of attack, span ratio, camber and downstroke ratio. We also determined how different parameters vary throughout a wingbeat over the flight speeds studied. Both the camber and the angle of attack varied over the wingbeat differently at different speeds, suggesting active control of these parameters to adjust to the changing aerodynamic conditions. This study of the kinematics strongly indicates that the flight of bats is governed by an unsteady high-lift mechanism at low flight speeds and points to differences between birds and bats.

A quantitative comparison of bird and bat wakes.

Qualitative comparison of bird and bat wakes has demonstrated significant differences in the structure of the far wake. Birds have been found to have a unified vortex wake of the two wings, while bats have a more complex wake with gradients in the circulation along the wingspan, and with each wing generating its own vortex structure. Here, we compare quantitative measures of the circulation in the far wake of three bird and one bat species. We find that bats have a significantly stronger normalized circulation of the start vortex than birds. We also find differences in how the circulation develops during the wingbeat as demonstrated by the ratio of the circulation of the dominant start vortex and the total circulation of the same sense. Birds show a more prominent change with changing flight speed and a relatively weaker start vortex at minimum power speed than bats. We also find that bats have a higher normalized wake loading based on the start vortex, indicating higher relative induced drag and therefore less efficient lift generation than birds. Our results thus indicate fundamental differences in the aerodynamics of bird and bat flight that will further our understanding of the evolution of vertebrate flight.

The near and far wake of Pallas' long tongued bat (Glossophaga soricina).

The wake structures of a bat in flight have a number of characteristics not associated with any of the bird species studied to this point. Unique features include discrete vortex rings generating negative lift at the end of the upstroke at medium and high speeds, each wing generating its own vortex loop, and a systematic variation in the circulation of the start and stop vortices along the wingspan, with increasing strength towards the wing tips. Here we analyse in further detail some previously published data from quantitative measurements of the wake behind a small bat species flying at speeds ranging from 1.5 to 7 m s(-1) in a wind tunnel. The data are extended to include both near- and far-wake measurements. The near-/far-wake comparisons show that although the measured peak vorticity of the start and stop vortices decreases with increasing downstream distance from the wing, the total circulation remains approximately constant. As the wake evolves, the diffuse stop vortex shed at the inner wing forms a more concentrated vortex in the far wake. Taken together, the results show that studying the far wake, which has been the standard procedure, nevertheless risks missing details of the wake. Although study of the far wake alone can lead to the misinterpretation of the wake topology, the net, overall circulation of the main wake vortices can be preserved so that approximate momentum balance calculations are not unreasonable within the inevitably large experimental uncertainties.