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The 225 million-year-old reptile Sharovipteryx mirabilis was the world's first delta-winged glider; this remarkable animal had a flight surface composed entirely of a hind-limb membrane. We use standard delta-wing aerodynamics to reconstruct the flight of S. mirabilis demonstrating that wing shape could have been controlled simply by protraction of the femora at the knees, and by variation in incidence of a small forelimb canard. Our method has allowed us to address the question of how identifying realistic glide performance can be used to set limits on aerodynamic design in this small animal. Our novel interpretation of the bizarre flight mode of S. mirabilis is the first based directly on interpretation of the fossil itself and the first grounded in aerodynamics.

By analyzing a homogenous dataset we show, in contradiction to a previous study, that the scaling of body frontal area (S(b)) with body mass (m(b)) does not differ between passerine and nonpasserine birds. It is likely that comparison of data collected from live passerines with data collected from frozen nonpasserines had led to the incorrect conclusion that the scaling of S(b) varied between the taxa. We suggest that body dimensions collected from frozen specimens, or specimens stored in alcohol, are not applicable to live birds, and that both the current equations presented in the literature for predicting S(b) from m(b) may lead to inaccurate estimates. Using data from preserved specimens, we found that S(b) scales isometrically with m(b) (S(b) proportional, variant m(b) (0.66)), and therefore we found no evidence for larger birds being more streamlined than smaller birds. S(b) scales with negative allometry against wingspan (b), however, and b scales with positive allometry against m(b), so larger birds have smaller S(b) relative to b. In addition, it appears that dorsoventral flattening of the body is a general characteristic of bird's bodies but that it is more pronounced in larger birds, suggesting perhaps a function in terms of increased lift during forward flight. It appears that bird's bodies obey the surface-to-area geometric scaling law, but bird body shape may vary in relation to aerodynamic function. We suggest that a large-scale study, simultaneously measuring S(b) and m(b) in live passerines and nonpasserines, is required to improve the predictive power of S(b) upon m(b) scaling equations, which play a key role in the estimation of mechanical power consumption in flight in birds. Furthermore, the relations between bird body shape and axial skeleton dimensions, with reference to aerodynamic adaptation, warrant further investigation.

It is technically demanding to measure the energetic cost of animal flight. Each of the previously available techniques has some disadvantage as well advantages. We compared measurements of the energetic cost of flight in a wind tunnel by four European starlings Sturnus vulgaris made using three independent techniques: heat transfer modelling, doubly labelled water (DLW) and mask respirometry. We based our heat transfer model on thermal images of the surface temperature of the birds and air flow past the body and wings calculated from wing beat kinematics. Metabolic power was not sensitive to uncertainty in the value of efficiency when estimated from heat transfer modelling. A change in the assumed value of whole animal efficiency from 0.19 to 0.07 (the range of estimates in previous studies) only altered metabolic power predicted from heat transfer modelling by 13%. The same change in the assumed value of efficiency would cause a 2.7-fold change in metabolic power if it were predicted from mechanical power. Metabolic power did not differ significantly between measurements made using the three techniques when we assumed an efficiency in the range 0.11-0.19, although the DLW results appeared to form a U-shaped power-speed curve while the heat transfer model and respirometry results increased linearly with speed. This is the first time that techniques for determining metabolic power have been compared using data from the same birds flying under the same conditions. Our data provide reassurance that all the techniques produce similar results and suggest that heat transfer modelling may be a useful method for estimating metabolic rate.

We trained two starlings (Sturnus vulgaris) to fly in a wind tunnel whilst wearing respirometry masks. We measured the metabolic power (P(met)) from the rates of oxygen consumption and carbon dioxide production and calculated the mechanical power (P(mech)) from two aerodynamic models using wingbeat kinematics measured by high-speed cinematography. P(met) increased from 10.4 to 14.9 W as flight speed was increased from 6.3 to 14.4 m s(-1) and was compatible with the U-shaped power/speed curve predicted by the aerodynamic models. Flight muscle efficiency varied between 0.13 and 0.23 depending upon the bird, the flight speed and the aerodynamic model used to calculate P(mech). P(met) during flight is often estimated by extrapolation from the mechanical power predicted by aerodynamic models by dividing P(mech) by a flight muscle efficiency of 0.23 and adding the costs of basal metabolism, circulation and respiration. This method would underestimate measured P(met) by 15-25 % in our birds. The mean discrepancy between measured and predicted P(met) could be reduced to 0.1+/-1.5 % if flight muscle efficiency was altered to a value of 0.18. A flight muscle efficiency of 0.18 rather than 0.23 should be used to calculate the flight costs of birds in the size range of starlings (approximately 0.1 kg) if P(met) is calculated from P(mech) derived from aerodynamic models.

Variation with tail spread of the lift generated by a bird tail was measured on mounted, frozen European starlings (Sturnus vulgaris) in a wind tunnel at a typical air speed and body and tail angle of attack in order to test predictions of existing aerodynamic theories modelling tail lift. Measured lift at all but the lowest tail spread angles was significantly lower than the predictions of slender wing, leading edge vortex and lifting line models of lift production. Instead, the tail lift coefficient based on tail area was independent of tail spread, tail aspect ratio and maximum tail span. Theoretical models do not predict bird tail lift reliably and, when applied to tail morphology, may underestimate the aerodynamic optimum tail feather length. Flow visualization experiments reveal that an isolated tail generates leading edge vortices as expected for a low-aspect ratio delta wing, but that in the intact bird body-tail interactions are critical in determining tail aerodynamics: lifting vortices shed from the body interact with the tail and degrade tail lift compared with that of an isolated tail.

The power required for flight in any flying animal is a function of flight speed. The power curve that describes this function has become an icon of studies of flight mechanics and physiology because it encapsulates the accessible animal's flight performance. The mechanical or aerodynamic power curve, describing the increase in kinetic energy of the air due to the passage of the bird, is necessarily U-shaped, for aerodynamic reasons, and can be estimated adequately by lifting-line theory. Predictions from this and related models agree well with measured mechanical work in flight and with results from flow visualization experiments. The total or metabolic power curve also includes energy released by the animal as heat, and is more variable in shape. These curves may be J-shaped for smaller birds and bats, but are difficult to predict theoretically owing to uncertainty about internal physiological processes and the efficiency of the flight muscles. The limitations of some existing models aiming to predict metabolic power curves are considered. The metabolic power curve can be measured for birds or bats flying in wind tunnels at controlled speeds. Simultaneous determination in European starlings Sturnus vulgaris of oxygen uptake, total metabolic rate (using labelled isotopes), aerodynamic power output and heat released (using digital video thermography) enable power curves to be determined with confidence; flight muscle efficiency is surprisingly low (averaging 15-18 %) and increases moderately with flight speed, so that the metabolic power curve is shallower than predicted by models. Accurate knowledge of the power curve is essential since extensive predictions of flight behaviour have been based upon it. The hypothesis that the power curve may not in fact exist, in the sense that the cost of flight may not be perceived by a bird as a continuous smooth function of air speed, is advanced but has not yet formally been tested. This hypothesis is considered together with evidence from variation in flight behaviour, wingbeat kinematics and flight gait with speed. Possible constraints on flight behaviour can be modelled by the power curves: these include the effect of a maximum power output and a constraint on maximum speed determined by downstroke wingbeat geometry and the relationship between thrust and lift.

The vortex wakes of flying and swimming animals provide evidence of the history of aero- and hydrodynamic force generation during the locomotor cycle. Vortex-induced momentum flux in the wake is the reaction of forces the animal imposes on its environment, which must be in equilibrium with inertial and external forces. In flying birds and bats, the flapping wings generate lift both to provide thrust and to support the weight. Distinct wingbeat and wake movement patterns can be identified as gaits. In flow visualization experiments, only two wake patterns have been identified: a vortex ring gait with inactive upstroke, and a continuous vortex gait with active upstroke. These gaits may be modelled theoretically by free vortex and lifting line theory to predict mechanical energy consumption, aerodynamic forces and muscle activity. Longer-winged birds undergo a distinct gait change with speed, but shorter-winged species use the vortex ring gait at all speeds. In swimming fish, the situation is more complex: the wake vortices form a reversed von Kármán vortex street, but little is known about the mechanism of generation of the wake, or about how it varies with speed and acceleration or with body form and swimming mode. An unresolved complicating factor is the interaction between the drag wake of the flapping fish body and the thrusting wake from the tail.

Flight energy is an important factor in the lives of birds. Many strategies and adaptations serve to minimize energy cost and to allow a range of performance consistent with a bird's ecological needs. Theoretical methods can produce good estimates of flight energy that suggest why flight adaptations occur; but remarkably little is known of the physiological adaptations required by flight, or of how these change, a I believe they must, in relation to ecology and flight behavior. More data on the metabolic power consumption of birds in natural flight would be valuable, but it is more important to determine the changes in internal metabolic processes associated with different levels of flight activity. Muscle efficiency in flight, in particular, may have substantial implications for our understanding of the energetic performance of birds. This is but one of a variety of unknown quantities, and only when the mechanisms that determine these are more deeply investigated can flight adaptations be completely understood.