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[Figure][1] The ability of certain animal groups to fly by themselves has always stirred our imaginations. Even in the 15th century Leonardo da Vinci was famously inspired to try to build bird-like ornithopters. However, it was not until the 19th century that the nature of aerodynamic lift

We have studied the nocturnal flight behaviour of the common swift (Apus apusL.), by the use of a tracking radar. Birds were tracked from Lund University in southern Sweden during spring migration, summer roosting flights and autumn migration. Flight speeds were compared with predictions from flight mechanical and optimal migration theories. During spring, flight speeds were predicted to be higher than during both summer and autumn due to time restriction. In such cases, birds fly at a flight speed that maximizes the overall speed of migration. For summer roosting flights, speeds were predicted to be lower than during both spring and autumn since the predicted flight speed is the minimum power speed that involves the lowest energy consumption per unit time. During autumn, we expected flight speeds to be higher than during summer but lower than during spring since the expected flight speed is the maximum range speed, which involves the lowest energy consumption per unit distance. Flight speeds during spring were indeed higher than during both summer and autumn, which indicates time-selected spring migration. Speeds during autumn migration were very similar to those recorded during summer roosting flights. The general result shows that swifts change their flight speed between different flight behaviours to a smaller extent than expected. Furthermore, the difference between flight speeds during migration and roosting among swifts was found to be less pronounced than previously recorded.

Birds and bats have evolved powered flight independently, which makes a comparison of evolutionary 'design' solutions potentially interesting. In this paper we highlight similarities and differences with respect to flight characteristics, including morphology, flight kinematics, aerodynamics, energetics and flight performance. Birds' size range is 0.002-15 kg and bats' size range is 0.002-1.5 kg. The wingbeat kinematics differ between birds and bats, which is mainly due to the different flexing of the wing during the upstroke and constraints by having a wing of feathers and a skin membrane, respectively. Aerodynamically, bats appear to generate a more complex wake than birds. Bats may be more closely adapted for slow maneuvering flight than birds, as required by their aerial hawking foraging habits. The metabolic rate and power required to fly are similar among birds and bats. Both groups share many characteristics associated with flight, such as for example low amounts of DNA in cells, the ability to accumulate fat as fuel for hibernation and migration, and parallel habitat-related wing shape adaptations.

SUMMARYThe 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.

Staying aloft when hovering and flying slowly is demanding. According to quasi–steady-state aerodynamic theory, slow-flying vertebrates should not be able to generate enough lift to remain aloft. Therefore, unsteady aerodynamic mechanisms to enhance lift production have been proposed. Using digital particle image velocimetry, we showed that a small nectar-feeding bat is able to increase lift by as much as 40% using attached leading-edge vortices (LEVs) during slow forward flight, resulting in a maximum lift coefficient of 4.8. The airflow passing over the LEV reattaches behind the LEV smoothly to the wing, despite the exceptionally large local angles of attack and wing camber. Our results show that the use of unsteady aerodynamic mechanisms in flapping flight is not limited to insects but is also used by larger and heavier animals.

Studies of bird migration in the Beringia region of Alaska and eastern Siberia are of special interest for revealing the importance of bird migration between Eurasia and North America, for evaluating orientation principles used by the birds at polar latitudes and for understanding the evolutionary implications of intercontinental migratory connectivity among birds as well as their parasites. We used tracking radar placed onboard the ice-breaker Oden to register bird migratory flights from 30 July to 19 August 2005 and we encountered extensive bird migration in the whole Beringia range from latitude 64° N in Bering Strait up to latitude 75° N far north of Wrangel Island, with eastward flights making up 79% of all track directions. The results from Beringia were used in combination with radar studies from the Arctic Ocean north of Siberia and in the Beaufort Sea to make a reconstruction of a major Siberian–American bird migration system in a wide Arctic sector between longitudes 110° E and 130° W, spanning one-third of the entire circumpolar circle. This system was estimated to involve more than 2 million birds, mainly shorebirds, terns and skuas, flying across the Arctic Ocean at mean altitudes exceeding 1 km (maximum altitudes 3–5 km). Great circle orientation provided a significantly better fit with observed flight directions at 20 different sites and areas than constant geographical compass orientation. The long flights over the sea spanned 40–80 degrees of longitude, corresponding to distances and durations of 1400–2600 km and 26–48 hours, respectively. The birds continued from this eastward migration system over the Arctic Ocean into several different flyway systems at the American continents and the Pacific Ocean. Minimization of distances between tundra breeding sectors and northerly stopover sites, in combination with the Beringia glacial refugium and colonization history, seemed to be important for the evolution of this major polar bird migration system.

Avian migration, which involves billions of birds flying vast distances, is known to influence all aspects of avian life. Here we investigate how birds fit moult into an annual cycle determined by the need to migrate. Large variation exists in moulting patterns in relation to migration: for instance, moult can occur after breeding in the summer or after arrival in the wintering quarters. Here we use an optimal annual routine model to investigate why this variation exists. The modelled bird's decisions depend on the time of year, its energy reserves, breeding status, experience, flight feather quality and location. Our results suggest that the temporal and spatial variations in food are an important influence on a migratory bird's annual cycle. Summer moult occurs when food has a high peak on the breeding site in the summer, but it is less seasonal elsewhere. Winter moult occurs if there is a short period of high food availability in summer and a strong winter peak at different locations (i.e. the food is very seasonal but in opposite phase on these areas). This finding might explain why only long-distance migrants have a winter moult.

The flapping flight of animals generates an aerodynamic footprint as a time-varying vortex wake in which the rate of momentum change represents the aerodynamic force. We showed that the wakes of a small bat species differ from those of birds in some important respects. In our bats, each wing generated its own vortex loop. Also, at moderate and high flight speeds, the circulation on the outer (hand) wing and the arm wing differed in sign during the upstroke, resulting in negative lift on the hand wing and positive lift on the arm wing. Our interpretations of the unsteady aerodynamic performance and function of membranous-winged, flapping flight should change modeling strategies for the study of equivalent natural and engineered flying devices.

The wingbeat kinematics and wake structure of a trained house martin in free, steady flight in a wind tunnel have been studied over a range of flight speeds, and compared and contrasted with similar measurements for a thrush nightingale and a pair of robins. The house martin has a higher aspect ratio (more slender) wing, and is a more obviously agile and aerobatic flyer, catching insects on the wing. The wingbeat is notable for the presence at higher flight speeds of a characteristic pause in the upstroke. The essential characteristics of the wing motions can be reconstructed with a simple two-frequency model derived from Fourier analysis. At slow speeds, the distribution of wake vorticity is more simple than for the other previously measured birds, and the upstroke does not contribute to weight support. The upstroke becomes gradually more significant as the flight speed increases, and although the vortex wake shows a signature of the pause phase, the global circulation measurements are otherwise in good agreement with surprisingly simple aerodynamic models, and with predictions across the different species, implying quite similar aerodynamic performance of the wing sections. The local Reynolds numbers of the wing sections are sufficiently low that the well-known instabilities of attached laminar flows over lifting surfaces, which are known to occur at two to three times this value, may not develop.

Hovering means stationary flight at zero net forward speed, which can be achieved by animals through muscle powered flapping flight. Small bats capable of hovering typically do so with a downstroke in an inclined stroke plane, and with an aerodynamically active outer wing during the upstroke. The magnitude and time history of aerodynamic forces should be reflected by vorticity shed into the wake. We thus expect hovering bats to generate a characteristic wake, but this has until now never been studied. Here we trained nectar-feeding bats,Leptonycteris yerbabuenae, to hover at a feeder and using time-resolved stereoscopic particle image velocimetry in conjunction with high-speed kinematic analysis we show that hovering nectar-feeding bats produce a series of bilateral stacked vortex loops. Vortex visualizations suggest that the downstroke produces the majority of the weight support, but that the upstroke contributes positively to the lift production. However, the relative contributions from downstroke and upstroke could not be determined on the basis of the wake, because wake elements from down- and upstroke mix and interact. We also use a modified actuator disc model to estimate lift force, power and flap efficiency. Based on our quantitative wake-induced velocities, the model accounts for weight support well (108%). Estimates of aerodynamic efficiency suggest hovering flight is less efficient than forward flapping flight, while the overall energy conversion efficiency (mechanical power output/metabolic power) was estimated at 13%.

The wakes of two individual robins were measured in digital particle image velocimetry (DPIV) experiments conducted in the Lund wind tunnel. Wake measurements were compared with each other, and with previous studies in the same facility. There was no significant individual variation in any of the measured quantities. Qualitatively, the wake structure and its gradual variation with flight speed were exactly as previously measured for the thrush nightingale. A procedure that accounts for the disparate sources of circulation spread over the complex wake structure nevertheless can account for the vertical momentum flux required to support the weight, and an example calculation is given for estimating drag from the components of horizontal momentum flux (whose net value is zero). The measured circulations of the largest structures in the wake can be predicted quite well by simple models, and expressions are given to predict these and other measurable quantities in future bird flight experiments.

Flight feather moult in small passerines is realized in several ways. Some species moult once after breeding or once on their wintering grounds; others even moult twice. The adaptive significance of this diversity is still largely unknown. We compared the resistance to mechanical fatigue of flight feathers from the chiffchaffPhylloscopus collybita, a migratory species moulting once on its breeding grounds, with feathers from the willow warblerPhylloscopus trochilus, a migratory species moulting in both its breeding and wintering grounds. We found that flight feathers of willow warblers, which have a shaft with a comparatively large diameter, become fatigued much faster than feathers of chiffchaffs under an artificial cyclic bending regime. We propose that willow warblers may strengthen their flight feathers by increasing the diameter of the shaft, which may lead to a more rapid accumulation of damage in willow warblers than in chiffchaffs.

We analysed five data sets of night directions of migrating arctic waders ill relation to,winds, recorded by tracking radar and optical range finder, in order to find out if these birds compensate for wind drift, or allow themselves to be drifted by winds. Our purpose was to investigate whether arctic waders use adaptive wind drift strategies or not. The data sets were collected in Siberia (two sets) and Canada during post-breeding (autumn) migration, and in Mauritania and South Sweden during pre-breeding (spring) migration. Both significant drift and compensation effects were found in three of the data sets, Canada, Mauritania and South Sweden. Almost no compensation was found in birds departing in easterly directions from the Siberian tundra (complete drift), while no drift effect was found in birds departing in westerly directions (complete compensation). There were indications that at least some populations of waders may use an adaptive drift strategy consisting of drift at high altitude and/or in high wind speed combined with compensation at low altitude and/or in lower wind speeds, but support for this idea was rather weak and not consistent. Our results were instead more in accordance with the adaptive drift theory that predicts initial drift during the migratory journey, followed by compensation during later stages as the birds are approaching their destinations. Such a strategy implies that arctic waders, at least adult birds, have the capacity of true navigation. A comparison with earlier studies of migrating arctic waders from different parts of the world show that all results so far may be interpreted in accordance with this general adaptive drift strategy An element of non-adaptive drift can, however, not be completely ruled out.

OF VIEW Point-of-view contains short papers ( _< 5 printed pages, about 4000 words) on selected topics that are intended for dis ussion and c mment on recent issues. Authors are encouraged t be provocative and argue their own views on controversial issues. Point-of-View is also open for communication of new ideas and re-evaluations of previously published data and hypotheses. Readers are welcome to suggest suitable topics and/or potential authors at JAB@ekol.lu.se. An abstract is not required.

In an influential paper published in Oikos, Pennycuick (1978) presented a list of “fifteen testable predictions about bird flight” derived from principles of flight mechanical theory (Pennycuick 1975, 1989). This paper has served greatly as a condensed agenda for bird flight research since 1978, as it focused straightforwardly on direct relationships between entities that could easily be measured, in some cases providing some technical innovations. By testing such predictions, one also implicitly tests the validity of flight-mechanical theory. Pennycuick’s (1978) paper was directly addressed to field ornithologists interested in bird flight to design experiments to test the theory. Many took his advice and were usually richly rewarded by interesting results and fine publications. The subsequent research on bird flight mechanics and migration has also generated additional predictions about flight (e.g. Hedenstrom and Alerstam 1995). Here I review tests of Pennycuick’s 15 original predictions and evaluate the outcome of these. In many cases, the predictions have not been tested explicitly, but in such cases I have compiled published information that shed some light on the relationship in question. I will also update the list of predictions, and so this chapter could serve as a guide to those interested in finding further relevant research problems.

SUMMARYIn view of the complexity of the wing-beat kinematics and geometry, an important class of theoretical models for analysis and prediction of bird flight performance entirely, or almost entirely, ignores the action of the wing itself and considers only the resulting motions in the air behind the bird. These motions can also be complicated, but some success has previously been recorded in detecting and measuring relatively simple wake structures that can sometimes account for required quantities used to estimate aerodynamic power consumption. To date, all bird wakes, measured or presumed,seem to fall into one of two classes: the closed-loop, discrete vortex model at low flight speeds, and the constant-circulation, continuous vortex model at moderate to high speeds. Here, novel and accurate quantitative measurements of velocity fields in vertical planes aligned with the freestream are used to investigate the wake structure of a thrush nightingale over its entire range of natural flight speeds. At most flight speeds, the wake cannot be categorised as one of the two standard types, but has an intermediate structure, with approximations to the closed-loop and constant-circulation models at the extremes. A careful accounting for all vortical structures revealed with the high-resolution technique permits resolution of the previously unexplained wake momentum paradox. All the measured wake structures have sufficient momentum to provide weight support over the wingbeat. A simple model is formulated and explained that mimics the correct, measured balance of forces in the downstroke- and upstroke-generated wake over the entire range of flight speeds. Pending further work on different bird species, this might form the basis for a generalisable flight model.

Projected body frontal area is used when estimating the parasite drag of bird flight. We investigated the relationship between projected frontal area and body mass among passerine birds, and compared it with an equation based on waterfowl and raptors, which is used as default procedure in a widespread software package for flight performance calculations. The allometric equation based on waterfowl/raptors underestimates the frontal area compared to the passerine equation presented here. Consequently, revising the actual frontal areas of small birds will concomitantly change the values of the parasite drag coefficient. We suggest that the new equation S-b = 0.0129m(B)(0.61) (m(2)) where m(B) is body mass (kg) should be used when a value of frontal area is needed for passerines. (Less)

Birds appear to use their tails during flight, but until recently the aerodynamic role that tails fulfil was largely unknown. In recent years delta-wing theory, devised to predict the aerodynamics of high-performance aircraft, has been applied to the tails of birds and has been successful in providing a model for the aerodynamics of a bird's tail. This theory now provides the conventional explanation for how birds' tails work. A delta-wing theory (slender-wing theory) has been used, as part of a variable-geometry model to predict how tail and wing shape should vary during flight at different airspeeds. We tested these predictions using barn swallows flying in a wind tunnel. We show that the predictions are not quantitatively well supported. This suggests that a new theory or a modified version of delta-wing theory is needed to adequately explain the way in which morphology varies during flight.

SUMMARYTwo barn swallows (Hirundo rustica) flying in the Lund wind tunnel were filmed using synchronised high-speed cameras to obtain posterior, ventral and lateral views of the birds in horizontal flapping flight. We investigated wingbeat kinematics, body tilt angle, tail spread and angle of attack at speeds of 4–14ms−1. Wingbeat frequency showed a clear U-shaped relationship with air speed with minima at 8.9ms−1(bird 1) and 8.7ms−1 (bird 2). A method previously used by other authors of estimating the body drag coefficient (CD,par) by obtaining agreement between the calculated minimum power (Vmin) and the observed minimum wingbeat frequency does not appear to be valid in this species, possibly due to upstroke pauses that occur at intermediate and high speeds, causing the apparent wingbeat frequency to be lower. These upstroke pauses represent flap-gliding, which is possibly a way of adjusting the force generated to the requirements at medium and high speeds, similar to the flap-bound mode of flight in other species. Body tilt angle, tail spread and angle of attack all increase with decreasing speed, thereby providing an additional lift surface and suggesting an important aerodynamic function for the tail at low speeds in forward flight. Results from this study indicate the high plasticity in the wingbeat kinematics and use of the tail that birds have available to them in order to adjust the lift and power output required for flight.

Flight directions of birds migrating at high geographic and magnetic latitudes can be used to test bird orientation by celestial or geomagnetic compass systems under polar conditions. Migration patterns of arctic shorebirds, revealed by tracking radar studies during an icebreaker expedition along the Northwest Passage in 1999, support predicted sun compass trajectories but cannot be reconciled with orientation along either geographic or magnetic loxodromes (rhumb lines). Sun compass routes are similar to orthodromes (great circle routes) at high latitudes, showing changing geographic courses as the birds traverse longitudes and their internal clock gets out of phase with local time. These routes bring the shorebirds from high arctic Canada to the east coast of North America, from which they make transoceanic flights to South America. The observations are also consistent with a migration link between Siberia and the Beaufort Sea region by way of sun compass routes across the Arctic Ocean.

Among birds, falcons are high-performance flyers, in many cases adapted for aerial hunting and hence suitable targets for investigating limits to flight performance. Using an optical range finder, we measured flight tracks of Eleonora’s falcon (Falco eleonorae), a species breeding in the Mediterranean region and specialised for hunting autumn passage bird migrants, when commuting between their nesting colony and offshore hunting areas (straight transportation flight) and when searching for prey (transecting and searching flight). Airspeed during searching flight was significantly slower than during straight transportation and transecting flight, but there was no significant difference in airspeed between the latter two flight modes. Straight transportation flight was significantly faster than predicted minimum power speed. Also, during straight transportation flight, the falcons responded to head- and tailwinds by increasing their airspeed when flying into the wind. However, they did not show any significant airspeed adjustments with respect to the angle between the track and the heading, as would be expected in birds trying to maintain a constant track direction. Mean sustainable climb rate (during ≥:240 s) was 1.4±0.31 m s−1 (mean ± S.D., N=13), which is rather a high rate for a bird the size of an Eleonora’s falcon. The climb rate was used to calculate maximum load-carrying capacity and maximum sustained horizontal flapping flight speed. The mean wingbeat frequency during powered climbing flight was 4.68 Hz, which was used to estimate the mass-specific muscle work. When falcons were leaving the colony for offshore hunting, they gained altitude by slope-soaring when there was an onshore wind. We formulated a simple criterion for the required gliding-flight rate of climb during an initial slope-soaring episode when minimizing the energy cost of reaching a certain altitude far out over the sea (which is where the prey is to be found). This climb rate was 0.36 m s−1, and our observations indicated that the falcons experienced climb rates above this value when soaring in slope-lift.

I report here for the first time measured flight speeds of Ross's and Sabine's gulls, obtained by optical range finder on the northeast Taymyr peninsula, Siberia. For two flocks of Ross's gulls, airspeeds were 12.2 m/s and 14.6 m/s. Mean airspeed for the Sabine's gulls was 13.9 m/s (SD=3.4 m/s. N=6). A comparison of these measured airspeeds with characteristic speeds predicted from aerodynamic theory showed that the Sabine's gulls were flying significantly faster than predicted minimum power speed, but their speed was not significantly different from maximum range speed. Sabine's gulls adjusted the airspeed in relation to head- and tailwinds, which is in agreement with aerodynamic theory. Key words: flight speed, optical range finder, wind response, Rhodostethia rosea , Larus sabini

The ability to fly in animals is one of the masterpieces of evolution. The ecological advantages include a rapid escape mechanism, and rapid and relatively (energetically) cheap transport. Even though humans have been interested in bird flight for centuries, the interest remains at a high level and modern research is steadily revealing new findings. The flying animal is the ideal model system for the organismal biologist, because it offers challenges in aerodynamics, material science, physiology, neurobiology and ecology, and their interactions.

We measured the air speeds of 31 bird species, for which we had body mass and wing measurements, migrating along the east coast of Sweden in autumn, using a Vectronix Vector 21 ornithodolite and a Gill WindSonic anemometer. We expected each species’ average air speed to exceed its calculated minimum-power speed (Vmp), and to fall below its maximum-range speed (Vmr), but found some exceptions to both limits. To resolve these discrepancies, we first reduced the assumed induced power factor for all species from 1.2 to 0.9, attributing this to splayed and up-turned primary feathers, and then assigned body drag coefficients for different species down to 0.060 for small waders, and up to 0.12 for the mute swan, in the Reynolds number range 25 000–250 000. These results will be used to amend the default values in existing software that estimates fuel consumption in migration, energy heights on arrival and other aspects of flight performance, using classical aeronautical theory. The body drag coefficients are central to range calculations. Although they cannot be measured on dead bird bodies, they could be checked against wind tunnel measurements on living birds, using existing methods.

SUMMARYIn this paper we describe the flight characteristics of a swift (Apus apus) in cruising flight at three different flight speeds (8.0, 8.4 and 9.2 m s–1) in a low turbulence wind tunnel. The wingbeat kinematics were recorded by high-speed filming and the wake of the bird was visualized by digital particle image velocimetry (DPIV). Certain flight characteristics of the swift differ from those of previously studied species. As the flight speed increases, the angular velocity of the wingbeat remains constant, and so as the wingbeat amplitude increases, the frequency decreases accordingly, as though the flight muscles were contracting at a fixed rate. The wings are also comparatively inflexible and are flexed or retracted rather little during the upstroke. The upstroke is always aerodynamically active and this is reflected in the wake, where shedding of spanwise vorticity occurs throughout the wingbeat. Although the wake superficially resembles those of other birds in cruising flight, with a pair of trailing wingtip vortices connected by spanwise vortices, the continuous shedding of first positive vorticity during the downstroke and then negative vorticity during the upstroke suggests a wing whose circulation is gradually increasing and then decreasing during the wingbeat cycle. The wake (and implied wing aerodynamics)are not well described by discrete vortex loop models, but a new wake-based model, where incremental spanwise and streamwise variations of the wake impulse are integrated over the wingbeat, shows good agreement of the vertical momentum flux with the required weight support. The total drag was also estimated from the wake alone, and the calculated lift:drag ratio of approximately 13 for flapping flight is the highest measured yet for birds.

[Figure][1] When swift fledglings depart the nest, there's no going back. They may not touch down again for another two years, and many swifts only perch to mate and raise young. According to Per Henningsson from Lund University, Sweden, the bird's aerial life style is reflected in their

A novel application of DPIV methods is presented for measuring velocity and vorticity distributions in vertical cross sections through the wake of a freely flying bird (thrush nightingale) in a wind tunnel. A dual-camera system is used, and successive cross-correlation operations remove lens/camera distortions, and then the undisturbed background flow, so that the final operation simply examines the disturbance effect of the bird alone. The concentration and tuning of processing methods to the disturbance quantities allows full exploitation of the correlation calculation and estimation algorithms. Since the ultimate objective is to deduce forces and power requirements on the bird itself from the wake structure, the analytical procedure is followed through an example on a fixed airfoil, before sample results from extensive bird flight tests are described. The wake structure of the thrush nightingale in slow (5-m/s) flight is qualitatively quite similar to those previously described in the literature, but certain quantitative details are different in important respects.

The Eleonora's Falcon Falco eleonorae breeds on islands and islets in the Mediterranean region and feeds its young on migratory birds caught in the air. The breeding season is scheduled to coincide with the peak of bird migration. Between 12 and 20 September 1997 we measured flight tracks of falcons within a range of 4 km from their breeding cliffs on Isola di San Pietro 6 km off the south-west coast of Sardinia, by using an optical range finder. The male falcons left the colony in directions between W and NW to hunt far off-shore, probably intersecting migrants departing from mainland Europe in southern France. Flight tracks of falcons leaving for off-shore hunting and falcons searching for prey closer to the colony differed significantly, as revealed by a straightness index. Falcons leaving the colony for the far off-shore hunting area climbed to altitudes over 1000 m. The highest altitude recorded was 1649 m. The altitudes reported from radar studies on nocturnal migrants in the Mediterranean match the flight altitudes observed in the departing Eleonora's Falcons. Calculations based on the times taken for falcons to return with prey to the nest indicate a maximum distance to the hunting area of about 24 km. The falcons compensated partially for wind drift caused by cross winds when departing for off-shore hunting. Complete compensation might not be possible if the wave pattern of the sea is used for orientation, due to movements of the reference system relative the ground. The flight time of Eleonora's Falcons did not seem to be severely restricted by a metabolic ceiling, although such a ceiling does not allow extensive day and night hunting.

Despite evidence that wind conditions are an important factor in determining stopover decisions, models of time-minimizing bird migration have up to now emphasized the optimal response of the migrants to variations in fuel acquisition rates. We present a simple model of a time-minimizing migrant faced with two potential wind conditions on each day, which occur with a fixed probability. Wind assistance is modelled as a multiplicative factor in the flight range equation. We identify conditions under which birds leave the stopover site even with no tailwinds and conditions where the birds leave only with tailwinds in cases with global and local variation of the fuel deposition rate. The optimal policy depends on the probability and amount of wind assistance. In all cases there is an initial period at a stopover site when the bird should stay and build up its initially small fuel reserves irrespective of wind. After this initial time, there is a period when the optimal departure decision is to leave when tailwinds occur but stay and continue fuel deposition in other winds. If the probability of tailwinds is low the bird should at some later time change its policy to leave even in unfavourable winds. However, if a certain threshold value of the probability of favourable winds is reached the birds should never leave without wind assistance. These patterns lead to a complex relationship between departure load and fuel deposition rate. We compare our predictions with a null-model where the birds simply leave as soon as favourable winds occur. We further show that the inclusion of wind assistance cannot explain the discrepancy between observed and predicted values of departure loads under local variation in fuelling rates.

Conventional models of bird flight combine metabolic, mechanical, and aerodynamic components to calculate the rate of fuel consumption and power required for flight, from which we may derive the optimal speeds flying animals should use in different situations. These models contain the implicit assumption that the metabolic and mechanical components of power output are constant and show no systematic variation with speed. This assumption underlies the optimum flight speeds, optimum climb protocol, maximum endurance and maximum flight range predicted by these models. Here we consider alternatives to the assumption that power is independent of speed and show that if the aerodynamic power output from a constant rate of fuel consumption varies with flight speed, then the optima derived from current models of animal flight need to be revised. In some cases (e.g. optimum flight speeds) the current models give answers that are only quantitatively wrong, but in some cases (e.g. optimal cruising altitude) current models give answers that are qualitatively wrong.

Lidar is a powerful technique normally associated with atmospheric monitoring. However, lidar techniques, also of the laser-induced fluorescence and laser-induced breakdown spectroscopy varieties, provide many new possibilities in unconventional fields including cultural heritage and ecological applications.

Optimal migration theory predicts that birds minimizing the overall time of migration should adjust stopover duration with respect to the rate of fuel accumulation. Recent theoretical developments also take into account the wind situation and predict that there is a time window (a set of days) during which birds should depart when assisted by winds but will not do so if there are head winds. There is also a final day when birds will depart irrespective of wind conditions. Hence, the wind model of optimal migration theory predicts that birds should be sensitive to winds and that there should be a correlation between departures and winds blowing towards the intended migration direction. We tested this assumption by tracking the departures of radio-tagged passerines during autumn migration in southern Sweden. Our birds were moderately to very fat when released and therefore energetically ready for departure. There was a significant correlation between direction of departure and wind direction. We also found that during days when birds departed there was a significantly larger tail wind component than during days when birds were present but did not depart. Our results show that passerines do take the current wind situation into account when departing on migratory flights. We also briefly discuss possible clues that birds use when estimating wind direction and strength. The inclusion of wind is an important amendment to optimal migration theory of birds and should be explored further.

Song flight in birds serves as a signal involved in mate choice and territoriality. In the Skylark Alauda arvensis the song flight is a noticeable feature during the breeding season. Single song flights sometimes lasted for more than 30 min, but the average duration was about 4 min. In this study I dissect the flight performance during the three phases of Skylark song flight: climbing, level and descending flight. During climb, the Skylark exhibits a power expenditure well below the estimated maximum capacity. The average climb speed was 0.98 m s-1. During level flight, which makes up the longest part of the whole song flight, the larks seem to minimize the energy cost by flying at the minimum power speed Vmp if the wind allows. This was evident from the occurrence of circling and turning flight in calm conditions and more stationary wind hovering at wind speeds above Vmp. Finally, descent is most often by parachuting at a mean sinking speed of 1.5 m s-'; diving with folded wings gave a mean sinking speed of 8.4 m s-'; flapping descent resulted in a mean sinking speed of 1.6 m s-1.

Sustained climb rates and airspeeds in flapping flight were measured by radar tracking fifteen species of migrating birds ranging in body mass between 10 g and 10 kg. There was an inverse correlation between body size and climb rate: the lowest mean climb rate, 0.32 m s−1, was observed in the mute swan and the highest mean value, 1.63 m s−1, in the dunlin. Some dunlin flocks achieved sustained climb rates exceeding 2ms−1, up to 2.14ms−1. Assuming that the migrants expend maximum sustained power during their climbs, the climbing power can be used as a conservative estimate of the power margin. Estimates of climbing power for the species tracked by radar were used, in conjunction with aerodynamic theory, to calculate the amount of extra load the migrants should be able to carry if their power margin was used for load transportation rather than for climbing. Calculated ratios of total body mass with maximum load to lean body mass ranged between 1.28 and 2.75, showing an overall negative correlation with body size. There was a broad agreement with maximum fuel loads observed among free-living birds, indicating that the upper limits of fuel-carrying capacities and flight ranges in migrating birds are determined by power margin constraints in sustained flapping flight. Markedly reduced climb rates have been recorded for shorebirds departing with very large fuel reserves from W. Africa and Iceland, supporting the calculated trade-off between climb rate and fuel load. Total power consumption was estimated as the sum of calculated aerodynamic power for forward flight and climbing power. The ratio of total power to the expected minimum aerodynamic power was 1.1–1.3 in the three largest species, increasing to 2–4 in the smallest species. Medium-and small-sized species seem to allocate power in excess of the minimum aerodynamic power not only for climbing but also for maintaining a forward speed faster than the minimum power speed. Given provisional estimates of flight muscle masses and wingbeat frequencies, the mass-specific sustained muscle work for the different climbing bird species was calculated to be in the range 16–41 joules per kilogram muscle mass, showing a significant positive correlation with body mass.

This paper introduces some theoretical concepts based on optimality theory that may be applied to bat migration. Most predictions are based on flight mechanics and potential flight range. Optimal behavior will depend on the relevant immediate currency, such as safety, time, or energy. Predictions concern optimal flight speeds, stopover duration, fuel load at departure, and the adaptive use of winds. I also consider a criterion for fuel management when bats are migrating between summer and hibernation sites. The overall migration speed is predicted to be 46 km/day for a small (10-g) bat, and banding recoveries in Nathusius's bat (Pipistrellus nathusii) showed a mean speed of 47 km/day. The timing of migration in bats that minimizes the time spent in migration is considered a result of the trade-off between seasonally increased length of the night (decreased in spring) and reduction of aerial insect abundance. Most predictions remains to be tested; the theoretical framework herein offers a point of departure when designing new field or laboratory studies of bat migration.

Abstract. A prediction of optimal climb rate up to the cruising altitude for birds setting out on a migration flight is developed. The argument is based on aerodynamic flight theory and minimization of energy expended during the whole flight. Predicted climb rates for knots, Calidris canutus, and turnstones, Arenaria interpres, are compared with observational data of these species when departing on spring migration from north-west Iceland. The data agree reasonably well with the predicted values, but the birds failed to show an expected negative relationship between airspeed and climb rate. It is argued, on the basis of measurements by weather balloons, that observations are likely to have been affected by turbulent air as a result of orographic wind effects at the hilly study site. Shorebirds seem to be able to locate and take advantage of lift while climbing. The test of the predictions should be regarded as provisional and future studies are required to refute or give support to the model.

Abstract A flying animal generates a trail of wake vortices that contain information about the time history and magnitude of aerodynamic forces developed on the wings and body. Methods for visualising and recording wake vortices have been developed, allowing quantitative measurements by digital particle image velocimetry (DPIV). Results from DPIV experiments in a wind tunnel are presented for four passerine species of differing size and morphology. The normalised vorticity and its integrated quantity, circulation (Γ) both decline gradually with increasing flight speed. The measured circulations are successfully explained by a simple aerodynamic model where a normalised circulation, Γ/Uc, represents half the time-averaged lift coefficient, which is >2 at 4 m s−1 for a thrush nightingale.