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Body-mounted accelerometers provide a new prospect for estimating power use in flying birds, as the signal varies with the two major kinematic determinants of aerodynamic power: wingbeat frequency and amplitude. Yet wingbeat frequency is sometimes used as a proxy for power output in isolation. There is, therefore, a need to understand which kinematic parameter birds vary and whether this is predicted by flight mode (e.g. accelerating, ascending/descending flight), speed or morphology. We investigate this using high-frequency acceleration data from (i) 14 species flying in the wild, (ii) two species flying in controlled conditions in a wind tunnel and (iii) a review of experimental and field studies. While wingbeat frequency and amplitude were positively correlated, R 2 values were generally low, supporting the idea that parameters can vary independently. Indeed, birds were more likely to modulate wingbeat amplitude for more energy-demanding flight modes, including climbing and take-off. Nonetheless, the striking variability, even within species and flight types, highlights the complexity of describing the kinematic relationships, which appear sensitive to both the biological and physical context. Notwithstanding this, acceleration metrics that incorporate both kinematic parameters should be more robust proxies for power than wingbeat frequency alone.

Space-based tracking technology using low-cost miniature tags is now delivering data on fine-scale animal movement at near-global scale. Linked with remotely sensed environmental data, this offers a biological lens on habitat integrity and connectivity for conservation and human health; a global network of animal sentinels of environmental change. Publisher PDF

The flight speeds that animals should adopt to minimise energy expenditure in different scenarios can be predicted by the curve of power against speed. However, relatively few studies have examined how animals respond to the landscape below them, which could affect speed through modifications in climb rate and perceived predation risk. We equipped homing pigeons(Columba livia)with high frequency loggers to examine how flight speed varies in relation to topography and land cover, predicting that these parameters may have a substantial impact on flight speed and hence cost. Pigeons showed mixed evidence for an energy saving strategy, as they minimised climb rates by starting their ascent ahead of hills, but selected rapid speeds in their ascents. Birds did not modify their speed substantially in relation to land cover, but higher speeds were observed during descending flight, highlighting the importance to consider the rate of change in altitude before estimating power use from speed. Finally, we document an unexpected variability in speed and altitude over fine scales; a source of substantial inefficiency. We suggest this may a form of protean behaviour adopted to reduce predation risk when flocking is not an option, and that such a strategy could be widespread.

Spectacular long-distance migration has evolved repeatedly in animals enabling exploration of resources separated in time and space. In birds, these patterns are largely driven by seasonality, cost of migration, and asymmetries in competition leading most often to leapfrog migration, where northern breeding populations winter furthest to the south. Here, we show that the highly aerial common swiftApus apus, spending the nonbreeding period on the wing, instead exhibits a rarely found chain migration pattern, where the most southern breeding populations in Europe migrate to wintering areas furthest to the south in Africa, whereas the northern populations winter to the north. The swifts concentrated in three major areas in sub-Saharan Africa during the nonbreeding period, with substantial overlap of nearby breeding populations. We found that the southern breeding swifts were larger, raised more young, and arrived to the wintering areas with higher seasonal variation in greenness (Normalized Difference Vegetation Index) earlier than the northern breeding swifts. This unusual chain migration pattern in common swifts is largely driven by differential annual timing and we suggest it evolves by prior occupancy and dominance by size in the breeding quarters and by prior occupancy combined with diffuse competition in the winter.

We develop an optimization model with two decision variables to explore optimal migration mechanisms to facilitate optimal breeding timing in migratory songbirds. In the model, fitness is a function of date-dependent mortality, speed-dependent predation risk, and phenological match at arrival. The model determines the optimal combination of departure date for spring migration and migration speed, which can be mediated either by the power requirement for flight (P) or foraging effort at stopover sites (k). Our model predicts that earlier departure for spring migration should be the primary mechanism underlying earlier breeding timing, with a lesser role for faster migration via lower P or higher k. In contrast, longer migration to breeding areas selects for both earlier departure and faster migration. Empirical data on sex-specific migration traits largely conform to model predictions, since males generally migrate earlier than females but not faster than females. In contrast, empirical data on age-specific migration traits show some disagreement with model predictions, thus implicating additional tradeoffs. In partial agreement with the model, a comparative analysis of 25 songbird species showed that populations with longer migrations migrate more quickly, but do not initiate migration earlier. Our model proves to be a useful framework for interpreting migration strategies in animals making costly seasonal migrations. (Less)

Every year, billions of seasonal migrants connect continents by transporting nutrients, energy, and pathogens between distant communities and ecosystems. For animals that power their movements by endogenous energy stores, the daily energy intake rates strongly influence the speed of migration. If access to food resources varies cyclically over the season, migrants sensitive to changes in daily energy intake rates may adjust timing of migration accordingly. As an effect, individuals adjusting to a common temporal cycle are expected to approach synchrony in foraging and movement. A large-scale periodic pattern, such as the dark–light cycle of the moon, could thus synchronize migrations across animal populations. However, such cyclic effects on the temporal regulation of migration has not been considered. Here, we show the temporal influence of the lunar cycle on the movement activity and migration tactics in a visual hunting nocturnal insectivore and long-distance migrant, the European nightjar, Caprimulgus europeaus. We found that the daily foraging activity more than doubled during moonlit nights, likely driven by an increase in light-dependent fuelling opportunities. This resulted in a clear cyclicity also in the intensity of migratory movements, with occasionally up to 100% of the birds migrating simultaneously following periods of full moon. We conclude that cyclic influences on migrants can act as an important regulator of the progression of individuals and synchronize pulses of migratory populations, with possible downstream effects on associated communities and ecosystems., If the fuelling rate of birds varies periodically, it may act as a synchroniser of migrant populations over large spatial scales. Tracking of the nightjar, a long-distance nocturnal migrant bird, reveals that it adjusts its movements in relation to the lunar cycle.

For studies of how flying animals control their flight, seabirds are of particular interest to track with a biologger because they forage offshore where the visual environment can be simply modeled by a flat world textured by waves. This study suggests that optic flow can explain gull’s altitude control over seas. In particular, a new flight model that includes both energy and optical invariants (called theventral optic flow regulation) explain the dynamics of gulls’ altitude control during offshore takeoff and cruising flight. A linear statistical model applied to 352 flights from 16 individual lesser black backed gulls (Larus fuscus) gave a strong correlation between wind assistance and gulls’ altitude. Thereafter, an optic flow-based flight model was applied to 18 offshore takeoff flights from 1. individual gulls. By introducing an upper limit in climb rate in a non-linear first order parametric model on the gull’s elevation dynamics, coupled with an optic-flow set-point, the predicted altitude gives an optimized fit factor value of 63% on average (min value: 30%, max value: 83%) with respect to GPS data. We conclude that the optic-flow regulation principle (here running close to 25°/s) allows gulls to adjust their altitude over sea without having to directly measure their current altitude.

How aerodynamic power required for animal flight varies with flight speed determines optimal speeds during foraging and migratory flight. Despite its relevance, aerodynamic power provides an elusive quantity to measure directly in animal flight. Here, we determine the aerodynamic power from wake velocity fields, measured using tomographical particle image velocimetry, of pied flycatchers flying freely in a wind tunnel. We find a shallow U-shaped power curve, which is flatter than expected by theory. Based on how the birds vary body angle with speed, we speculate that the shallow curve results from increased body drag coefficient and body frontal area at lower flight speeds. Including modulation of body drag in the model results in a more reasonable fit with data than the traditional model. From the wake structure, we also find a single starting vortex generated from the two wings during the downstroke across flight speeds (1–9 m s−1). This is accomplished by the arm wings interacting at the beginning of the downstroke, generating a unified starting vortex above the body of the bird. We interpret this as a mechanism resulting in a rather uniform downwash and low induced power, which can help explain the higher aerodynamic performance in birds compared with bats.

Slotted wing tips of birds are commonly considered an adaptation to improve soaring performance, despite their presence in species that neither soar nor glide. We used particle image velocimetry to measure the airflow around the slotted wing tip of a jackdaw ( Corvus monedula ) as well as in its wake during unrestrained flight in a wind tunnel. The separated primary feathers produce individual wakes, confirming a multi-slotted function, in both gliding and flapping flight. The resulting multi-cored wingtip vortex represents a spreading of vorticity, which has previously been suggested as indicative of increased aerodynamic efficiency. Considering benefits of the slotted wing tips that are specific to flapping flight combined with the wide phylogenetic occurrence of this configuration, we propose the hypothesis that slotted wings evolved initially to improve performance in powered flight.

Bats navigate the dark using echolocation. Echolocation is enhanced by external ears, but external ears increase the projected frontal area and reduce the streamlining of the animal. External ears are thus expected to compromise flight efficiency, but research suggests that very large ears may mitigate the cost by producing aerodynamic lift. Here we compare quantitative aerodynamic measures of flight efficiency of two bat species, one large-eared (Plecotus auritus) and one small-eared (Glossophaga soricina), flying freely in a wind tunnel. We find that the body drag of both species is higher than previously assumed and that the large-eared species has a higher body drag coefficient, but also produces relatively more ear/body lift than the small-eared species, in line with prior studies on model bats. The measured aerodynamic power ofP. aurituswas higher than predicted from the aerodynamic model, while the small-eared species aligned with predictions. The relatively higher power of the large-eared species results in lower optimal flight speeds and our findings support the notion of a trade-off between the acoustic benefits of large external ears and aerodynamic performance. The result of this trade-off would be the eco-morphological correlation in bat flight, with large-eared bats generally adopting slow-flight feeding strategies.

Summary The morphology and kinematics of a flying animal determines the resulting aerodynamic lift through the regulation of the speed of the air moving across the wing, the wing area and the lift coefficient. We studied the detailed three-dimensional wingbeat kinematics of the bat, Leptonycteris yerbabuenae, flying in a wind tunnel over a range of flight speeds (0–7 m/s), to determine how factors affecting the lift production vary across flight speed and within wingbeats. We found that the wing area, the angle of attack and the camber, which are determinants of the lift production, decreased with increasing speed. The camber is controlled by multiple mechanisms along the span, including the deflection of the leg relative to the body, the bending of the fifth digit, the deflection of the leading edge flap and the upward bending of the wing tip. All these measures vary throughout the wing beat suggesting active or aeroelastic control. The downstroke Strouhal number, Std, is kept relatively constant, suggesting that favorable flow characteristics are maintained during the downstroke, across the range of speeds studied. The Std is kept constant through changes in the stroke plane, from a strongly inclined stroke plane at low speeds to a more vertical stroke plane at high speeds. The mean angular velocity of the wing correlates with the aerodynamic performance and shows a minimum at the speed of maximum lift to drag ratio, suggesting a simple way to determine the optimal speed from kinematics alone. Taken together our results show the high degree of adjustments that the bats employ to fine tune the aerodynamics of the wings and the correlation between kinematics and aerodynamic performance.

Flying insects typically possess two pairs of wings. In beetles, the front pair has evolved into short, hardened structures, the elytra, which protect the second pair of wings and the abdomen. This allows beetles to exploit habitats that would otherwise cause damage to the wings and body. Many beetles fly with the elytra extended, suggesting that they influence aerodynamic performance, but little is known about their role in flight. Using quantitative measurements of the beetle's wake, we show that the presence of the elytra increases vertical force production by approximately 40 per cent, indicating that they contribute to weight support. The wing-elytra combination creates a complex wake compared with previously studied animal wakes. At mid-downstroke, multiple vortices are visible behind each wing. These include a wingtip and an elytron vortex with the same sense of rotation, a body vortex and an additional vortex of the opposite sense of rotation. This latter vortex reflects a negative interaction between the wing and the elytron, resulting in a single wing span efficiency of approximately 0.77 at mid downstroke. This is lower than that found in birds and bats, suggesting that the extra weight support of the elytra comes at the price of reduced efficiency.

Most hovering animals, such as insects and hummingbirds, enhance lift by producing leading edge vortices (LEVs) and by using both the downstroke and upstroke for lift production. By contrast, most hovering passerine birds primarily use the downstroke to generate lift. To compensate for the nearly inactive upstroke, weight support during the downstroke needs to be relatively higher in passerines when compared with, e.g. hummingbirds. Here we show, by capturing the airflow around the wing of a freely flying pied flycatcher, that passerines may use LEVs during the downstroke to increase lift. The LEV contributes up to 49 per cent to weight support, which is three times higher than in hummingbirds, suggesting that avian hoverers compensate for the nearly inactive upstroke by generating stronger LEVs. Contrary to other animals, the LEV strength in the flycatcher is lowest near the wing tip, instead of highest. This is correlated with a spanwise reduction of the wing's angle-of-attack, partly owing to upward bending of primary feathers. We suggest that this helps to delay bursting and shedding of the particularly strong LEV in passerines.

Plumage constitutes a significant component of the somatic investment of birds. A detailed investigation of feathers and moult can help to uncover trade-offs involved in somatic investment decisions, the sources of some of the costs birds have to pay and the potential fitness consequences. We used micro-computed tomography imaging to study the second moment of area, a structural parameter that is one determinant of bending stiffness and the cortex volume of flight feather shafts of two sister taxa, the willow warbler Phylloscopus trochilus, a migratory species with two annual moults, and the chiffchaff Phylloscopus collybita, a migrant with one annual post-nuptial moult. Juvenile and adult willow warbler and chiffchaff feathers, all grown on the breeding grounds, are structurally very similar to each other. Willow warbler feathers grown during moult on the wintering grounds, however, have a significantly higher second moment of area and a significantly larger cortex volume than all the other feather types. We discuss the possibility that the seasonal variability of willow warbler feathers may be an adaptive structural reflection of a moult-migration strategy that has allowed this species to occupy large breeding and wintering ranges.

Migration causes temporal and energetic constraints during plumage development, which can compromise feather structure and function. In turn, given the importance of a good quality of flight feathers in migratory movements, selection may have favoured the synthesis of feathers with better mechanical properties than expected from a feather production constrained by migration necessities. However, no study has assessed whether migratory behaviour affects the relationship between the mechanical properties of feathers and their structural characteristics. We analysed bending stiffness (a feather mechanical property which is relevant to birds’ flight), rachis width and mass (two main determinants of variation in bending stiffness) of wing and tail feathers in migratory and sedentary blackcaps Sylvia atricapilla. Migratory blackcaps produced feathers with a narrower rachis in both wing and tail, but their feathers were not significantly lighter; in addition, bending stiffness was higher in migratory blackcaps than in sedentary blackcaps. Such unexpected result for bending stiffness remained when we statistically controlled for individual variation in rachis width and feather mass, which suggests the existence of specific mechanisms that help migratory blackcaps to improve the mechanical behaviour of their feathers under migration constraints.

Previous studies on wake flow visualization of live animals using DPIV have typically used low repetition rate lasers and 2D imaging. Repetition rates of around 10 Hz allow ~1 image per wingbeat in small birds and bats, and even fewer in insects. To accumulate data representing an entire wingbeat therefore requires the stitching-together of images captured from different wingbeats, and at different locations along the wing span for 3D-construction of wake topologies. A 200 Hz stereo DPIV system has recently been installed in the Lund University wind tunnel facility and the high-frame rate can be used to calculate all three velocity components in a cube, whose third dimension is constructed using the Taylor hypothesis. We studied two bat species differing in body size, Glossophaga soricina and Leptonycteris curasoa. Both species shed a tip vortex during the downstroke that was present well into the upstroke, and a vortex of opposite sign to the tip vortex was shed from the wing root. At the transition between upstroke/downstroke, a vortex loop was shed from each wing, inducing an upwash. Vorticity iso-surfaces confirmed the overall wake topology derived in a previous study. The measured dimensionless circulation, Γ/Uc, which is proportional to a wing section lift coefficient, suggests that unsteady phenomena play a role in the aerodynamics of both species.

Gliding birds continually change the shape and size of their wings(1-6), presumably to exploit the profound effect of wing morphology on aerodynamic performance(7-9). That birds should adjust wing sweep to suit glide speed has been predicted qualitatively by analytical glide models(2,10), which extrapolated the wing's performance envelope from aerodynamic theory. Here we describe the aerodynamic and structural performance of actual swift wings, as measured in a wind tunnel, and on this basis build a semiempirical glide model. By measuring inside and outside swifts' behavioural envelope, we show that choosing the most suitable sweep can halve sink speed or triple turning rate. Extended wings are superior for slow glides and turns; swept wings are superior for fast glides and turns. This superiority is due to better aerodynamic performance - with the exception of fast turns. Swept wings are less effective at generating lift while turning at high speeds, but can bear the extreme loads. Finally, our glide model predicts that cost-effective gliding occurs at speeds of 8 - 10 m s(-1), whereas agility-related figures of merit peak at 15 - 25 m s(-1). In fact, swifts spend the night ('roost') in flight at 8 - 10 m s(-1) ( ref. 11), thus our model can explain this choice for a resting behaviour(11,12). Morphing not only adjusts birds' wing performance to the task at hand, but could also control the flight of future aircraft(7).

New imaging research in the blowfly reveals the dynamic movement of the steering muscles and the complex hinge mechanism in flies that enable astonishing flight maneuverability., Insights into how exactly a fly powers and controls flight have been hindered by the need to unpick the dynamic complexity of the muscles involved. The wingbeats of insects are driven by two antagonistic groups of power muscles and the force is funneled to the wing via a very complex hinge mechanism. The hinge consists of several hardened and articulated cuticle elements called sclerites. This articulation is controlled by a great number of small steering muscles, whose function has been studied by means of kinematics and muscle activity. The details and partly novel function of some of these steering muscles and their tendons have now been revealed in research published in this issue of PLOS Biology. The new study from Graham Taylor and colleagues applies time-resolved X-ray microtomography to obtain a three-dimensional view of the blowfly wingbeat. Asymmetric power output is achieved by differential wingbeat amplitude on the left and right wing, which is mediated by muscular control of the hinge elements to mechanically block the wing stroke and by absorption of work by steering muscles on one of the sides. This new approach permits visualization of the motion of the thorax, wing muscles, and the hinge mechanism. This very promising line of work will help to reveal the complete picture of the flight motor of a fly. It also holds great potential for novel bio-inspired designs of fly-like micro air vehicles.

In this chapter the physics behind animal movement, the adaptations found in locomotion to generate forces, and the mechanisms reducing the cost of transport are discussed. The ability to minimize costs and maximize movement speed is part of the biomechanical physics of animal locomotion. For any type of active movement (e.g. walking, running, swimming, or flying), the animal is required to produce forces to overcome resistance and in many cases also gravity. For terrestrial locomotion, this is achieved by generating ground reaction forces, while in air and water by generating fluid dynamic forces. Due to scaling laws in physics, the speed that can be achieved and the cost of transport are correlated with the size of the animal. Moving across scales thus has consequences for our expectations regarding animal movement, including the occurrence of seasonal migrations, which may be limited by the speed and cost of locomotion.

Slow and hovering animal flight creates high demands on the lift production of animal wings. Steady state aerodynamics is unable to explain the forces required and the most commonly used mechanism to enhance the lift production is a leading edge vortex (LEV). Although LEVs increase the lift, they come at the cost of high drag. Here we determine the flow above the wing of lesser long-nosed bats at slow and cruising speed using particle image velocimetry (PIV). We find that a prominent LEV is present during the downstroke at slow speed, but not at cruising speed. Comparison with previously published LEV data from a robotic flapper inspired by lesser long-nosed bats suggests that bats should be able to generate LEVs at cruising speeds, but that they avoid doing so, probably to increase flight efficiency. In addition, at slow flight speeds we find LEVs of opposite spin at the inner and outer wing during the upstroke, potentially providing a control challenge to the animal. We also note that the LEV stays attached to the wing throughout the downstoke and does not show the complex structures found in insects. This suggests that bats are able to control the development of the LEV and potential control mechanisms are discussed.

The present interest in micro air vehicles has given the research on bat flight a new impulse. With the use of high speed cameras and improved PIV techniques, the kinematics and aerodynamics of bats have been studied in great detail. A robotic flapper makes it possible to do measurements by systematically changing only one parameter at a time and investigate the parameter space outside the natural flight envelope of bats without risking animal safety. For this study, a robotic flapper (RoBat), inspired by Leptonycteris yerbabuenae was developed and tested over the speed range 1-7 m/s, with variable maximum angles of attacks (AoA(max) = 55 degrees and 15 degrees, respectively) and constant AoA(max). = 55 degrees. These measurements show the presence of a leading edge vortex (LEV) for low speeds and a fully attached flow for high speeds at low AoA(max), which is in line with natural bat flight. A LEV occurs for AoA(max) = 55 degrees throughout the complete flight speed range, and throughout which the LEV circulation coefficient remains rather constant. This implies that bats and micro air vehicles could use LEVs for high load maneuvers also at relatively high flight speeds. However, at high flight speeds the LEV bursts, which causes increased drag, most likely due to a decrease in Strouhal number. (C) 2011 Academie des sciences. Published by Elsevier Masson SAS. All rights reserved. (Less)

Recent shifts in phenology in response to climate change are well established but often poorly understood. Many animals integrate climate change across a spatially and temporally dispersed annual life cycle, and effects are modulated by ecological interactions, evolutionary change and endogenous control mechanisms. Here we assess and discuss key statements emerging from the rapidly developing study of changing spring phenology in migratory birds. These well-studied organisms have been instrumental for understanding climate-change effects, but research is developing rapidly and there is a need to attack the big issues rather than risking affirmative science. Although we agree poorly on the support for most claims, agreement regarding the knowledge basis enables consensus regarding broad patterns and likely causes. Empirical data needed for disentangling mechanisms are still scarce, and consequences at a population level and on community composition remain unclear. With increasing knowledge, the overall support ('consensus view') for a claim increased and between-researcher variability in support ('expert opinions') decreased, indicating the importance of assessing and communicating the knowledge basis. A proper integration across biological disciplines seems essential for the field's transition from affirming patterns to understanding mechanisms and making robust predictions regarding future consequences of shifting phenologies.

Previous studies on wake flow visualization of live animals using DPIV have typically used low repetition rate lasers and 2D imaging. Repetition rates of around 10 Hz allow ~1 image per wingbeat in small birds and bats, and even fewer in insects. To accumulate data representing an entire wingbeat therefore requires the stitching-together of images captured from different wingbeats, and at different locations along the wing span for 3D-construction of wake topologies. A 200 Hz stereo DPIV system has recently been installed in the Lund University wind tunnel facility and the high-frame rate can be used to calculate all three velocity components in a cube, whose third dimension is constructed using the Taylor hypothesis. We studied two bat species differing in body size, Glossophaga soricina and Leptonycteris curasoa. Both species shed a tip vortex during the downstroke that was present well into the upstroke, and a vortex of opposite sign to the tip vortex was shed from the wing root. At the transition between upstroke/downstroke, a vortex loop was shed from each wing, inducing an upwash. Vorticity iso-surfaces confirmed the overall wake topology derived in a previous study. The measured dimensionless circulation, Γ/Uc, which is proportional to a wing section lift coefficient, suggests that unsteady phenomena play a role in the aerodynamics of both species.

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.

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.

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.