Drought in Africa caused delayed arrival of European songbirds.

Despite an overall advancement in breeding area arrival, one of the latest spring arrivals in northwest Europe since 1950 of several trans-Saharan songbird species occurred in 2011. Year-round tracking of red-backed shrikes and thrush nightingales revealed that the cause of the delay was a prolongation of stopover time during spring migration at the Horn of Africa, which was affected by extreme drought. Our results help to establish a direct link at the individual level between changes in local climate during migration and arrival and breeding condition in Europe thousands of kilometers further north.

Flight speeds of swifts (Apus apus): seasonal differences smaller than expected.

We have studied the nocturnal flight behaviour of the common swift (Apus apus L.), 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.

Confronting the winds: orientation and flight behaviour of roosting swifts, Apus apus.

Swifts, Apus apus, spend the night aloft and this offers an opportunity to test the degree of adaptability of bird orientation and flight to different ecological situations. We predicted the swifts' behaviour by assuming that they are adapted to minimize energy expenditure during the nocturnal flight and during a compensatory homing flight if they become displaced by wind. We tested the predictions by recording the swifts' altitudes, speeds and directions under different wind conditions with tracking radar; we found an agreement between predictions and observations for orientation behaviour, but not for altitude and speed regulation. The swifts orientated consistently into the head wind, with angular concentration increasing with increasing wind speed. However, contrary to our predictions, they did not select altitudes with slow or moderate winds, nor did they increase their airspeed distinctly when flying into strong head winds. A possible explanation is that their head-wind orientation is sufficient to keep nocturnal displacement from their home area within tolerable limits, leaving flight altitude to be determined by other factors (correlated with temperature), and airspeed to show only a marginal increase in strong winds. The swifts were often moving "backwards", heading straight into the wind but being overpowered by wind speeds exceeding their airspeed. The regular occurrence of such flights is probably uniquely associated with the swifts' remarkable habit of roosting on the wing.

Detours in bird migration.

Bird migration routes often follow detours where passages across ecological barriers are reduced in extent. This occurs in spite of the fact that long barrier crossings are within the birds' potential flight range capacity. Long-distance flights are associated with extra energy costs for transport of the heavy fuel loads required. This paper explores how important the fuel transport costs, estimated on the basis of flight mechanics, may be to explain detours for birds migrating by flapping flight. Maximum detours in relation to expanse of the barrier are predicted for cases where birds travel along the detour by numerous short flights and small fuel reserves, divide the detour into a limited number of flight steps, and where a reduced barrier passage is included in the detour. The principles for determining the optimum route, often involving a shortcut across part of the barrier, are derived. Furthermore, the effects of differences in fuel deposition rates and in transport costs for the profitability of detours are briefly considered. An evaluation of a number of observed and potential detours in relation to the general predictions of maximum detours, indicates that reduction of fuel transport costs may well be a factor of widespread importance for the evolution of detours in bird migration at wide ecological barriers.

Migration along orthodromic sun compass routes by arctic birds.

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.

Bird orientation at high latitudes: flight routes between Siberia and North America across the Arctic Ocean

Bird migration and orientation at high latitudes are of special interest because of the difficulties associated with different compass systems in polar areas and because of the considerable differences between flight routes conforming to loxodromes (rhumblines) or orthodromes (great circle routes). Regular and widespread east-north-east migration of birds from the northern tundra of Siberia towards North America across the Arctic Ocean (without landmark influences) were recorded by ship-based tracking radar studies in July and August. Field observations indicated that waders, including species such as Phalaropusfulicarius and Calidris melanotos, dominated, but also terns and skuas may have been involved. Analysis of flight directions in relation to the wind showed that these movements are not caused by wind drift. Assuming possible orientation principles based on celestial or geomagnetic cues, different flight trajectories across the Arctic Ocean were calculated: geographical loxodromes, sun compass routes, magnetic loxodromes and magnetoclinic routes. The probabilities of these four alternatives are evaluated on the basis of both the availability of required orientation cues and the predicted flight paths. This evaluation supports orientation along sun compass routes. Because of the longitudinal time displacement sun compass routes show gradually changing compass courses in close agreement with orthodromes. It is suggested that an important migration link between Siberia and North American stopover sites 1000-2500km apart across the Arctic Ocean has evolved based on sun compass orientation along orthodrome-like routes.

A new low-turbulence wind tunnel for bird flight experiments at Lund University, Sweden

A new wind tunnel for experiments on bird flight was completed at Lund University, Sweden, in September 1994. It is a closed-circuit design, with a settling section containing five screens and a contraction ratio of 12.25. The test section is octagonal, 1.20 m wide by 1.08 m high. The first 1.2 m of its length is enclosed by acrylic walls, and the last 0.5 m is open, giving unrestricted access. Experiments can be carried out in both the open and closed parts, and comparison between them can potentially be used to measure the lift effect correction. The fan is driven by an a.c. motor with a variable-frequency power supply, allowing the wind speed to be varied continuously from 0 to 38 m s-1. The whole machine can be tilted to give up to 8 ° descent and 6 ° climb. A pitot-static survey in the test section showed that the air speed was within ±1.3 % of the mean at 116 out of 119 sample points, exceeding this deviation at only three points at the edges. A hot-wire anemometer survey showed that the turbulence level in the closed part of the test section was below 0.04 % of the wind speed throughout most of the closed part of the test section, rising to approximately 0.06 % in the middle of the open part. No residual rotation from the fan could be detected in the test section. No decrease in wind speed was detectable beyond 3 cm from the side walls of the closed part, and turbulence was minimal beyond 10 cm from the walls. The installation of a safety net at the entrance to the test section increased the turbulence level by a factor of at least 30, to 1.2 % longitudinally and 1.0 % transversely.

Optimum fuel loads in migratory birds: distinguishing between time and energy minimization

By combining the potential flight range of fuel with different migration policies, the optimum departure fuel load for migratory birds can be calculated. We evaluate the optimum departure fuel loads associated with minimization of three different currencies: (1) overall time of migration, (2) energy cost of transport and (3) total energy coast of migration. Predicted departure loads are highest for (1), lowest for (2) and intermediate for (3). Further, currencies (1) and (3) show departure loads dependent on the fuel accumulation rate at stopovers, while (2) is not affected by variation in the rate of fuel accumulation. Furthermore, fuel loads optimized with respect to currency (3) will differ depending on the size (body mass) of the bird and the energy density of the fuel. We review ecological situations in which the various currencies may apply, and suggest how a combination of stopover decisions and observations of flight speed may be used to decide among the three cases of migration policies. Finally, we calculate that the total energy cost of migration is roughly divided between flight and stopover as 1:2. The total time of migration is similarly divided between flight and stopover as 1:7, probably with a relatively longer stopover time in larger species. Hence, we may expect strong selection pressures to optimize the fuel accumulation strategies during stopover episodes.Copyright 1997 Academic Press Limited Copyright 1997 Academic Press Limited

The geographical scale factor in orientation of migrating birds

Migration routes of birds throw light on orientation performance at different geographic scales, over distances ranging from a few kilometres to more than 104 km. Detailed knowledge about the flight routes may be used to test predictions about optimal orientation according to theoretical principles and about the use of compasses based on celestial or magnetic cues. Ringing recoveries demonstrate that the migratory journey of many species, such as the wheatear and willow warbler, is divided into successive legs with different main orientation. Autumn and spring migration routes are often different, sometimes diverging on a continental scale. Aerial radiotracking of whooping cranes in North America and satellite tracking of brent geese migrating from Iceland across the Greenland ice cap point to the significant role of large-scale topography for the shaping of migration routes. Compass and position control are also required, e.g. during long passages across featureless sea or ice, but how these elements are integrated into the birds' orientation system remains unclear. Radar studies from the Arctic Ocean illustrate the importance of map projections for interpreting flight paths and suggest that birds accomplish approximate great circle orientation. Gradual course changes shown by migrating knots monitored by radar in Scandinavia are at variance with expected changes if the birds were to use a star, sun or magnetic compass over longer distances. Accurate recording of short flight segments shows how flying birds respond to visual, audible and electromagnetic cues, and also documents orientation precision and capacity to integrate rapidly shifting courses into a consistent resulting orientation. Analyses of flight patterns are crucial for understanding how birds find and follow their migration routes over different ranges of geographical scale.

Optimal fat loads in migrating birds: a test of the time-minimization hypothesis.

We tested the hypothesis that birds are selected to minimize the time spent on migration, that is, to migrate as fast as possible. Optimal fat loads in time-selected migration were predicted for different rates of fat accumulation at stopover sites. We analyzed departure fat loads of migrating bluethroats Luscinia svecica svecica, experimentally provided with extra food at a stopover site, and of migrating rufous hummingbirds Selasphorus rufus, which showed considerable individual variation in fat-deposition rate, in relation to these predictions. We found qualitative agreement with the time-minimization hypothesis. However, quantitative agreement requires that specific assumptions be fulfilled for both species: (1) consistent differences in expected speed of migration should exist between different individuals of the same species and/or (2) the expected speed of migration should increase along the route Both of these assumptions are probably valid, and ringing data suggest an increase in bluethroat autumn migration speed along the route Physiological and flight mechanical constraints will prevent birds from depositing excessively large amounts of fuel. These assumptions and constraints should be taken into account in future critical tests of the hypothesis that natural selection operates to maximize the speed of migration.

Ecological causes and consequences of bird orientation.

An advanced orientation capability offers possibilities for birds to optimize movement patterns in a wide variety of ecological situations. The adaptive significance of various patterns of angular dispersion and of orientation responses to topography and sociality are elucidated. The orientation capacity is characterized by flexibility, exemplified by reorientation, promoting safety and restoration of fat reserves during migration. There are also limitations to the orientation process, leading to costs of migration through mis- or disorientation, and to constraints on the evolution of routes and timing of migratory flights. Young migrants may acquire an erroneous compass sense, and misorient several thousands of kilometers off their normal course. Widespread and dense fog of long duration causes disorientation and mortality among land birds migrating over the sea. Orientational constraints in the evolution of migration routes may be most easily disclosed at high geographic and magnetic latitudes. Here the birds are faced with special difficulties in using their celestial as well as their magnetic compasses. The sun compass could be used for great circle orientation, but observed spring flight trajectories of high-arctic waders and geese seem to conform with rhumbline routes.

Bird flight and optimal migration.

The development of the mechanical and aerodynamical theory of bird flight has greatly stimulated research at widely different levels in the field of bird movement. Recent work has shown that the drag of bird bodies is less than was previously assumed. Furthermore, the structure and circulation of wingtip vortices in the wake of flying birds have been revealed, with implications for estimating flight performance on the basis of vortex theory. Predictions about optimal speed and flight behaviour have been successfully tested by field studies using optical and radar registration. Flight theory also allows predictions about optimal fuel deposition rules for migrating birds. Research about bird flight, with the dynamic interplay between theoretical development and empirical work in biophysics, physiology and ecology, represents a fine example of a highly successful use of the optimality approach in biology.