Testing an emerging paradigm in migration ecology shows surprising differences in efficiency between flight modes

To maximize fitness, flying animals should maximize flight speed while minimizing energetic expenditure. Soaring speeds of large-bodied birds are determined by flight routes and tradeoffs between minimizing time and energetic costs. Large raptors migrating in eastern North America predominantly glide between thermals that provide lift or soar along slopes or ridgelines using orographic lift (slope soaring). It is usually assumed that slope soaring is faster than thermal gliding because forward progress is constant compared to interrupted progress when birds pause to regain altitude in thermals. We tested this slope-soaring hypothesis using high-frequency GPS-GSM telemetry devices to track golden eagles during northbound migration. In contrast to expectations, flight speed was slower when slope soaring and eagles also were diverted from their migratory path, incurring possible energetic costs and reducing speed of progress towards a migratory endpoint. When gliding between thermals, eagles stayed on track and fast gliding speeds compensated for lack of progress during thermal soaring. When thermals were not available, eagles minimized migration time, not energy, by choosing energetically expensive slope soaring instead of waiting for thermals to develop. Sites suited to slope soaring include ridges preferred for wind-energy generation, thus avian risk of collision with wind turbines is associated with evolutionary trade-offs required to maximize fitness of time-minimizing migratory raptors.     

Individual variation in orientation promotes a 3000‐km latitudinal change in wintering grounds in a long‐distance migratory raptor

Migrating juvenile birds rely on endogenous information in choosing the direction in which to fly, but such input may be overridden by social interactions with experienced individuals. We tagged seven juvenile Short‐toed Eagles Circaetus gallicus with GPS transmitters in southern Italy. This trans‐Saharan migrant flies mainly by soaring and is therefore not well adapted to performing long water crossings. Five of the seven tagged juveniles used the longer but apparently safer route towards the Strait of Gibraltar, and two migrated along a southerly trajectory and subsequently spent the winter in Sicily, apparently forced to do so by the 150‐km‐wide Sicily Channel. One of these individuals took the longer route the following autumn. These results, combined with long‐term (15 years) visual field observations involving thousands of individuals, suggest that inexperienced Short‐toed Eagles may learn their migratory routes from experienced adults, whereas some of them migrate south in response to an innate orientation instinct. Transport costs, inherited information and geography apparently interact, forcing some Short‐toed Eagles to winter 3000 km to the north of the majority of their conspecifics.     

Narrow sea crossings present major obstacles to migrating Griffon Vultures Gyps fulvus

The flight behaviour of Griffon Vultures Gyps fulvus was studied at a major migration bottleneck, the Strait of Gibraltar in southernmost Spain, during the autumns of 2004 to 2007. The 14‐km‐wide sea channel significantly impeded the southern migration of the species into Africa, with many birds attempting repeated passage for weeks before crossing, and others not crossing at all and overwintering in Southern Spain. Water‐crossing attempts were restricted to times between 11:00 and 14:00 h on days with light or variable winds, or on days with strong winds from the north or west. No crossing attempts were made on days with strong winds from the south or east. Vultures attempted to cross the Strait in large flocks and never attempted to do so alone. Although 29% of the birds soared during crossing attempts, at least until they flew beyond visible range of approximately 4 km, most engaged in considerable flapping flight when attempting to cross. Overall, birds flying over water flapped more than 10 times as frequently as those flying over land prior to crossing attempts. Vultures did not flap continuously, but intermittently in brief bouts of flapping interspersed with periods of gliding or soaring flight. The number of flaps per bout over water was significantly greater than the number of flaps per bout over land. Vultures flying over water that flapped at rates of 20 flaps or more per minute typically aborted attempted crossings and returned to Spain in intermittent flapping and gliding flight. There are numerous reports of Vultures falling into the Strait and drowning while attempting to cross, as well as reports of returning Vultures collapsing on the beach having reached Spain in spring ( Barrios Partida 2006 ). Our observations indicate that passage of Griffon Vultures at the Strait of Gibraltar is limited by the species’ over‐water flapping‐flight abilities, including its inability to flap continuously for even short periods of time. We suggest that even relatively short sea crossings represent significant obstacles to migrating Vultures and discuss the implications of this limitation on the distribution and abundance of the species.     

Effect of wind,thermal convection,and variation in flight strategies on the daily rhythm and flight paths of migrating raptors at Georgia's Black Sea coast

Every autumn, large numbers of raptors migrate through geographical convergence zones to avoid crossing large bodies of water. At coastal convergence zones, raptors may aggregate along coastlines because of convective or wind conditions. However, the effect of wind and thermal convection on migrating raptors may vary depending on local landscapes and weather, and on the flight strategies of different raptors. From 20 August to 14 October 2008 and 2009, we studied the effect of cloud development and crosswinds on the flight paths of raptors migrating through the eastern Black Sea convergence zone, where coastal lowlands at the foothills of the Pontic Mountains form a geographical bottleneck 5‐km‐wide near Batumi, the capital of the Independent Republic of Ajaria in southwestern Georgia. To identify key correlates of local aggregation, we examined diurnal variation in migration intensity and coastal aggregation of 11 species of raptors categorized based on size and flight strategies. As reported at other convergence zones, migration intensity of large obligate‐soaring species peaked during the core period of thermal activity at mid‐day. When clouds developed over interior mountains and limited thermal convection, these large obligate‐soaring species aggregated near the coast. However, medium‐sized soaring migrants that occasionally use flapping flight did not aggregate at the coast when clouds over the mountains weakened thermal convection. Numbers of alternate soaring‐flapping harriers (Circus spp.) peaked during early morning, with these raptors depending more on flapping flight during a time of day with poor thermal convection. Small sparrowhawks (Accipiter spp.) aggregated at the coast during periods when winds blew offshore, suggesting aggregation caused by wind drift. Thus, weather conditions, including cloud cover and wind speed and direction, can influence the daily rhythm and flight paths of migrating raptors and, therefore, should be accounted for before inferring population trends from migration counts.     

The annual cycle of a trans-equatorial Eurasian-African passerine migrant: different spatio-temporal strategies for autumn and spring migration

The small size of the billions of migrating songbirds commuting between temperate breeding sites and the tropics has long prevented the study of the largest part of their annual cycle outside the breeding grounds. Using light-level loggers (geolocators), we recorded the entire annual migratory cycle of the red-backed shrike Lanius collurio, a trans-equatorial Eurasian-African passerine migrant. We tested differences between autumn and spring migration for nine individuals. Duration of migration between breeding and winter sites was significantly longer in autumn (average 96 days) when compared with spring (63 days). This difference was explained by much longer staging periods during autumn (71 days) than spring (9 days). Between staging periods, the birds travelled faster during autumn (356 km d(-1)) than during spring (233 km d(-1)). All birds made a protracted stop (53 days) in Sahelian sub-Sahara on southbound migration. The birds performed a distinct loop migration (22 000 km) where spring distance, including a detour across the Arabian Peninsula, exceeded the autumn distance by 22 per cent. Geographical scatter between routes was particularly narrow in spring, with navigational convergence towards the crossing point from Africa to the Arabian Peninsula. Temporal variation between individuals was relatively constant, while different individuals tended to be consistently early or late at different departure/arrival occasions during the annual cycle. These results demonstrate the existence of fundamentally different spatio-temporal migration strategies used by the birds during autumn and spring migration, and that songbirds may rely on distinct staging areas for completion of their annual cycle, suggesting more sophisticated endogenous control mechanisms than merely clock-and-compass guidance among terrestrial solitary migrants. After a century with metal-ringing, year-round tracking of long-distance migratory songbirds promises further insights into bird migration.     

Migration of red‐backed shrikes from the Iberian Peninsula: optimal or sub‐optimal detour?

The current Northern Hemisphere migration systems are believed to have arisen since the last glaciation. In many cases, birds do not migrate strait from breeding to non‐breeding areas but fly via a detour. All western European populations of red‐backed shrikes Lanius collurio are assumed to reach their southern African wintering grounds detouring via southeast Europe. Based on theoretical considerations under an optimality framework this detour is apparently optimal. Here, we use individual geolocator data on red‐backed shrikes breeding in Spain to show that these birds do indeed detour via southeast Europe en route to southern Africa where they join other European populations of red‐backed shrikes and return via a similar route in spring. Disregarding potential wind assistance, the routes taken for the tracked birds in autumn were not optimal compared to crossing the barrier directly. For spring migration the situation was quite different with the detour apparently being optimal. However, when considering potential wind assistance estimated total air distances during autumn migration were overall similar and the barrier crossing shorter along the observed routes. We conclude that considering the potential benefit of wind assistance makes the route via southeast Europe likely to be less risky in autumn. However, it cannot be ruled out that other factors, such as following a historical colonisation route could still be important.     

Real-time weather analysis reveals the adaptability of direct sea-crossing by raptors

Many animals seasonally travel between their breeding and wintering grounds. With their advanced mobility, birds often migrate over thousands of kilometres. Recently, satellite-tracking studies have revealed peculiar migration routes for some avian species at a global scale. However, the adaptability of such migration routes has not been clearly demonstrated. Using satellite-tracking data for 33 individuals, we show that the Japanese population of Oriental honey-buzzards (Pernis ptilorhynchus) directly crosses the 650-km-wide East China Sea during their autumn migration, although they fly a longer route around the sea rather than directly crossing it during their spring migration. By applying aerodynamic theory, we show that the buzzards could cross the sea by soaring and gliding flight. Moreover, using a high-resolution meteorological-prediction analysis, we demonstrate that the migratory trajectory of the birds strongly depends on the wind direction at their estimated locations. In the area, northeastern tailwinds blow stably only during autumn. Thermals were abundant ca. 500–1,000 m over the East China Sea in autumn, but that was not the case in spring. We suggest that the autumn-migration route across the East China Sea is likely to have evolved in response to the specific weather conditions over the sea. Animations showing movements of Oriental honey-buzzards and temporal change in weather conditions are available at: , , , , , and .     

Soaring across continents: decision‐making of a soaring migrant under changing atmospheric conditions along an entire flyway

Thermal soaring birds reduce flight‐energy costs by alternatingly gaining altitude in thermals and gliding across the earth's surface. To find out how soaring migrants adjust their flight behaviour to dynamic atmospheric conditions across entire migration routes, we combined optimal soaring migration theory with high‐resolution GPS tracking data of migrating honey buzzards Pernis apivorus and wind data from a global numerical atmospheric model. We compared measurements of gliding air speeds to predictions based on two distinct behavioural benchmarks for thermal soaring flight. The first being a time‐optimal strategy whereby birds alter their gliding air speeds as a function of climb rates to maximize cross‐country air speed over a full climb– glide cycle (V_opt). The second a risk‐averse energy‐efficient strategy at which birds alter their gliding air speed in response to tailwinds/headwinds to maximize the distance travelled in the intended direction during each glide phase (V_bgw). Honey buzzards were gliding on average 2.05 ms^{– 1} slower than V_opt and 3.42 ms^{– 1} faster than V_bgw while they increased air speeds with climb rates and reduced air speeds in tailwinds. They adopted flexible flight strategies gliding mostly near V_bgw under poor soaring conditions and closer to V_opt in good soaring conditions. Honey buzzards most adopted a time‐optimal strategy when crossing the Sahara, and at the onset of spring migration, where and when they met with the best soaring conditions. The buzzards nevertheless glided slower than V_opt during most of their journeys, probably taking time to navigate, orientate and locate suitable thermals, especially in areas with poor thermal convection. Linking novel tracking techniques with optimal migration models clarifies the way birds balance different tradeoffs during migration.     

African Odyssey project – satellite tracking of black storks Ciconia nigra breeding at a migratory divide

The African Odyssey project focuses on studying the migration of the black stork Ciconia nigra breeding at a migratory divide. In 1995–2001, a total of 18 black storks breeding in the Czech Republic were equipped with satellite (PTT) and VHF transmitters. Of them, 11 birds were tracked during at least one migration season and three birds were tracked repeatedly. The birds migrated either across western or eastern Europe to spend the winter in tropical west or east Africa, respectively. One of the juveniles made an intermediate route through Italy where it was shot during the first autumn migration. The mean distance of autumn migration was 6,227 km. The eastern route was significantly longer than the western one (7,000 km and 5,667 km respectively). Important stopover sites were discovered in Africa and Israel. Wintering areas were found from Mauritania and Sierra Leone in the west to Ethiopia and Central African Republic in the east and south. One of the storks migrating by the eastern migration route surprisingly reached western Africa. Birds that arrived early in the wintering areas stayed longer than those arriving later. On the average, birds migrating via the western route spent 37 d on migration compared to 80 d for birds migrating via the eastern route. The mean migration speed in the autumn was 126 km/d and the fastest stork flew 488 km/d when crossing the Sahara. The repeatedly tracked storks showed high winter site fidelity.     

Do Levant Sparrowhawks Accipiter brevipes also migrate at night?

Flight paths of visually identified Levant Sparrowhawks Accipiter brevipes on autumn migration were analysed with a tracking radar in the Arava Valley, Israel. This time of the year there are no significant numbers of other species with a similar wing-beat pattern. This wing-beat pattern was found not only in daytime but also frequently at night. It is suggested that the Levant Sparrowhawk uses two strategies of migration: (1) soaring and gliding to reduce energy consumption; (2) flapping flight to reduce time spent on migration. The latter may be more important towards the end of the migratory season and/or when birds have become separated from the main migratory stream.     

The gliding speed of migrating birds: slow and safe or fast and risky?

Aerodynamic theory postulates that gliding airspeed, a major flight performance component for soaring avian migrants, scales with bird size and wing morphology. We tested this prediction, and the role of gliding altitude and soaring conditions, using atmospheric simulations and radar tracks of 1346 birds from 12 species. Gliding airspeed did not scale with bird size and wing morphology, and unexpectedly converged to a narrow range. To explain this discrepancy, we propose that soaring‐gliding birds adjust their gliding airspeed according to the risk of grounding or switching to costly flapping flight. Introducing the Risk Aversion Flight Index (RAFI, the ratio of actual to theoretical risk‐averse gliding airspeed), we found that inter‐ and intraspecific variation in RAFI positively correlated with wing loading, and negatively correlated with convective thermal conditions and gliding altitude, respectively. We propose that risk‐sensitive behaviour modulates the evolution (morphology) and ecology (response to environmental conditions) of bird soaring flight.     

Variable detours in long‐distance migration across ecological barriers and their relation to habitat availability at ground

Migration detours, the spatial deviation from the shortest route, are a widespread phenomenon in migratory species, especially if barriers must be crossed. Moving longer distances causes additional efforts in energy and time, and to be adaptive, this should be counterbalanced by favorable condition en route. We compared migration patterns of nightingales that travelled along different flyways from their European breeding sites to the African nonbreeding sites. We tested for deviations from shortest routes and related the observed and expected routes to the habitat availability at ground during autumn and spring migration. All individuals flew detours of varying extent. Detours were largest and seasonally consistent in western flyway birds, whereas birds on the central and eastern flyways showed less detours during autumn migration, but large detours during spring migration (eastern flyway birds). Neither migration durations nor the time of arrival at destination were related to the lengths of detours. Arrival at the breeding site was nearly synchronous in birds flying different detours. Flying detours increased the potential availability of suitable broad‐scale habitats en route only along the western flyway. Habitat availability on observed routes remained similar or even decreased for individuals flying detours on the central or the eastern flyway as compared to shortest routes. Thus, broad‐scale habitat distribution may partially explain detour performance, but the weak detour‐habitat association along central and eastern flyways suggests that other factors shape detour extent regionally. Prime candidate factors are the distribution of small suitable habitat patches at local scale as well as winds specific for the region and altitude.     

Age-dependent migration strategy in honey buzzards Pernis apivorus tracked by satellite

Six adult and three juvenile honey buzzards Pernis apivorus were radio-tracked by satellite during autumn migration from southwestern Sweden. All adults crossed the Mediterranean Sea at the Strait of Gibraltar and continued across the Sahara desert to winter in West Africa, from Sierra Leone to Cameroon. Analysing three main steps of the migration, (1) from the breeding site to the southern Mediterranean region, (2) across the Sahara and (3) from the southern Sahara to the wintering sites, the adults changed direction significantly between these steps, and migrated along a distinct large-scale detour. In contrast, the juveniles travelled in more southerly directions, crossed the Mediterranean Sea at various places, but still ended up in the same wintering areas as the adults. Average speeds maintained on travelling days were similar for the two age groups, about 170 km/day in Europe, 270 km/day across Sahara and 125 km/day in Africa south of Sahara. However, as the adults used fewer stopover days en route, they maintained higher mean overall speeds and completed migration in a shorter time (42 days) than the juveniles (64 days). Although the juveniles set out on more direct courses towards the wintering grounds, they did not cover significantly shorter distances than the adults, as they tended to show a larger directional scatter between shorter flight segments. The results corroborate previous suggestions that adult and juvenile honey buzzards follow different routes during autumn migration, and that the birds change migration strategy during their lifetime. While juveniles may use individual vector orientation, social influences and learning may be of great importance for the detour migration of adults. The remarkable and distinct age-dependent shift in migratory route and orientation of the honey buzzard provides a challenging evolutionary problem.     

Regional wind patterns likely shape a seasonal migration detour

Migrating animals should optimise time and energy use when migrating, travelling directly to their destination. Detours from the most direct route may arise however because of barriers and weather conditions. Identifying how such situations arise from variable weather conditions is crucial to understand population response in the light of increased anthropogenic climate change. Here we used light-level geolocators to follow Cyprus wheatears for their full annual cycle in two separate years migrating between Cyprus, over the Mediterranean and the Sahara to winter in north–east sub-Saharan Africa. We predicted that any route detours would be related to wind conditions experienced during migration. We found that spring migration for all birds included an eastern detour, whilst autumn migrations were direct across the Sahara. The direct autumn migration was likely a consequence of consistent tail-winds, whilst the eastern detour in spring is likely to be more efficient given the wind conditions which are against a direct route. Such variable migration routes shaped by coincidence with prevailing winds are probably common suggesting that some birds may be able to adapt to future changes in wind conditions.     

Limitations and mechanisms influencing the migratory performance of soaring birds

Migration is costly in terms of time, energy and safety. Optimal migration theory suggests that individual migratory birds will choose between these three costs depending on their motivation and available resources. To test hypotheses about use of migratory strategies by large soaring birds, we used GPS telemetry to track 18 adult, 13 sub‐adult and 15 juvenile Golden Eagles Aquila chrysaetos in eastern North America. Each age‐class had potentially different motivations during migration. During spring, the migratory performance (defined here as the directness of migratory flight) of adults was higher than that of any other age‐classes. Adults also departed earlier and spent less time migrating. Together, these patterns suggest that adults were primarily time‐limited and the other two age‐classes were energy‐limited. However, adults that migrated the longest distances during spring also appeared to take advantage of energy‐conservation strategies such as decreasing their compensation for wind drift. During autumn, birds of all age‐classes were primarily energy‐minimizers; they increased the length of stopovers, flew less direct routes and migrated at a slower pace than during spring. Nonetheless, birds that departed later in autumn flew more directly, indicating that time limitations may have affected their decision‐making. During both seasons, juveniles had the lowest performance, sub‐adults intermediate performance and adults the highest performance. Our results show age‐ and seasonal variation in time and energy‐minimization strategies that are not necessarily exclusive of one another. Beyond time and energy, a complex suite of factors, including weather, experience and navigation ability, influences migratory performance and decision‐making.     

Flight mode affects allometry of migration range in birds

Billions of birds migrate to exploit seasonally available resources. The ranges of migration vary greatly among species, but the underlying mechanisms are poorly understood. I hypothesise that flight mode (flapping or soaring) and body mass affect migration range through their influence on flight energetics. Here, I compiled the tracks of migratory birds (196 species, weighing 12–10 350 g) recorded by electronic tags in the last few decades. In flapping birds, migration ranges decreased with body mass, as predicted from rapidly increasing flight cost with increasing body mass. The species with higher aspect ratio and lower wing loading had larger migration ranges. In soaring birds, migration ranges were mass‐independent and larger than those of flapping birds, reflecting their low flight costs irrespective of body mass. This study demonstrates that many animal‐tracking studies are now available to explore the general patterns and the underlying mechanisms of animal migration.     

Flight strategies of migrating raptors; a comparative study of interspecific variation in flight characteristics

The comparison of flight styles and flight parameters of migrating raptors in Israel revealed the following. (1) Climbing rate in thermal circling did not differ between species, indicating that chiefly the strength of thermal updrafts determined the climbing rate and that morphological features were less relevant. (2) In interthermal gliding, air speed was positively and gliding angle negatively related to the species' average body mass. Heavier species glided faster and had smaller gliding angles. (3) In soaring and gliding flight, cross-country speed relative to the air was positively related to the species' body mass; it was obviously the result of the gliding ability increasing with body mass. (4) Eagles and buzzards used soaring and gliding flight for more than 95% of the observation time. Additional soaring in a straight line whilst gliding was extensively used by the Steppe Eagle Aquila nipalensis, Lesser Spotted Eagle Aquila pomarina and Booted Eagle Hieraætus pennatus and even more frequently by the resident species, the Griffon Vulture Gyps fulvus and Shorttoed Eagle Circaetus gallicus. Smaller species, such as the Levant Sparrowhawk Accipiter brevipes, harriers (Circus sp.) and small falcons (Falco sp.). showed the highest proportion of flapping and gliding flight (9–33%). (5) In a comparison of the flight parameters and proportions of flight styles, a cluster analysis distinguished two main groups: The first consisted of Montagu's Harrier Circus pygargus, Pallid Harrier Circus macrourus, Levant Sparrowhawk and small falcons; their flight behaviour was characterized by both the high proportion of flapping and the low gliding performance. The second group comprised the typical soaring migrants: Steppe Eagle, Lesser Spotted Eagle, Booted Eagle, Steppe Buzzard Buteo buteo vulpinus, Honey Buzzard Pernis apivorus and Egyptian Vulture Neophron percnopterus, and they had very similar flight behaviour and were closely clustered. The Black Kite Milvus migrans and Marsh Harrier Circus aeruginosus were intermediate between typical soarers and flappers. The two resident species, Griffon Vulture and Short-toed Eagle, were grouped separately from the soaring migrants.     

Wing size but not wing shape is related to migratory behavior in a soaring bird

Both wing size and wing shape affect the flight abilities of birds. Intra and inter‐specific studies have revealed a pattern where high aspect ratio and low wing loading favour migratory behaviour. This, however, have not been studied in soaring migrants. We assessed the relationship between the wing size and shape and the characteristics of the migratory habits of the turkey vulture Cathartes aura, an obligate soaring migrant. We compared wing size and shape with migration strategy among three fully migratory, one partially migratory and one non‐migratory (resident) population distributed across the American continent. We calculated the aspect ratio and wing loading using wing tracings to characterize the wing morphology. We used satellite‐tracking data from the migratory populations to calculate distance, duration, speed and altitude during migration. Wing loading, but not aspect ratio, differed among the populations, segregating the resident population from the completely migratory ones. Unlike what has been reported in species using flapping flight during migration, the migratory flight parameters of turkey vultures were not related to the aspect ratio. By contrast, wing loading was related to most flight parameters. Birds with lower wing loading flew farther, faster, and higher during their longer journeys. Our results suggest that wing morphology in this soaring species enables lower‐cost flight, through low wing‐loading, and that differences in the relative sizes of wings may increase extra savings during migration. The possibility that wing shape is influenced by foraging as well as migratory flight is discussed. We conclude that flight efficiency may be improved through different morphological adaptations in birds with different flight mechanisms.     

Migration routes and strategies in a highly aerial migrant, the common swift Apus apus, revealed by light-level geolocators

The tracking of small avian migrants has only recently become possible by the use of small light-level geolocators, allowing the reconstruction of whole migration routes, as well as timing and speed of migration and identification of wintering areas. Such information is crucial for evaluating theories about migration strategies and pinpointing critical areas for migrants of potential conservation value. Here we report data about migration in the common swift, a highly aerial and long-distance migrating species for which only limited information based on ringing recoveries about migration routes and wintering areas is available. Six individuals were successfully tracked throughout a complete migration cycle from Sweden to Africa and back. The autumn migration followed a similar route in all individuals, with an initial southward movement through Europe followed by a more southwest-bound course through Western Sahara to Sub-Saharan stopovers, before a south-eastward approach to the final wintering areas in the Congo basin. After approximately six months at wintering sites, which shifted in three of the individuals, spring migration commenced in late April towards a restricted stopover area in West Africa in all but one individual that migrated directly towards north from the wintering area. The first part of spring migration involved a crossing of the Gulf of Guinea in those individuals that visited West Africa. Spring migration was generally wind assisted within Africa, while through Europe variable or head winds were encountered. The average detour at about 50% could be explained by the existence of key feeding sites and wind patterns. The common swift adopts a mixed fly-and-forage strategy, facilitated by its favourable aerodynamic design allowing for efficient use of fuel. This strategy allowed swifts to reach average migration speeds well above 300 km/day in spring, which is higher than possible for similar sized passerines. This study demonstrates that new technology may drastically change our views about migration routes and strategies in small birds, as well as showing the unexpected use of very limited geographical areas during migration that may have important consequences for conservation strategies for migrants.     

Around the Mediterranean: an extreme example of loop migration in a long‐distance migratory passerine

An important issue in migration research is how small‐bodied passerines pass over vast geographical barriers; in European–African avian migration, these are represented by the Mediterranean Sea and the Sahara Desert. Eastern (passing eastern Mediterranean), central (passing Apennine Peninsula) and western (via western Mediterranean) major migration flyways are distinguished for European migratory birds. The autumn and spring migration routes may differ (loop migration) and there could be a certain level of individual flexibility in how individuals navigate themselves during a single migration cycle. We used light‐level loggers to map migration routes of barn swallows Hirundo rustica breeding in the centre of a wide putative contact zone between the northeastern and southernwestern European populations that differ in migration flyways utilised and wintering grounds. Our data documented high variation in migration patterns and wintering sites of tracked birds (n = 19 individuals) from a single breeding colony, with evidence for loop migration in all but one of the tracked swallows. In general, two migratory strategies were distinguished. In the first, birds wintering in a belt stretching from southcentral to southern Africa that used an eastern route for both the spring and autumn migration, then shifted their spring migration eastwards (anti‐clockwise loops, n = 12). In the second, birds used an eastern or central route to their wintering grounds in central Africa, shifting the spring migration route westward (clockwise loops, n = 7). In addition, we observed an extremely wide clockwise loop migration encompassing the entire Mediterranean, with one individual utilising both the eastern (autumn) and western (spring) migratory flyway during a single annual migration cycle. Further investigation is needed to ascertain whether clockwise migratory loops encircling the entire Mediterranean also occur other small long‐distance passerine species.