The migration of the great snipe Gallinago media: intriguing variations on a grand theme

The migration of the great snipe Gallinago media was previously poorly known. Three tracks in 2010 suggested a remarkable migratory behaviour including long and fast overland non‐stop flights. Here we present the migration pattern of Swedish male great snipes, based on 19 individuals tracked by light‐level geolocators in four different years. About half of the birds made stopover(s) in northern Europe in early autumn. They left the breeding area 15 d earlier than those which flew directly to sub‐Sahara, suggesting two distinct autumn migration strategies. The autumn trans‐Sahara flights were on average 5500 km long, lasted 64 h, and were flown at ground speeds of 25 m s^?1 (90 km h^?1). The arrival in the Sahel zone of west Africa coincided with the wet season there, and the birds stayed for on average three weeks. The birds arrived at their wintering grounds around the lower stretches of the Congo River in late September and stayed for seven months. In spring the great snipes made trans‐Sahara flights of similar length and speed as in autumn, but the remaining migration through eastern Europe was notably slow. All birds returned to the breeding grounds within one week around mid‐May. The annual cycle was characterized by relaxed temporal synchronization between individuals during the autumn–winter period, with maximum variation at the arrival in the wintering area. Synchronization increased in spring, with minimum time variation at arrival in the breeding area. This suggests that arrival date in the breeding area is under strong stabilizing selection, while there is room for more flexibility in autumn and arrival to the wintering area. The details of the fast non‐stop flights remain to be elucidated, but the identification of the main stopover and wintering areas is important for future conservation work on this red‐listed bird species.     

Skipping‐type migration in a small Arctic wader,the Temminck's stint Calidris temminckii

By using morphometric data and geolocator tracking we investigated fuel loads and spatio‐temporal patterns of migration and non‐breeding in Temminck's stints Calidris temminckii. Body masses in stints captured at autumn stopover sites from Scandinavia to northern Africa were generally not much higher than during breeding and did not vary geographically. Thus, we expected migrating stints to make several stopovers and either circumventing the Sahara desert with low fuel loads or fuelling at north African stopover sites before desert crossing. Geolocation revealed that birds (n = 6) departed their Norwegian breeding site in the last part of July and all but one migrated south‐west over continental western Europe. A single bird headed south‐east to the Balkan Peninsula where the geolocator died. As predicted, southbound migration proceeded in a typical skipping manner with 1–4 relatively short stopovers (median 4 d) during 10–27 d of migration before reaching north‐west Africa. Here birds spent 11–20 d before crossing the Sahara. The non‐breeding sites were located at or near the Niger River in Mali and were occupied continuously for more than 215 d with no indications of itinerancy. Spring migration commenced in late April/early May when birds crossed the desert and used stopover sites in the western Mediterranean basin in a similar manner as during autumn. The lowest body masses were recorded in spring at islands in the central Mediterranean basin, indicating that crossing the Sahara and Mediterranean barriers is exhausting to these birds. Hence, the skipping‐type pattern of migration revealed by geolocators is likely to be natural in this species and not an effect of instrumentation.     

Discovering the migration and non‐breeding areas of sand martins and house martins breeding in the Pannonian basin (central‐eastern Europe)

The central‐eastern European populations of sand martin and house martin have declined in the last decades. The drivers for this decline cannot be identified as long as the whereabouts of these long distance migrants remain unknown outside the breeding season. Ringing recoveries of sand martins from central‐eastern Europe are widely scattered in the Mediterranean basin and in Africa, suggesting various migration routes and a broad non‐breeding range. The European populations of house martins are assumed to be longitudinally separated across their non‐breeding range and thus narrow population‐specific non‐breeding areas are expected. By using geolocators, we identified for the first time, the migration routes and non‐breeding areas of sand martins (n = 4) and house martins (n = 5) breeding in central‐eastern Europe. In autumn, the Carpathian Bend and northern parts of the Balkan Peninsula serve as important pre‐migration areas for both species. All individuals crossed the Mediterranean Sea from Greece to Libya. Sand martins spent the non‐breeding season in northern Cameroon and the Lake Chad Basin, within less than a 700 km radius, while house martins were widely scattered in three distinct regions in central, eastern, and southern Africa. Thus, for both species, the expected strength of migratory connectivity could not be confirmed. House martins, but not sand martins, migrated about twice as fast in spring compared to autumn. The spring migration started with a net average speed of > 400 km d^–1 for sand martins, and > 800 km d^–1 for house martins. However, both species used several stopover sites for 0.5–4 d and were stationary for nearly half of their spring migration. Arrival at breeding grounds was mainly related to departure from the last sub‐Saharan non‐breeding site rather than distance, route, or stopovers. We assume a strong carry‐over effect on timing in spring.     

First geolocator tracks of Swedish red‐necked phalaropes reveal the Scandinavia‐Arabian Sea connection

We studied migration and wintering patterns of a wader with a pelagic lifestyle during the non‐breeding period, the red‐necked phalarope Phalaropus lobatus. Using light‐level geolocation, we obtained three full annual tracks and one autumn migration track of male red‐necked phalaropes caught during breeding in Scandinavia. These tracks confirmed expectations that individuals from the Scandinavian population winter in the Arabian Sea. Migration was accomplished in two to four migration leaps, staging for a few days in the Gulf of Finland (autumn) or the southern Baltic Sea (spring) and for up to a month in or near the Black and Caspian Sea (autumn and spring). In addition, travel speeds suggested that only the flights between the Baltic and Black/Caspian Sea are non‐stop, and thus the birds seem to make additional short stops during the other flights. Stopover time in the Black/Caspian Sea is only 8–10 d in spring but up to 36 d in autumn, which is longer than expected if only used for pre‐migratory fattening to cover the ca 2000 km to the Gulf of Oman. After entering the Arabian Sea via the Gulf of Oman, birds dispersed over the entire presumed winter range. Winter movements appear to correspond to the spatio‐temporal patterns in primary production linked to seasonally changing monsoon winds. These are not only the first tracks of Scandinavian red‐necked phalaropes, but also the first seabird tracks in the Arabian Sea, one of the most productive and dynamic marine areas on the planet.     

Migration of the Little Ringed Plover Charadrius dubius breeding in South Sweden tracked by geolocators

Capsule Little Ringed Plovers breeding in South Sweden migrate towards the southeast in the autumn, via the Middle East, to winter in Saharan and sub-Saharan locations or in India, while the spring migration is more directly towards the north.

Aims To study the migration routes and wintering area of Little Ringed Plovers (Charadrius dubius) breeding in South Sweden, and to investigate the migration strategy and speed for this little studied shorebird.

Methods We use light-level geolocators to track the year-round movements of Little Ringed Plovers breeding in South Sweden.

Results Autumn migration proceeded towards the southeast, in three birds via lengthy stopovers in the Middle East, followed by movements towards the west and southwest to final winter destinations in Africa, while one male made a long stopover in northwestern Iran before migrating to India. The birds wintering in Africa probably stayed at freshwater locations in the Sahara or just south or north of the Sahara. Spring migration was more directly back to the breeding area. Overall migration speeds were similar during autumn and spring migration at about 189 and 209?km/day, respectively. The migration was carried out mainly as many short flights between stopovers. In particular, autumn migration was longer than the direct distance between breeding and wintering sites.

Conclusions This study shows that the geolocator method can successfully be used with relatively small (40?g) shorebirds. We found that a local population of Little Ringed Plover may have widely differing wintering sites (low connectivity), from sub-Saharan Africa to the Indian subcontinent. The migration strategy of the Little Ringed Plover, with multiple short flights, deviates from that of many other long-distance migrating shorebirds that, instead, make one or a few long flights.     

Seasonal migration strategies of Common Ringed Plovers Charadrius hiaticula

How individual birds schedule their movements and use different sites during the non‐breeding season are fundamental issues in avian migration ecology, and studies have often revealed strong seasonal variation in such strategies. Using geolocators we tracked Common Ringed Plovers Charadrius hiaticula from northern Norway to West Africa and back to assess whether there were differences in migratory speed, duration and stopover use between autumn and spring migration and whether birds used multiple sites during the non‐breeding season. Although the pace of migration was similar between autumn and spring, the length of flight bouts and duration of the preceding stopovers were positively correlated only in autumn. Four of five birds showed a marked southward movement in mid‐winter.     

Annual Cycle and Migration Strategies of a Trans-Saharan Migratory Songbird: A Geolocator Study in the Great Reed Warbler

Recent technological advancements now allow us to obtain geographical position data for a wide range of animal movements. Here we used light-level geolocators to study the annual migration cycle in great reed warblers (Acrocephalus arundinaceus), a passerine bird breeding in Eurasia and wintering in sub-Saharan Africa. We were specifically interested in seasonal strategies in routes and schedules of migration. We found that the great reed warblers (all males, no females were included) migrated from the Swedish breeding site in early August. After spending up to three weeks at scattered stopover sites in central to south-eastern Europe, they resumed migration and crossed the Mediterranean Sea and Sahara Desert without lengthy stopovers. They then spread out over a large overwintering area and each bird utilised two (or even three) main wintering sites that were spatially separated by a distinct mid-winter movement. Spring migration initiation date differed widely between individuals (1-27 April). Several males took a more westerly route over the Sahara in spring than in autumn, and in general there were fewer long-distance travels and more frequent shorter stopovers, including one in northern Africa, in spring. The shorter stopovers made spring migration on average faster than autumn migration. There was a strong correlation between the spring departure dates from wintering sites and the arrival dates at the breeding ground. All males had a high migration speed in spring despite large variation in departure dates, indicating a time-minimization strategy to achieve an early arrival at the breeding site; the latter being decisive for high reproductive success in great reed warblers. Our results have important implications for the understanding of long-distance migrants’ ability to predict conditions at distant breeding sites and adapt to rapid environmental change.     

Spring migration strategies of Whinchat Saxicola rubetra when successfully crossing potential barriers of the Sahara and the Mediterranean Sea

The flexibility for migrant land birds to be able to travel long distances rapidly without stopovers, and thus to cross wide inhospitable areas such as deserts and oceans, is likely to be a major determinant of their survival during migration. We measured variation in flight distance, speed and duration of major stopovers (more than 2 days), using geolocator tracks of 35 Whinchats Saxicola rubetra that migrated successfully from central Nigeria to Eastern Europe in spring, and examined how these measures changed, or depended on age, when crossing the barriers of the Sahara or the Mediterranean Sea. In all, 31% of Whinchats crossed at least the Sahara and the Mediterranean before a major stopover and 17% travelled over 4751 km on average without any major stopovers. Flight distance and speed during, and duration of major stopovers after, crossing the Mediterranean Sea were indistinguishable from migration over Continental Europe. Speed during a migration leg was lowest crossing Continental Europe and fastest, with longer duration major stopovers afterwards, when crossing the Sahara, but there was much individual variation, and start date of migration was also a good predictor of stopover duration. As the distance travelled during a leg increased, so major stopover duration afterwards increased (1 day for every 1000 km), but the speed of travel during the leg had no effect. There were no differences in any migration characteristics with age, other than an earlier start date for adult birds. The results suggest that adaptive shortening or even dropping of daily stopovers may occur often, allowing rapid, long‐distance migration at the cost of major stopovers afterwards, but such behaviour is not restricted to or always found when crossing barriers, even for birds on their first spring migration. The results may highlight the importance of stopover sites rather than barrier width as the likely key component to successful migration. Individual variation in spring migration may indicate that small passerine migrants like Whinchats may be resilient to future changes in the extent of barriers they encounter, although this may not be true of first autumn migrations or if stopover sites are lost.     

Migration timing and routes,and wintering areas of Flammulated Owls

Determining patterns in annual movements of animals is an important component of population ecology, particularly for migratory birds where migration timing and routes, and wintering habitats have key bearing on population dynamics. From 2009 to 2011, we used light‐level geolocators to document the migratory movements of Flammulated Owls (Psiloscops flammeolus). Four males departed from breeding areas in Colorado for fall migration between ≤5 and 21 October, arrived in wintering areas in Mexico between 11 October and 3 November, departed from wintering areas from ≤6 to 21 April, and returned to Colorado between 15 and 21 May. Core wintering areas for three males were located in the Trans‐Mexican Volcanic Belt Mountains in the states of Jalisco, Michoacán, and Puebla in central and east‐central Mexico, and the core area for the other male was in the Sierra Madre Oriental Mountains in Tamaulipas. The mean distance from breeding to wintering centroids was 2057 ± 128 km (SE). During fall migration, two males took a southeastern path to eastern Mexico, and two males took a path due south to central Mexico. In contrast, during spring migration, all four males traveled north from Mexico along the Sierra Madre Oriental Mountains to the Rio Grande Valley and north through New Mexico. The first stopovers in fall and last stopovers in spring were the longest in duration for all males and located 300–400 km from breeding areas. Final spring stopovers may have allowed male Flammulated Owls to fine tune the timing of their return to high‐elevation breeding areas where late snows are not uncommon. One male tracked in both years had similar migration routes, timing, and wintering areas each year. Core wintering and final stopover areas were located primarily in coniferous forests and woodlands, particularly pine‐oak forests, suggesting that these are important habitats for Flammulated Owls throughout their annual cycle.     

Wind conditions facilitate the seasonal water‐crossing behaviour of Oriental Honey‐buzzards Pernis ptilorhynchus over the East China Sea

Migratory raptors rarely fly over stretches of water larger than 25 km, although different species undertake water crossings of varying lengths, depending mainly on their wing morphology. Oriental Honey‐buzzards fly c. 680 km over the East China Sea in autumn from breeding areas in Japan to wintering areas in Southeast Asia, but avoid this long water crossing in spring. We investigated the effects of weather on this exceptional migratory behaviour and its seasonality through a maximum entropy niche modelling approach. We used data collected through satellite tracking of 31 adult birds as presence points and a set of variables related to wind, precipitation and convective condition as environmental predictors. Results of modelling showed very different, almost non‐overlapping, areas suitable for migration over the East China Sea region in autumn and spring. Suitable migration routes in autumn mostly occurred over the sea, whereas suitable areas for spring migration mostly occurred over land, suggesting that circumnavigating the East China Sea is preferable in spring. At the regional scale, wind conditions facilitate water‐crossing behaviour of Oriental Honey‐buzzards in autumn, but not in spring. Specifically, suitable tailwinds over the sea enable water‐crossing in autumn, whereas in spring, wind support and convective conditions are best over land. Our modelling did not suggest any importance of convective conditions for autumn migration. However, we expect that at smaller temporal scales, convective conditions would be a considerable facilitator of the water‐crossing behaviour in this species.     

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.     

Blackaps Sylvia atricapilla stopping over at the desert edge; physiological state and flight-range estimates

Migrating Blackcaps Sylvia atricapilla were mist netted at the desert edge in northern Israel and in Elat (southern Israel) during spring and autumn migrations between 1970 and 1991. Birds in spring in northern Israel were representative of birds that had completed the crossing of the Sahara, while those in Elat still had to cross the 150 km of the Negev Desert, which separates Elat and northern Israel. In autumn, birds captured in northern Israel were representative of those about to cross the Sahara Desert, while those in Elat had already started to cross the desert. The data allowed analysis of seasonal and location differences in the physiological state of Blackcaps before and after crossing the Sahara. Data analysed included body mass, visible fat score and calculated fat content. Autumn migrants were in better physiological condition than spring migrants at both locations, probably as a consequence of their migration route through fertile areas in autumn compared with the crossing of the Sahara in spring. Body mass was less variable after the Sahara crossing in spring than before the crossing in autumn. In spring, 71% and 67% of the birds were fat depleted (fat scores 0 and 1) at Elat and in northern Israel, respectively, while in autumn 34% and 42% were fat depleted. Blackcaps at Elat were 1.6 g lighter than those in northern Israel in autumn and 1.9 g lighter in spring. Potential flight ranges were estimated on the basis of meteorological conditions and flight altitude of passerines above the Negev in Israel (northern Sahara edge) during migration and on a simulation model that considered both energy and water as potential limiting factors for flight duration and distance. The simulation model predicted that half of the Blackcaps that stopped over in Elat and the majority of those that stopped over in northern Israel could not make a nonstop flight over the Sahara Desert in autumn without the assistance of at least an 8 m per s tailwind. Such a wind would still not be sufficient for 34% of the birds in Elat and 42% in northern Israel, and clearly they had insufficient fat reserves to cross the Sahara in a single flight. Although the fattest Blackcaps had accumulated sufficient fat to enable them to traverse the Sahara in a single flight, they probably faced dehydration by at least 12% of their initial body mass when they reached the southern Sahara edge. These birds should use intermittent migration with stopovers at sites with drinking and feeding potential. Their decision to stop over during the day in the desert at sites with shade but without food and water would be beneficial if the meteorological conditions during daytime migration imposed greater risks of dehydration than at night. Spring migrants could not reach their breeding areas in Europe without feeding, but those examined in Elat could cross the remainder of the desert in a single flight.     

Individual variation in migratory movements and winter behaviour of Iberian Lesser Kestrels Falco naumanni revealed by geolocators

The population decline of the Lesser Kestrel Falco naumanni has been the subject of studies across its Western Palaearctic breeding range, but little is known about its use of pre‐migratory areas or African wintering quarters. We used geolocators to describe the temporal and spatial patterns of Portuguese Lesser Kestrel migration and wintering behaviour. Data on the complete migration were obtained from four individuals and another three provided further information. Prior to southward migration, Lesser Kestrels showed two different behaviours: northward‐orientated movements to Spain and movements in the proximity of the breeding area. Autumn migration took place mostly in late September; spring departures occurred mainly in the first half of February. Wintering grounds included Senegal, Mauritania and Mali, with individuals overlapping considerably in Senegal. Movements registered within the wintering grounds suggest itinerant behaviour in relation to local flushes of prey. During spring migration, birds crossed the Sahara Desert through Mauritania, Western Sahara and Morocco before passing over the Mediterranean to reach Portugal. Autumn migration lasted 4.8 ± 1.1 days, and spring migration lasted 4.1 ± 0.3 days. The mean daily flight range varied between approximately 300 and 850 km for an entire journey of around 2500 km. Effective protection of roosting sites in both pre‐migratory and wintering areas and maintaining grasshopper populations in Sahelian wintering quarters appear crucial in preserving this threatened migratory raptor across its African–Eurasian flyway. There was no evidence of any deleterious effects of fitting birds with loggers.     

Satellite telemetry reveals long‐distance migration in the Asian great bustard Otis tarda dybowskii

The range of the great bustard stretches 10 000 km across Eurasia, one of the largest ranges of any threatened species. While movement patterns of the western subspecies of great bustard are relatively well‐understood, this is the first research to monitor the movements of the more endangered Asian subspecies of great bustard through telemetry and to link a breeding population of Asian great bustards to their wintering grounds. Using Argos/GPS platform transmitter terminals, we identified the annual movement patterns of three female great bustards captured at their breeding sites in northern Mongolia. The 4000 km round‐trip migration we have recorded terminated at wintering grounds in Shaanxi, China. This route is twice as long as has previously been reported for great bustards, which are among the heaviest flying birds. The journey was accomplished in approximately two months each way, at ground velocities of 48–98 km h^?1, and incorporated multiple and variable stopover sites. On their wintering grounds these birds moved itinerantly across relatively large home ranges. Our findings confirm that migratory behavior in this species varies longitudinally. This variation may be attributable to longitudinal gradients in seasonality and severity of winter across Eurasia. The distance and duration of the migratory route taken by great bustards breeding in Mongolia, the crossing of an international border, the incorporation of many stopovers, and the use of a large wintering territory present challenges to the conservation of the Asian subspecies of great bustard in this rapidly changing part of the world.     

Towards a new understanding of migration timing: slower spring than autumn migration in geese reflects different decision rules for stopover use and departure

According to migration theory and several empirical studies, long‐distance migrants are more time‐limited during spring migration and should therefore migrate faster in spring than in autumn. Competition for the best breeding sites is supposed to be the main driver, but timing of migration is often also influenced by environmental factors such as food availability and wind conditions. Using GPS tags, we tracked 65 greater white‐fronted geese Anser albifrons migrating between western Europe and the Russian Arctic during spring and autumn migration over six different years. Contrary to theory, our birds took considerably longer for spring migration (83 days) than autumn migration (42 days). This difference in duration was mainly determined by time spent at stopovers. Timing and space use during migration suggest that the birds were using different strategies in the two seasons: In spring they spread out in a wide front to acquire extra energy stores in many successive stopover sites (to fuel capital breeding), which is in accordance with previous results that white‐fronted geese follow the green wave of spring growth. In autumn they filled up their stores close to the breeding grounds and waited for supportive wind conditions to quickly move to their wintering grounds. Selection for supportive winds was stronger in autumn, when general wind conditions were less favourable than in spring, leading to similar flight speeds in the two seasons. In combination with less stopover time in autumn this led to faster autumn than spring migration. White‐fronted geese thus differ from theory that spring migration is faster than autumn migration. We expect our findings of different decision rules between the two migratory seasons to apply more generally, in particular in large birds in which capital breeding is common, and in birds that meet other environmental conditions along their migration route in autumn than in spring.     

Body composition of north and southbound migratory blackcaps is influenced by the lay‐of‐the‐land ahead

The annual migration of small birds depends on the optimal management of time and energy. Since refueling at stopovers between flights consumes most of the birds’ time and energy, selection of food‐rich sites, and timely departure therefrom are likely crucial to success. We examined this concept quantifying body composition of 200 migrating blackcaps, Sylvia atricapilla, in Eilat, Israel, using dual‐energy x‐ray absorptiometry and generated a model to predict body composition as it changes with body mass (m_b). We then back‐calculated body composition of > 20 000 blackcaps ringed between 1984 and 2005, and tested the hypothesis that the amount of fuel that a bird stores determines the length of its stopover. We predicted that 1) if time‐constrained in spring, birds at the stopover site carry less than a maximum fuel load, but 2) if not time‐constrained, as in autumn, their fuel load is much higher than in spring. We found the change in body composition of blackcaps to be biphasic and correlated with increasing m_b. At m_b < ? 17.8 g, increasing m_b is due to increasing lean mass (m_l), while at m_b > ? 17.8 g increasing m_b results from increasing fat mass (m_f), which is accompanied by decreasing m_l. Body composition of blackcaps at a spring stopover site indicates that blackcaps leave stopovers as soon as they regain functionality of their digestive systems, but before laying down much m_f. In autumn blackcaps arrive with fuel stores much larger than in spring. For these birds, the Eilat stopover apparently serves to complete fat accumulation before crossing the deserts ahead. We conclude that in spring, the decision to depart is not determined by the bird's fuel stores, especially when early arrival at the breeding site, and therefore time, is of the essence. In autumn, accumulating enough fuel to ensure successful crossing of the deserts ahead probably dictates stopover time.     

Individual and sex‐related patterns of prolonged flights during both day and night by great reed warblers crossing the Mediterranean Sea and Sahara Desert

A wide variety of the barrier crossing strategies exist among migrating songbirds, ranging from strict nocturnal flights to non‐stop flights over a few days. We evaluate barrier crossing strategies in a nocturnally migrating songbird crossing the Mediterranean Sea and the Sahara Desert, the great reed warbler, exploring variation between the sexes and within individuals. We used data from 31 year‐round light‐level geolocators tracks from 26 individuals (13 males and 13 females), with four individuals tracked for 2–3.5 consecutive years. Almost all individuals (25 of 26) prolonged their flights into the day at least on one occasion. The mean duration of these prolonged flights was 19.9 h and did not differ between sexes or seasons. Fifteen birds performed non‐stop flights during more than one full day and night (≥ 24 h; mean = 31.9 h; max = 55 h) in autumn and/or spring, but these flights were generally too short to cross an entire barrier (such as the Sahara Desert) in one non‐stop flight. Patterns of prolonged flights showed considerable within‐individual variation in females between seasons (autumn versus spring) and in both males and females between years, suggesting high individual flexibility in migration strategy. Significantly more males than females performed prolonged flights during autumn migration, but not spring, possibly reflecting sex‐specific carry‐over effects. We conclude that great reed warblers have the ability to conduct prolonged continuous flights for up to several nights and days, which potentially would allow them to cross the Sahara Desert in one non‐stop flight. However, they typically use a mixed strategy of several nocturnal flights with intermittent stopovers in combination with 1–3 prolonged flights. Prolonged flights covered less than half (44%) of the total flight time across the barriers, and the diurnal parts of the flights covered only 18% of this time.     

Carry‐over effects and compensation: late arrival on non‐breeding grounds affects wing moult but not plumage or schedules of departing bar‐tailed godwits Limosa lapponica baueri

In the annual cycle of migratory birds, temporal and energetic constraints can lead to carry‐over effects, in which performance in one life history stage affects later stages. Bar‐tailed godwits Limosa lapponica baueri, which achieve remarkably high pre‐migratory fuel loads, undertake the longest non‐stop migratory flights yet recorded, and breed during brief high‐latitude summers, may be particularly vulnerable to persistent effects of disruptions to their rigidly‐timed annual routines. Using three years of non‐breeding data in New Zealand, we asked how arrival timing after a non‐stop flight from Alaska (>11 000 km) affected an individual godwit's performance in subsequent flight feather moult, contour feather moults, and migratory departure. Late arrival led to later wing moult, but godwits partially compensated for delayed moult initiation by increasing moult rate and decreasing the total duration of moult. Delays in arrival and wing moult up to 34–37 d had no apparent effect on an individual's migratory departure or extent of breeding plumage at departure, both of which were extraordinarily consistent between years. Thus, ‘errors’ in timing early in the non‐breeding season were essentially corrected in New Zealand prior to spring migration. Variation in migration timing also had no apparent effect on an individual's likelihood of returning the following season. The bar‐tailed godwits’ rigid maintenance of plumage and spring migration schedules, coupled with high annual survival, imply a surprising degree of flexibility to address unforeseen circumstances in the annual cycle.     

Timing of migration and residence areas during the non‐breeding period of barn swallows Hirundo rustica in relation to sex and population

We investigated sex‐ and year‐dependent variation in the temporal and spatial movement pattern of barn swallows Hirundo rustica during the non‐breeding period. Hundred and three individuals equipped with miniaturized light‐level geolocators at three different breeding areas in southern Switzerland and northern Italy provided data for the analysis. We identified a region 1000 km in radius centred in Cameroon as the main non‐breeding residence area of these three geographical populations. Five residence areas of males only were in southern Africa, south of 19°S. Most individuals occupied a single site during their stay south of the Sahara. The timing of migration broadly overlapped between sexes and all geographical breeding populations. Between the two study years there was a distinct difference of 5 to 10 d in departure dates from and arrival at the breeding sites. Remarkably, the period of residence in sub‐Saharan Africa was very similar (157 d) in the two study years, but their positions in the first year (2010–2011) were about 400 km more to the north than in the second (2011–2012). Independent of the year, individuals with sub‐Saharan residence areas further north and east had a shorter pre‐breeding migration and arrived earlier than those staying further south and west. In addition, birds breeding in southern Switzerland arrived at their breeding colony 7–10 d later than those breeding only 100 km south, in the Po river plain. Our study provides new information on the variance in migration phenology and the distribution of residence areas in sub‐Saharan Africa in relation to sex, population and year. It supports the usefulness of light‐level geolocators for the study of annual routines of large samples of small birds.     

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.