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 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.     

Migrating songbirds on stopover prepare for,and recover from,oxidative challenges posed by long‐distance flight

Managing oxidative stress is an important physiological function for all aerobic organisms, particularly during periods of prolonged high metabolic activity, such as long‐distance migration across ecological barriers. However, no previous study has investigated the oxidative status of birds at different stages of migration and whether that oxidative status depends on the condition of the birds. In this study, we compared (1) energy stores and circulating oxidative status measures in (a) two species of Neotropical migrants with differing migration strategies that were sampled at an autumn stopover site before an ecological barrier; and (b) a species of trans‐Saharan migrant sampled at a spring stopover site after crossing an ecological barrier; and (2) circulating oxidative measures and indicators of fat metabolism in a trans‐Saharan migrant after stopovers of varying duration (0–8 nights), based on recapture records. We found fat stores to be positively correlated with circulating antioxidant capacity in Blackpoll Warblers and Red‐eyed Vireos preparing for fall migration on Block Island, USA, but uncorrelated in Garden Warblers on the island of Ponza, Italy, after a spring crossing of the Sahara Desert and Mediterranean Sea. In all circumstances, fat stores were positively correlated with circulating lipid oxidation levels. Among Garden Warblers on the island of Ponza, fat anabolism increased with stopover duration while oxidative damage levels decreased. Our study provides evidence that birds build antioxidant capacity as they build fat stores at stopover sites before long flights, but does not support the idea that antioxidant stores remain elevated in birds with high fuel levels after an ecological barrier. Our results further suggest that lipid oxidation may be an inescapable hazard of using fats as the primary fuel for flight. Yet, we also show that birds on stopover are capable of recovering from the oxidative damage they have accrued during migration, as lipid oxidation levels decrease with time on stopover. Thus, the physiological strategy of migrating songbirds may be to build prophylactic antioxidant capacity in concert with fuel stores at stopover sites before a long‐distance flight, and then repair oxidative damage while refueling at stopover sites after long‐distance flight.     

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.     

Inability to regain normal body mass despite extensive refuelling in great reed warblers following the trans‐Sahara crossing during spring migration

Migratory birds wintering in Africa face the challenge of passing the Sahara desert with few opportunities to forage. During spring migration birds thus arrive in the Mediterranean area with very low energy reserves after crossing the desert. Since early arrival to the breeding grounds often is of importance to maximize reproductive success, finding stopover sites with good refuelling possibilities after the Saharan passage is of utmost importance. Here we report on extensive fuelling in the great reed warbler Acrocephalus arundinaceus on the south coast of Crete in spring, the first land that they encounter after crossing the Sahara desert and the Mediterranean Sea in this area. Birds were studied at a river mouth and due to an exceptional high recapture rate (45 and 51% in two successive years), we were able to get information about stopover behaviour in 56 individual great reed warblers during two spring seasons. The large proportion of trapped great reed warbler compared to other species and the large number of recaptures suggest that great reed warblers actively choose this area for stopover. They stayed on average 3–4 d, increased on average about 3.5 g in body mass and the average rate of body mass increase was 4.8% of lean body mass d^–1. Wing length affected the rate of increase and indicated that females have a slower increase than males. The results found show that great reed warblers at this site regularly deposit larger fuel loads than needed for one continued flight stage. The low body mass found in great reed warblers (also in birds with high fat scores) is a strong indication that birds staging at Anapodaris still had not been able to rebuild their structural tissue after the strenuous Sahara crossing, suggesting that rebuilding structural tissue may take longer time than previously thought.     

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.     

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.     

When and where to fuel before crossing the Sahara desert – extended stopover and migratory fuelling in first-year garden warblers Sylvia borin

Large numbers of passerine migrants cross the Sahara desert every year on their way to-and-from wintering areas in tropical Africa. In the desert, hardly any fuelling opportunities exist and most migrants have to prepare in advance. A central question is how inexperienced birds know where to fuel. Inexperienced garden warblers Sylvia borin were studied in Greece just before the desert crossing in autumn. Body mass data collected at two sites indicate that most birds do not fuel for the desert crossing further north. For the first time, detailed information about stopover duration close to the Sahara desert was studied by using light weight radio-transmitters. Results from Crete show that most first-year garden warblers arrive with relatively small fuel loads in relation to lean body mass (<30%), stay for 13–20 d and depart with an average fuel load of about 100%. Radio-tagged birds performed small scale movements initially and took advantage of fig fruits. Birds trapped at fig trees were heavier than birds trapped with tape lures, showing that tape lures can bias the sample of migrants trapped. The precise fuelling pattern found indicates that first-year migrants must also include external spatial cues to make the preparation for crossing the desert in the right area.     

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.     

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.     

Migration strategies of the Baltic dunlin: rapid jump migration in the autumn but slower skipping type spring migration

Migration during spring is usually faster than during autumn because of competition for breeding territories. In some cases, however, the costs and benefits associated with the environment can lead to slower spring migration, but examples are quite rare. We compared seasonal migration strategies of the endangered Baltic population of the dunlin Calidris alpina schinzii using light‐level geolocator data from 26 individuals breeding in Finland. Autumn migration was faster, with individuals showing a ‘jump’ and ‘skipping’ migration strategy characterised by fewer stationary periods, shorter total stopping time and faster flight. Spring migration was slower, with individuals using a ‘skipping’ strategy. The duration of migration was longer for early departing birds during spring but not during autumn suggesting that early spring migrants are prevented from arriving to the breeding areas or that fueling conditions are worse on the stopover sites for early arriving individuals. Dunlins showed high migratory connectivity. All individuals had one long staging at the Wadden Sea in the autumn after which half of the individuals flew 4500 km non‐stop to Banc d’Arguin, Mauritania. The other half stopped briefly on the Atlantic coast on their way to Mauritania. One bird wintered on the coast of Portugal. Nine individuals that carried geolocators for two years were site faithful to their final non‐breeding sites. Based on the strategies during the non‐breeding period we identified, Baltic dunlin may be especially vulnerable to rapid environmental changes at the staging and non‐breeding areas. Consequently, the preservation of the identified non‐breeding areas is important for their conservation.     

Strategies of passerine migration across the Mediterranean Sea and the Sahara Desert: a radar study

Radar observations of the diurnal timing of bird migration in the Sahara Desert are presented for autumn migration. Study sites were on a transect along the north-south migratory direction. Three groups of birds migrating either during day, evening or night in the northern part of the Western desert in Egypt were identified. The maximum of day and night groups occurred later the further south the study sites were. Based on the distance between sites and the timing of peak migration, birds were flying at an estimated ground speed of about 20 m/s. The maximum of the evening group was at about 21:00 h at all sites. The three groups were classified according to three different strategies of migration across the Mediterranean Sea and the Sahara Desert: (1) the day group of birds performed a non-stop flight across the sea and at least the northern part of the desert; [2] the night group performed an intermittent migratory strategy with stopover at the coast of Egypt to continue migration the next evening; (3) the evening group birds were also intermittent migratory fliers, but they stopped somewhere in the desert after a continuous flight across the sea and part of the desert. About 20% of all migrants are involved in non-stop migration and 80% in intermittent migration with stopover at the coast (70%) or with stopover in the desert (10%). It is argued that any species of small passerine has the option to use any of the three strategies.     

Migration strategies and annual space‐use in an Afro‐Palaearctic aerial insectivore – the European nightjar Caprimulgus europaeus

Obligate insectivorous birds breeding in high latitudes travel thousands of kilometres during annual movements to track the local seasonal peaks of food abundance in a continuously fluctuating resource landscape. Avian migrants use an array of strategies when conducting these movements depending on e.g. morphology, life history traits and environmental factors encountered en route. Here we used geolocators to derive data on the annual space‐use, temporal pattern and migratory strategies in an Afro‐Palaearctic aerial insectivorous bird species – the European nightjar Caprimulgus europaeus. More specifically, we aimed to test a set of hypothesises pertaining to the migration of a population of nightjars breeding in south‐eastern Sweden. We found that the birds wintered across the central and western parts of the southern tropical Africa almost entirely outside the currently described wintering range of the species. The nightjars performed a narrow loop migration across Sahara, with spring Sahel stopovers significantly to the west of autumn stops indicative to an adaptive response to winds during migration. To our surprise, the migration speed was faster in the autumn (119 km d^{? 1}) than in the spring (99 km d^{? 1}), possibly due to the prevailing wind regimes over the Sahara. The estimated flight fraction in both autumn (14%) and spring (12%) was almost exactly as the theoretically predicted 1:7 time relationship between flights and stopovers for small birds. The temporal patterns within the annual cycle indicate that individuals follow alternative spatiotemporal schedules that converge towards the breeding season. The positive relationship between the spatially and temporally distant winter departure and breeding arrival suggests that individuals´ temporal fine‐tuning to breeding may be constrained, leading to potential negative fitness consequences.     

Stopover duration of Palearctic passerine migrants in the western Sahara – independent of fat stores?

Birds on migration spend much more time on stopover sites to refuel for the next migration step than aloft, but empirical data on stopover duration are rare, especially for Palearctic trans-Sahara migrants whilst crossing the desert. Previous studies suggest that stopover duration of fat birds in oases is much shorter than that of lean birds. During 2003 and 2004 capture–recapture data of migrating passerines from two inland oases in spring and from one coastal site in autumn in Mauritania, West Africa, were analysed to test whether the probability of being a transient and the stopover duration depend on fuel stores at first capture. The application of capture–recapture models revealed that during autumn migration at the coast the proportion of transients (individuals that stop over only for 1 day) was relatively high (77–90%) in three out of four species investigated and stopover duration was short (1.9–4.6 days). In the inland oases in spring, transients were detected in only four out of 12 analyses. Stopover duration was longer than at the coast in autumn and surprisingly long in some species with durations of up to 30 days. Models taking into account the initial fat load of birds on the first capture occasion were, with one exception, never the most parsimonious ones. This indicates that the time spent after and before capture at the stopover site did not depend on the fat stores at first capture. Therefore, we cannot confirm the assumption that birds arriving at stopover sites in the desert with low fat loads stay longer than birds that arrive with high fat loads.     

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.     

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.     

How does a first year passerine migrant find its way? Simulating migration mechanisms and behavioural adaptations

In the Palaearctic-African migration system, birds face several trade-offs on their first autumn migration. The shortest route, minimising travelling time, would lead them directly south across the Mediterranean Sea and the Sahara Desert, involving long stretches of no refuelling possibilities. This route is risky because of flight range constraints. Besides taking longer, a detour along the Iberian peninsula may require a more complex orientation mechanism. We simulated migrants with stopover and orientation behaviour and investigated the effect of flight costs and behavioural rules (e.g. crossing or flying along coast lines, a shift in migratory direction) on the resulting flight path and especially on the evolution of endogenous directions. The simulation of autumn migration from southern Scandinavia to south of the Sahara showed that it would be possible to reach the winter quarters by vector summation with a constant endogenous direction, but then either orientation must be very accurate or flight costs must be small. For small passerines both requirements are so far not corroborated by empirical studies. Alternatively, flying along coast lines or shifting direction in northern Africa from south-westerly to southerly, resulted in similar survival rates as with a constant south-westerly endogenous direction, but with a larger range of feasible values. Although weather factors were not included, our results suggest that the Mediterranean Sea and the Sahara desert had a dominating influence on the evolution of endogenous directions. This influence is probably acting through flight range constraints.     

Wide ranging stopover movements and substantial fuelling in first year garden warblers at a northern stopover site

Migratory birds use stopovers to replenish their fuel reserves and they generally spend more time at stopover sites than they do in actual flight. When arriving at a new stopover site birds may need to search extensively to find a suitable feeding area and this search and settling period may affect the duration of stopover. Stopover behaviour can thus have profound effects on the migratory programme and studies on stopover behaviour are important to understand migratory strategies. We followed 51 first‐year garden warblers Sylvia borin with radio‐transmitters at an autumn stopover site on the island of Gotland in southern Sweden. Our aim was to determine the distance birds relocated from the coastal capture site when searching for an area to settle in, and also to establish the duration of stopover and put it in relation to refuelling rate by recapturing a subset of the radio‐tracked individuals. Sixteen birds made an extended stopover (> 2 d), relocated inland from the capture site and settled on average 5.6 km from the capture site, with the longest recorded relocation being fourteen kilometres. Birds that relocated nocturnally settled in areas further away than birds that relocated diurnally. Thirteen birds that continued migration after a short stop carried larger fuel stores than birds that stopped over longer and they remained close to the capture site until departure. Three birds were re‐trapped and showed high fuelling rates, between 0.3 and 1.1 g d^–1. They left the stopover site with fuel loads between 40–56 percent of lean body mass, which possibly would have allowed them to reach the Mediterranean area without additional refuelling stops.     

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.