A place to land: spatiotemporal drivers of stopover habitat use by migrating birds

Migrating birds require en route habitats to rest and refuel. Yet, habitat use has never been integrated with passage to understand the factors that determine where and when birds stopover during spring and autumn migration. Here, we introduce the stopover‐to‐passage ratio (SPR), the percentage of passage migrants that stop in an area, and use 8 years of data from 12 weather surveillance radars to estimate over 50% SPR during spring and autumn through the Gulf of Mexico and Atlantic coasts of the south‐eastern US, the most prominent corridor for North America’s migratory birds. During stopovers, birds concentrated close to the coast during spring and inland in forested landscapes during autumn, suggesting seasonal differences in habitat function and highlighting the vital role of stopover habitats in sustaining migratory communities. Beyond advancing understanding of migration ecology, SPR will facilitate conservation through identification of sites that are disproportionally selected for stopover by migrating birds.     

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.     

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.     

Faster spring migration in northern wheatears is not explained by an endogenous seasonal difference in refueling rates

A widespread phenomenon in migrant birds is that they travel faster in spring than in autumn. During migration birds spend most time at stopover sites and, correspondingly, the faster spring migration is mainly explained by shorter stopovers in spring than autumn. Because a main purpose of stopovers is to replenish the fuel used in flight, a higher rate of fuel deposition (FDR) in spring is thought to explain the shorter stopovers and hence shorter total duration of migration in spring. Critical migratory processes, including the onset and extent of pre‐migratory fueling, are endogenously regulated. It is therefore not unlikely that refueling at stopover sites is, at least partly, also under endogenous control. We here tested whether there is an endogenous seasonal difference in food intake and FDR, which could contribute to shorter stopovers and hence faster migration in spring. We measured daily food intake and daily FDR in two subspecies of the northern wheatear Oenanthe oenanthe, temporarily confined at stopover under identical constant indoor conditions in spring and autumn. The two wheatear subspecies differed markedly in absolute food intake and FDR. Within subspecies, however, food intake and FDR did not differ between spring and autumn, indicating that faster spring migration in northern wheatears is not explained by an endogenously controlled seasonal difference in birds’ motivation to refuel. To further substantiate this claim, similar measurements should be taken at other locations along northern wheatears’ migration routes. Comparable experiments in other species could test the generality of our results.     

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.     

Effects of migration distance and sex on stopover timing and refueling by Wilson's Warblers

Migration distance and sex are integral to avian migration strategies, yet these intrinsic factors are understudied with respect to their effects on stopover ecology and behavior. We investigated how individual variation in migration distance and sex affected body condition, refueling performance, and stopover durations of Wilson's Warblers (Cardellina pusilla) during spring and fall 2006 at a stopover site in southern Arizona. Migration distance of individuals was inferred using the hydrogen stable isotope ratio of feathers (δ²H_f) as an index of breeding latitude, refueling rate was assessed using plasma triglyceride and β‐OH‐butyrate, and minimum length of stay was estimated by re‐sighting color‐banded birds. In the spring, migration distance and sex were strong determinants of the timing of migration by Wilson's Warblers, with males and shorter‐distance migrants passing through the site earlier than females and longer‐distance migrants. Later‐arriving migrants also had higher fat scores and refueling rates. However, neither migration distance nor sex independently affected body condition (fat score or size‐corrected mass), refueling rate, or minimum length of stay. In a smaller sample of fall migrants, we found that longer‐distance migrants and males had higher refueling rates than shorter‐distance migrants and females. Our results show that differences in migration distance can, under some circumstances, affect how birds use stopover sites. Stable isotope analysis and other methods can provide information on migration distance, and should be integrated with measurements of timing, fuel stores, refueling performance, and departure behavior to gain a deeper understanding of bird migration.     

Fuel reserves affect migratory orientation of thrushes and sparrows both before and after crossing an ecological barrier near their breeding grounds

Fat reserves influence the orientation of migrating songbirds at ecological barriers, such as expansive water crossings. Upon encountering a body of water, fat migrants usually cross the barrier exhibiting 'forward' migration in a seasonally appropriate direction. In contrast, lean birds often exhibit temporary 'reverse' orientation away from the water, possibly to lead them to suitable habitats for refueling. Most examples of reverse orientation are restricted to autumn migration and, in North America, are largely limited to transcontinental migrants prior to crossing the Gulf of Mexico. Little is known about the orientation of lean birds after crossing an ecological barrier or on the way to their breeding grounds. We examined the effect of fat stores on migratory orientation of both long- and short-distance migrants before and after a water crossing near their breeding grounds; Catharus thrushes (Swainson's and gray-cheeked thrushes, C. ustulatus and C. minimus ) and white-throated sparrows Zonotrichia albicollis were tested for orientation at the south shore of Lake Ontario during spring and autumn. During both spring and autumn, fat birds oriented in a seasonally appropriate, forward direction. Lean thrushes showed a tendency for reverse orientation upon encountering water in the spring and axial, shoreline orientation after crossing water in the autumn. Lean sparrows were not consistently oriented in any direction during either season. The responses of lean birds may be attributable to their stopover ecology and seasonally-dependent habitat quality.     

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.     

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.     

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.     

Spring migration dynamics and sex-specific patterns in stopover strategy in the Wood Sandpiper Tringa glareola

Due to being a virtually monomorphic wader species, migration dynamics and sex-related migration patterns in the Wood Sandpiper (Tringa glareola) have rarely been investigated. We captured spring migrants at an important stopover site in northeastern Austria. Birds were individually color-marked, and sex was determined by an analysis of DNA from tail feather material. Among temporary residents (birds seen again after day of capture), males migrated on average 3 days earlier than females. However, since sexes did not differ in fat score, the length of stay and the proportion of transients (birds not seen again after day of capture) and temporary residents, we suggest that males and females adopt similar migration strategies in the spring. The large number of transients captured as well as shorter stopover durations in later temporary residents indicate that Wood Sandpipers minimize time at this stage of their northbound migration. Temporary residents earlier in the season exhibited lower fat stores than later ones. Nevertheless, since the fat stores of transients and temporary residents were similar even after the progress of the season had been accounted for, we assume that Wood Sandpipers may afford to exhibit individual flexibility in migration strategy and the use of stopover sites, especially early in the season. This variability may be a necessary adaptation to cope with possible varying environmental conditions at dynamic and unpredictable inland stopover sites. After having reached North Mediterranean regions, mean body mass of spring migrants gradually increases during successive stopovers, indicating that Wood Sandpipers follow a ‘hopping’ migration strategy. This emphasizes the high conservation value of even small artificial mudflat pools as important stepping stones in order to maintain a continuous network of wetland habitats for this continental migrant.     

Environmental conditions at arrival to the wintering grounds and during spring migration affect population dynamics of barn swallows Hirundo rustica breeding in Northern Italy

Several populations of long-distance migratory birds are currently suffering steep demographic declines. The identification of the causes of such declines is difficult because population changes may be driven by events occurring in distant geographical areas during different phases of the annual life-cycle of migrants. Furthermore, wintering areas and migration routes of populations of small-sized species are still largely unknown, with few exceptions. In this paper we identified the critical phases of the annual life-cycle that most influence the population dynamics of a small passerine, the Barn Swallow Hirundo rustica. We used information on temporal dynamics of a population breeding in Northern Italy, whose wintering range and timing of migration have been recently described by miniaturised tracking dataloggers. Our results indicated that primary productivity in the wintering grounds in the month when most individuals arrive from autumn migration and primary productivity in an area that is probably a stopover site during spring migration, influenced population dynamics more than habitat conditions at the breeding grounds. By using annual variation in primary productivity at the wintering grounds and stopover sites as predictors, we replicated the observed interannual population changes with great accuracy. However, the steep decline recently suffered by the population could be replicated only by including a constant annual decline in the model, suggesting that changes in primary productivity only predicted the interannual variation around the long-term trend. Our study therefore suggests the existence of critical periods during wintering and migration that may have large impact on population fluctuations of migrant birds.     

迁徙鸟类中途停歇期的生理生态学研究

大多数候鸟的迁徙活动由迁徙飞行和中途停歇两个部分组成。在迁徙过程中,鸟类要多次交替经历消耗能量的飞行阶段和积累能量的中途停歇阶段。从鸟类在中途停歇时期的能量积累速度、体重变化模式以及迁徙飞行中的禁食或食物限制、食物种类的改变、中途停歇的能量快速积累过程对消化器官的影响等方面,对目前迁徙鸟类的生理生态学研究成果进行回顾,并提出有待解决的问题及今后的研究方向。     

Faster migration in autumn than in spring: seasonal migration patterns and non‐breeding distribution of Icelandic whimbrels Numenius phaeopus islandicus

Migration is fundamental in the life of many birds and entails significant energetic and time investments. Given the importance of arrival time in the breeding area and the relatively short period available to reproduce (particularly at high latitudes), it is expected that birds reduce spring migration duration to a greater extent than autumn migration, assuming that pressure to arrive into the wintering area might be relaxed. This has previously been shown for several avian groups, but recent evidence from four tracked Icelandic whimbrels Numenius phaeopus islandicus, a long distance migratory wader, suggests that this subspecies tends to migrate faster in autumn than in spring. Here, we 1) investigate differences in seasonal migration duration, migration speed and ground speed of whimbrels using 56 migrations from 19 individuals tracked with geolocators and 2) map the migration routes, wintering and stopover areas for this population. Tracking methods only provide temporal information on the migration period between departure and arrival. However, migration starts with the fuelling that takes place ahead of departure. Here we estimate the period of first fuelling using published fuel deposition rates and thus explore migration speed using tracking data. We found that migration duration was shorter in autumn than in spring. Migration speed was higher in autumn, with all individuals undertaking a direct flight to the wintering areas, while in spring most made a stopover. Wind patterns could drive whimbrels to stop in spring, but be more favourable during autumn migration and allow a direct flight. Additionally, the stopover might allow the appraisal of weather conditions closer to the breeding areas and/or improve body condition in order to arrive at the breeding sites with reserves.     

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 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.     

红胁蓝尾鸲(Tarsiger cyanurus)在中国东北部帽儿山地区的迁徙中途停歇生态

中国迁徙鸣禽类的保护面对着与世界其他地区如欧洲和北美洲鸟类保护相似的挑战。迁徙鸣禽类具有复杂生活周期和很大的空间关联。迁徙过程中发生的事件对迁徙鸣禽类种群动态具有决定作用。对于鸣禽类迁徙中途停歇期的生态,比如停歇期的长短,能量的积累,生境的利用等,了解还非常有限。在中国东北部的一个鸟类迁徙停歇地对红胁蓝尾鸲(Tarsiger cyanurus)的中途停歇生态包括迁徙时间、停歇时间、能量状态和性比进行了研究。2002年秋和2003年春分别捕获了1751只和684只红胁蓝尾鸲。红胁蓝尾鸲的体重在秋季迁徙时要比在春季迁徙时重。春季雌性红胁蓝尾鸲停歇时的能量状态指数最低; 而秋季的红胁蓝尾鸲比春季的红胁蓝尾鸲停歇时间更长。无论季节和性别,红胁蓝尾鸲的能量状态指数和第1次捕获的时间早晚成正相关, 间接证明红胁蓝尾鸲在停歇期间能够比较快地积累能量。秋季雄性红胁蓝尾鸲日体重净增率最大。估测秋季停歇期的每日能量净增能维持红胁蓝尾鸲雌性0.6h和雄性3.1h的飞行。红胁蓝尾鸲的中途停歇生态与北美和欧洲一些迁徙鸣禽类很相似。比如,春季迁徙过境的时间和脂肪积累的变化与自然选择对雄性的要求：当食物和气候适宜时尽快到达繁殖地的假设是一致的。对迁徙中途的停歇生态研究有利于更好地了解鸟类的迁徙行为和更有效地保护迁徙鸣禽类。     

Comparison of autumn and spring migration strategies of Neotropical migratory landbirds in northeast Belize

ABSTRACT Migration represents one of the most vulnerable stages of a migrant's life cycle, but the strategies and stopover sites used by Neotropical migrants in Central America are not well known. We carried out constant‐effort mist netting and conducted censuses along transects during one autumn (2007) and one spring (2008) migration in northeast Belize. We recorded more landbird migrant species in autumn (63) than in spring (54), and spring abundance was >25% lower for 88% of transient species. These differences in presence and abundance indicate that routes and stopover strategies vary between seasons and species. In autumn, fuel loads, calculated as any increase in mass above lean body mass (LBM), were generally small (mean = 5.9% LBM and 10.1% LBM for wintering and transient species, respectively) and fuel deposition rates and minimum stopover durations suggest that some individuals replenished energy reserves in our study area. Variation in autumn fuel loads meant that some individuals had reserves sufficient for flights >1000 km. Fuel loads were larger in spring for 16 of 17 species, and the mean spring fuel load for transient species (32.5% LBM) was sufficient for a flight from northeast Belize to North America without refueling. The similarity in spring passage times between northeast Belize and the Gulf Coast of the United States also suggests that energy reserves were not replenished in northeast Belize prior to crossing the Gulf of Mexico. We hypothesize that sufficient energy reserves are accumulated during spring stopovers in northern South America or elsewhere in Mesoamerica to allow migrants to fly directly to North America without refueling.     

Repeatability of individual migration routes,wintering sites,and timing in a long‐distance migrant bird

Migratory birds are often faithful to wintering (nonbreeding) sites, and also migration timing is usually remarkably consistent, that is, highly repeatable. Spatiotemporal repeatability can be of advantage for multiple reasons, including familiarity with local resources and predators as well as avoiding the costs of finding a new place, for example, nesting grounds. However, when the environment is variable in space and time, variable site selection and timing might be more rewarding. To date, studies on spatial and temporal repeatability in short‐lived long‐distance migrants are scarce, most notably of first‐time and subsequent migrations. Here, we investigated repeatability in autumn migration directions, wintering sites, and annual migration timing in Hoopoes (Upupa epops), a long‐distance migrant, using repeated tracks of adult and first‐time migrants. Even though autumn migration directions were mostly the same, individual wintering sites often changed from year to year with distances between wintering sites exceeding 1,000 km. The timing of migration was repeatable within an individual during autumn, but not during spring migration. We suggest that Hoopoes respond to variable environmental conditions such as north–south shifts in rainfall during winter and differing onset of the food availability during spring migration.     

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.