Spring passerine migrants stopping over in the Sahara are not fall-outs

The strategy of migrants crossing the Sahara desert has been the subject of debate, but recent evidence from radar studies has confirmed that most passerines use an intermittent migration strategy. The latter has also been suggested from previous studies in oases during autumn migration. It was found that migrants with relatively high fuel loads rest in the desert during daytime and continue migration during the following night, whereas lean migrants stopover in oases for several days to refuel. However, data from the Sahara are scarce for spring migration. We captured passerine migrants near B?r Amrane (22°47′N, 8°43′W) in the plain desert of Mauritania for 3 weeks during spring migration in 2004. We estimated flight ranges of 85 passerines stopping over in the desert to test whether they carried sufficient fuel loads to accomplish migration across the Sahara successfully. High fat loads of the majority of birds indicated that they were neither “fall-outs” nor too weak to accomplish migration successfully. The flight range estimates, based on mean flight speeds derived from radar measurements (59 km/h), revealed that 85% of all birds were able to reach the northern fringe of the desert with an intermittent migration strategy. Furthermore, birds stopping over in an oasis (Ouadane, 370 km to the southwest of B?r Amrane) did not carry consistently lower fuel loads compared to the migrants captured in the desert.     

How do migratory songbirds cross the Sahara?

Many European songbirds winter in Africa south of the Sahara.Along their migratory routes they must fly over the huge desert belt of the Sahara twice a year. For decades ornithologists have assumed that most migrants cross this 'ecological barrier' in one long non-stop flight of thousands of kilometres. Results of recent research, however, suggest that many of the songbirds that migrate across the Sahara follow an intermittent migratory strategy with regular stopovers in the desert.     

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.     

Seasonal differences in energy requirements of Garden Warblers Sylvia borin migrating across the Sahara desert

The Sahara desert acts as an ecological barrier for billions of passerine birds on their way to and from their African wintering areas. The Garden Warbler Sylvia borin is one of the most common migrants involved. We used body mass of this species from Greece in autumn and spring to simulate the desert crossing and to assess how body mass relates to fuel requirement. The flight range estimates were adjusted to the seasonal extent of the desert, 2200 km in autumn and about 2800 km in spring. In autumn, with an average fuel load of about 100% of body mass without fuel, birds were not able to cross the desert in still air, but northerly winds prevail during September and with the average wind assistance only one in 14 was predicted to fail to make the crossing. Body mass data from spring, after the desert crossing, was used to estimate departure body mass from south of the desert. The average wind assistance in spring is close to zero and departure body mass of the average bird arriving at Antikythira, a small Greek island, under such conditions was estimated to be 34.6 g, which corresponded to a fuel load of 116%. Calculations based on 1% body mass loss per hour of flight showed slightly larger body mass loss than that calculated from flight range estimates. The results suggest that passerine birds about to cross the eastern part of the Sahara desert need to attain a larger fuel load in spring than in autumn.     

Flight altitude of trans-Sahara migrants in autumn: a comparison of radar observations with predictions from meteorological conditions and water and energy balance models

Radar observations on the altitude of bird migration and altitudinal profiles of meteorological conditions over the Sahara desert are presented for the autumn migratory period. Migratory birds fly at an average altitude of 1016 m (a.s.l.) during the day and 571 m during the night. Weather data served to calculate flight range using two models: an energy model (EM) and an energy-and-water model (EWM). The EM assumes that fuel supply limits flight range whereas the EWM assumes that both fuel and water may limit flight range. Flight ranges estimated with the EM were generally longer than those with the EWM. This indicates that trans-Sahara migrants might have more problems balancing their water than their energy budget. However, if we assume fuel stores to consist of 70% instead of 100% fat (the remainder consisting of 9% protein and 21% water), predicted flight ranges of the EM and EWM largely overlap. Increased oxygen extraction, reduced flight costs, reduced exhaled air temperature, reduced cutaneous water loss and increased tolerance to water loss are potential physiological adaptations that would improve the water budget in migrants. Both the EM and EWM predict optimal flight altitudes in agreement with radar observations in autumn. Optimal flight altitudes are differently predicted by the EM and EWM for nocturnal spring migration. During spring, the EWM predicts moderately higher and the EM substantially higher flight altitudes than during autumn. EWM predictions are therefore in better agreement with radar observations on flight altitude of migrants over the Negev desert in spring than EM predictions.     

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.     

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.     

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.     

SPRING MIGRATION THROUGH SOUTHEAST MOROCCO

Very few quantitative data existed for the spring migration of birds across the Sahara desert. Observations covering 105 days in spring 1963–66 were made at Defilia, on the extreme northern fringe of the desert in southeast Morocco. The physical features and climate of the area are described, with brief reference to neighbouring desert zones. The birds recorded are listed in weekly or part-weekly totals, followed by subspecific and other comments on selected species.
The movements of migrants are summarized, together with the effects of adverse winds on migration; very bad weather along the northern desert fringe may contribute to the late arrival of some species in the British Isles (and presumably elsewhere in Europe) in spring by causing delays and heavy mortality. Most visible migration was northeastward or eastward, but substantial westward movements of hirundines took place.
Comparison is made with migration through other areas in Morocco and Algeria, both from existing literature and from previously unpublished data, which provide evidence of broad-front migration across the entire width of the Moroccan Sahara. Birds recorded in northwest Algeria in 1966 are listed, together with notes on physical features, climate and field observations. Many terns and waders were recorded at Daiat Tchiour, including some predominantly coastal species, which suggests that the latter were migrating on a great-circle track from winter quarters in West Africa across the Sahara to their breeding grounds in Siberia.
Recoveries of ringed birds are discussed; some of these, together with records of species well west of their normal migratory range, suggest that birds may be subject to lateral displacement by winds when crossing the Sahara, leading to considerable annual variation in species and numbers of birds recorded on the northern desert edge, even in the same place.     

Songbird migration across the Sahara: the non-stop hypothesis rejected!

Billions of songbirds breeding in the Western Palaearctic cross the largest desert of the world, the Sahara, twice a year. While crossing Europe, the vast majority use an intermittent flight strategy, i.e. fly at night and rest or feed during the day. However, it was long assumed that they overcome the Sahara in a 40 h non-stop flight. In this study, we observed bird migration with radar in the plain sand desert of the Western Sahara (Mauritania) during autumn and spring migration and revealed a clear prevalence of intermittent migration. Massive departures of songbirds just after sunset independent of site and season suggests strongly that songbirds spent the day in the plain desert. Thus, most songbirds cross the Sahara predominantely by the intermittent flight strategy. Autumn migration took place mainly at low altitudes with high temperatures, its density decreased abruptly before sunrise, followed by very little daytime migration. Migration was highly restricted to night-time and matched perfectly the intermittent flight strategy. However, in spring, when migratory flights occurred at much higher altitudes than in autumn, in cool air, about 17% of the songbird migration occurred during the day. This suggests that flying in high temperatures and turbulent air, as is the case in autumn, may lead to an increase in water and/or energy loss and may prevent songbirds from prolonged flights into the day.     

WATER-BIRDS OVER THE SAHARA

An investigation is made of the extent to which Palaearctic birds belonging to typically water-bird families occur south of the Sahara in areas remote from both the sea and the Nile, namely, Darfur, Northern Nigeria and the Inundation Zone of the Niger, and hence may be presumed to cross the Sahara. Fifty-seven species fall to be considered and for these occurrences within the Sahara are also collated. The results owe a great deal to work done in the last few years, especially ringing. Most of the 57 species dealt with occur regularly in appreciable numbers in one or more of the areas under consideration and some of the species are enormously common. In general there is evidence to show that many birds winter in Africa far to the west of their Palaearctic breeding grounds, from which much diagonal passage of the Sahara is to be inferred. The results are considered in relation to the ecological conditions on the northern and southern edges of the desert and in relation to the meteorological conditions over it. A remarkable circumstance is that few water-birds indeed of any species have been observed in the oases of the Sahara, so that their crossing of the desert is clearly very efficient, although so much of it is by lengthy diagonal and although many birds entering Africa in the middle longitudes meet with no suitable habitat between Europe and the tropics. Except perhaps for storks, which soar, and certainly for herons, which proceed by heavy flapping, the air speed of water-birds is much greater than that of the small passerine migrants, the necessary potentiality of which has been calculated as 50–60 hours of flight without refuelling; but too little is known of the aerodynamics and physiology involved for it to be calculated whether water-birds need to start with the comparable accumulations of fat and no definite information exists except for the Wood Sandpiper, which has been shown to put on up to 32% of fat. By comparison with passerines, more individual water-bird species winter partly north and partly south of the Sahara; the factors affecting each bird's behaviour in this respect are unknown.     

The daily pattern of autumn bird migration in the northern Sahara

The temporal pattern of migration by passerine birds during the night, and their arrival during the day at the Egyptian coast and in the northern Sahara Desert was investigated. The mean direction of nocturnal migration at the coast was south-southeast, while at all desert sites it was south-southwest.
Birds arrived at the Egyptian coast only during the second half of the night which is explained by the fact that no birds could have taken off from the Mediterranean Sea. At least some of the birds landed at the coast where they spent the day before taking off shortly after sunset. These birds passed the desert sites at the expected time of day assuming a ground speed of 18 m per second. However, the origin of the birds passing the desert sites early at night is unclear. They must either have spent the day in the desert north of the study sites or they had overflown the Egyptian coast in the afternoon without landing.
The landing of birds during the day at the desert sites was bimodal. This pattern of arrival is explained either by some birds having landed at the Egyptian coast in the early morning before continuing, or by deteriorating conditions later in the day during flight or when resting in the desert, that obliged them to seek shelter at the desert sites.
A correlation between the number of migrants observed during the night and the number of resting birds in the desert on the following day suggests that an unknown proportion of birds might regularly use an intermittent migratory strategy that includes rest periods by day when crossing the desert, whereas others might adapt a non-stop migratory strategy.     

How hazardous is the Sahara Desert crossing for migratory birds? Indications from satellite tracking of raptors

We investigated the risk associated with crossing the Sahara Desert for migrating birds by evaluating more than 90 journeys across this desert by four species of raptors (osprey Pandion haliaetus, honey buzzard Pernis apivorus, marsh harrier Circus aeruginosus and Eurasian hobby Falco subbuteo) recorded by satellite telemetry. Forty per cent of the crossings included events of aberrant behaviours, such as abrupt course changes, slow travel speeds, interruptions, aborted crossings followed by retreats from the desert and failed crossings due to death, indicating difficulties for the migrants. The mortality during the Sahara crossing was 31 per cent per crossing attempt for juveniles (first autumn migration), compared with only 2 per cent for adults (autumn and spring combined). Mortality associated with the Sahara passage made up a substantial fraction (up to about half for juveniles) of the total annual mortality, demonstrating that this passage has a profound influence on survival and fitness of migrants. Aberrant behaviours resulted in late arrival at the breeding grounds and an increased probability of breeding failure (carry-over effects). This study also demonstrates that satellite tracking can be a powerful method to reveal when and where birds are exposed to enhanced risk and mortality during their annual cycles.     

Flight-range estimates for small trans-Sahara migrants

Arguments in support of the non-stop and the intermittent strategies for crossing the Sahara have been based on data on the fat reserves of birds before the crossing and of birds grounded in the desert. In this paper, flight-range estimates were calculated and the necessary assumptions about air speed, energy input during flight, and energy equivalent of body reserves were evaluated. As examples, Willow Warblers Phylloscopus trochilus and Garden Warblers Sylvia borin were investigated during autumn migration from two study sites north of the Sahara and two study sites in the desert. In still air, the flight-range for both species at all study sites was too short to reach the Sahel zone without refuelling. It is concluded that birds depend on tailwinds for a successful crossing, independent of a non-stop or an intermittent migratory strategy, and that weather conditions in autumn allow them to rely on tailwinds.     

STOPOVER AND FAT DEPOSITION BY BLACKCAPS SYLVIA ATRICAPILLA FOLLOWING SPRING MIGRATION OVER THE SAHARA

Maitav, A. & Izhaki, I. 1994. Stopover and fat deposition by Blackcaps Sylvia atricapilla following spring migration over the Sahara. Ostrich 65:160-166.

Length of stopover and rate of weight change were studied in Blackcaps that stopover in Eilat (southern Israel) in spring after a trans-Saharan flight. The birds studied had already completed 2000 km of desert crossing and faced just 200 km more. The potential predicted flight range for these transients suggests that the fat depleted Blackcaps (fat class=0) could not reach the east Mediterranean scrublands, which offer more extensive foraging opportunities than the desert area around Eilat, without refueling. However, 46% of the Blackcaps had sufficient fat (fat class > 0) to reach central or northern Israel in a single flight in still air. Recaptured birds stayed in Eilat for 2,4±1,9 days. Although the body mass of an average recaptured bird increased by 0,27±1,44 g/day, which was an increase of 0,8%±7,0% of its body mass per day during stopover, these results were not significantly different from zero. Stopover length, mass change during stopover, and the rate of change in mass did not differ between males and females and between those who stopped over for only one day to those which stayed > 1 day. However, there was a tendency of these lean birds to stay for a shorter period in Eilat than relatively fat birds. There was also some tendency for lean birds to gain more mass during their stopover than fat birds. It seems that in spring, when less time is allotted for the whole migration program, the migrants resume their journey before fat reserves have been replenished.     

THE IBIS

Some aspects of the weights of over 5,500 birds of 79 species, mostly migrants, captured in spring at an oasis on the northern edge of the Sahara, are discussed. Mean weights for five species were 26–44% below those of fat birds (those containing at least 30% lipid) weighed about the same time on the south side of the desert. Weights of fresh arrivals in Britain are higher than those in North Africa, but the differences are not great, suggesting that these have also made a long journey across Europe. Peak arrival times of day-migrants differed from those of night-migrants and the weights of some species varied with the time of day. There was no great advantage to migrants in remaining at the oasis; there were few weight changes greater than those normally occurring within an individual's daily range. Several species lost weight steadily during a cold spell, and many died. Male Swallows are heavier and larger than females, and in both sexes weight increases with wing-length.     

Interaction of bodymass,fat, foraging and stopover period in trans-Sahara migrating passerine birds

Summary The effect of body mass and fat reserves on the choice of the stopover place and on the stopover period was investigated in 3 species of passerine birds migrating through the Sahara in Egypt. Birds grounded in an oasis with food and water were more than 10% lighter than those from a desert stopover place which offered shade only. Stopover period was in general one day except for the light portion of oasis birds which stopped for up to three weeks during which they regained fat. A calculation of the maximum range which can be covered with the available fat depots revealed that nearly all birds from the desert stopover place should be able to cross the desert whereas about 60% of the oasis birds should not. An intermittant migratory strategy for the desert crossing is proposed with flight during the night and stopover during the day, even without drinking and feeding.     

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.     

Migratory and targeted flight in seasonal forms of the black bean aphid, Aphis fabae

ABSTRACT. Adult winged forms of Aphis fabae Scop., the spring migrants (fundatrigeniae; alate virginoparae), the summer migrants (alate virginoparae) and the autumn migrants (gynoparae), were flown in an automated vertical wind tunnel, with a small green target presented every 60 s. On average, targeted flight developed in the spring and summer migrants after 19 and 15 min, respectively. However, in autumn migrants targeted flight occurred after 184 min of flight. No evidence of target approach was found up to 30 min prior to this time, although some movement towards the target was observed in the 30 min before the first response of autumn migrants, suggesting that a gradual change from migratory to targeted flight occurred. No correlation was found between aphid weight and the time to first response for spring or summer migrants, but a positive correlation was found for autumn migrants. Flight patterns of autumn migrants, in the absence of a target, are described and compared with flight patterns previously reported for summer migrants.     

Interspecific comparison of the performance of soaring migrants in relation to morphology, meteorological conditions and migration strategies

Background

Performance of migrating birds can be affected by a number of intrinsic and extrinsic factors like morphology, meteorological conditions and migration strategies. We compared travel speeds of four raptor species during their crossing of the Sahara desert. Focusing the analyses on this region allows us to compare different species under equivalent conditions in order to disentangle which factors affect migratory performance.

Methodology/Principal Finding

We tracked raptors using GPS satellite transmitters from Sweden, Spain and Italy, and evaluated their migratory performance at both an hourly and a daily scale. Hourly data (flight speed and altitude for intervals of two hours) were analyzed in relation to time of day, species and season, and daily data (distance between roosting sites) in relation to species, season, day length and tailwind support.

Conclusions/Significance

Despite a clear variation in morphology, interspecific differences were generally very small, and did only arise in spring, with long-distance migrants (>5000 km: osprey and Western marsh-harrier) being faster than species that migrate shorter distances (Egyptian vulture and short-toed eagle). Our results suggest that the most important factor explaining hourly variation in flight speed is time of day, while at a daily scale, tailwind support is the most important factor explaining variation in daily distance, raising new questions about the consequences of possible future changes in worldwide wind patterns.