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

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

Migrating Montagu's harriers frequently interrupt daily flights in both Europe and Africa

Time budgets are a powerful but hitherto seldom used way to study how migrants organise their bi‐annual travels. We studied daily time budgets of travelling Montagu's harriers Circus pygargus, based on GPS tracking data, in which we were particularly interested in how time budgets differ between regions and seasons, and are affected by wind. We found that Montagu's harriers used a relatively broad daily time window for travelling by starting daily travels just after sunrise and ending daily travels just before sunset. Occasionally, flights were extended into the night. Montagu's harriers frequently interrupted their daily flights for on average 1.5 h d^–1. These interruptions occurred in all regions and seasons. The tracking data during interruptions suggested two different behaviours: in 41% of all interruptions the birds were moving (presumed foraging,) and in 32% they were stationary (presumed resting; the remaining interruptions could not be classified). The interruptions for foraging indicate that Montagu's harriers have a fly‐and‐forage migration strategy (i.e. combine travelling and foraging on the same day), but the interruptions for resting illustrate that their travels comprise of more than fly‐and‐forage behaviour alone. The large number of interruptions for foraging in the Sahara Desert indicates that this region is less hostile for a migrating raptor than presumed previously. Importantly, harriers spent more time on interruptions for resting on days with stronger headwinds, suggesting that interruptions for resting serve a function of waiting for more favourable weather conditions. Daily variation in time budgets was largely explained by wind; harriers flew more hours per day, and interrupted their flights fewer hours per day, on days they experienced stronger tailwinds. In contrast, time budgets were similar between regions and seasons, suggesting that wind rather than landscape and season shape travel routines of Montagu's harriers.     

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

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

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

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

Is there a connection between weather at departure sites,onset of migration and timing of soaring‐bird autumn migration in Israel?

Aims Different aspects of soaring‐bird migration are influenced by weather. However, the relationship between weather and the onset of soaring‐bird migration, particularly in autumn, is not clear. Although long‐term migration counts are often unavailable near the breeding areas of many soaring birds in the western Palaearctic, soaring‐bird migration has been systematically monitored in Israel, a region where populations from large geographical areas converge. This study tests several fundamental hypotheses regarding the onset of migration and explores the connection between weather, migration onset and arrival at a distant site. Location Globally gridded meteorological data from the breeding areas in north‐eastern Europe were used as predictive variables in relation to the arrival of soaring migrants in Israel. Methods Inverse modelling was used to study the temporal and spatial influence of weather on initiation of migration based on autumn soaring‐bird migration counts in Israel. Numerous combinations of migration duration and temporal influence of meteorological variables (temperature, sea‐level pressure and precipitable water) were tested with different models for meteorological sensitivity. Results The day of arrival in Israel of white storks, honey buzzards, Levant sparrowhawks and lesser spotted eagles was significantly and strongly related to meteorological conditions in the breeding area days or even weeks before arrival in Israel. The cumulative number of days or cumulative value above or below a meteorological threshold performed significantly better than other models tested. Models provided reliable estimates of migration duration for each species. Main conclusions The meteorological triggers of migration at the breeding grounds differed between species and were related to deteriorating living conditions and deteriorating migratory flight conditions. Soaring birds are sensitive to meteorological triggers at the same period every year and their temporal response to weather appears to be constrained by their annual routine.     

Loop migration in adult marsh harriers Circus aeruginosus,as revealed by satellite telemetry

Loop migration among birds is characterized by the spring route lying consistently west or east of the autumn route. The existence of loops has been explained by general wind conditions or seasonal differences in habitat distribution. Loop migration has predominantly been studied at the population level, for example by analysing ring recoveries. Here we study loop migration of individual marsh harriers Circus aeruginosus tracked by satellite telemetry. We show that despite a generally narrow migration corridor the harriers travelled in a distinct clockwise loop through Africa and southern Europe, following more westerly routes in spring than in autumn. We used the Normalized Difference Vegetation Index (NDVI) to identify potential feeding habitat in Africa. Suitable habitat seemed always more abundant along the western route, both in spring and autumn, and no important stopover site was found along the eastern route. Observed routes did thus not coincide with seasonal variation in habitat availability. However, favourable habitat might be more important during spring migration, when the crossing of the Sahara seems more challenging, and thus habitat availability might play an indirect role in the harriers’ route choice. Grid‐based wind data were used to reconstruct general wind patterns, and in qualitative agreement with the observed loop marsh harriers predominantly encountered westerly winds in Europe and easterly winds in Africa, both in autumn and in spring. By correlating tail‐ and crosswinds with forward and perpendicular movement rates, respectively, we show that marsh harriers are partially drifted by wind. Thus, we tentatively conclude that wind rather than habitat seems to have an overriding effect on the shape of the migration routes of marsh harriers. General wind conditions seem to play an important role also in the evolution of narrow migratory loops as demonstrated for individual marsh harriers.     

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.     

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

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

Local and regional weather patterns influencing post‐breeding migration counts of soaring birds at the Strait of Gibraltar,Spain

Migration is a significant event in the annual cycle of many avian species. During migration birds face many challenges, including unfamiliar foraging and refuge habitats, resulting in a much higher rate of mortality during migration than during other seasons of the year. Weather may significantly affect a bird's decision to initiate migration, the course and pace of migration, and its survival during migration. Each of these influences may impact the counts of migrating birds at geographical convergence zones or bottlenecks. It is important to quantify the effect of short‐term weather on these counts to appropriately interpret and use such counts in other analyses. To this end, we aim to assess the effects of local and regional weather conditions on the migration counts of soaring birds at the Strait of Gibraltar during post‐breeding migration. We used information‐theoretic approaches to analyse the influence of local weather and weather in northern Spain on the migration counts of five soaring bird species from two count sites near the Strait of Gibraltar. Migration counts were higher on days with local northerly and westerly winds, often following a day of easterly winds, on days with local high pressure systems, and often following a day of lower pressure. Weather conditions in northern Spain influenced migration counts at the Strait of Gibraltar, but the effects were much weaker than local weather conditions. We confirm that short‐term weather conditions, locally and regionally, can influence migration counts and should thus be considered when these counts are used in other analyses.     

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

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

Effect of abiotic factors on reproduction in the centre and periphery of breeding ranges: a comparative analysis in sympatric harriers

Variables such as weather or other abiotic factors should have a higher influence on demographic rates in border areas than in central areas, given that climatic adaptation might be important in determining range borders. Similarly, for a given area, the relationship between weather and reproduction should be dissimilar for species which are in the centre of their breeding range and those that are near the edge. We tested this hypothesis on two sympatric ground‐nesting raptors, the hen harrier Circus cyaneus and the Montagu's harrier Circus pygargus in Madrid, central Spain, where the hen harrier is at the southern edge of its breeding range in the western Palearctic and the Montagu's harrier is central in its distribution. We examined the reproductive success of both species during an 8‐yr period, and looked at the influence of the most stressful abiotic factors in the study area (between‐year variation in rainfall and within‐year variation in temperature) on reproductive parameters. In the hen harrier, low levels of rainfall during the breeding season had a negative influence on annual fledging success and thus on population fledgling production. The relationship between rainfall and reproduction was probably mediated through food abundance, which in Mediterranean habitat depends directly on rainfall levels. In the Montagu's harrier, no negative effect of dry seasons on productivity was found. Additionally, in the hen harrier, the proportion of eggs that did not hatch in each clutch increased with higher temperatures during the incubation period. No such relationship was found in the Montagu's harrier. We interpret these between‐species differences in terms of differences of breeding range and adaptations to the average conditions existing there. Hen harriers, commonest at northern latitudes, are probably best adapted to the most typical conditions at those latitudes, and have probably not developed thermoregulatory or behavioural mechanisms to cope with drought and high temperatures in Mediterranean habitats, in contrast to Montagu's harrier. Thus hen harrier distribution might be constrained by these variables, due to lower reproductive success or higher reproductive costs. Accordingly, a logistic regression analysis of the presence or absence of both species in 289 random points throughout the western Palearctic showed that the distribution of both species was related to temperature, but the relationship was in opposite directions for the two species: hen harriers had lower probability of breeding in areas with higher temperature (as expected in a species with a more northerly distribution).     

Diet specialisation and foraging efficiency under fluctuating vole abundance: a comparison between generalist and specialist avian predators

Specialist species, using a narrow range of resources, are predicted to be more efficient when foraging on their preferred food than generalist species consuming a wider range of foods. We tested whether the foraging efficiency of the pallid harrier Circus macrourus, a vole specialist, and of sympatric Montagu's harriers C. pygargus, a closely related generalist, differed in relation to inter‐annual variations in vole abundance over five years (including two peak‐ one intermediate and two low vole abundance years). We show that the hunting parameters of pallid harriers strongly varied with vole abundance (higher encounter rates, capture rates and proportion of successful strikes in high than intermediate and low vole abundance years, respectively), whereas Montagu's harriers showed stable capture rates and hunting success (proportion of strikes that were successful), irrespective of vole abundance. Encounter rates and capture rates were higher for pallid than for Montagu's harriers when voles were abundant, but lower when voles were scarce. The hunting success of pallid harriers was also lower than that of Montagu's harriers when voles were scarce, and when they had to target alternative preys, in particular birds. Overall, estimated biomass intake rate was 40% higher for pallid harriers than for Montagu's harriers when voles were abundant, but 50% lower when voles were scarce. Our results indicate that specialists predators, like pallid harriers, which evolve specific adaptations or breeding strategies, do better when their preferred prey is abundant, but may face a cost of specialisation, being not efficient enough when their preferred prey is scarce. These results have broader implications for understanding why specialist predators are, in general, more vulnerable than generalists, and for predicting how specialists can cope with rapid environmental changes affecting the abundance or predictability of their preferred resources.     

Flight mode affects allometry of migration range in birds

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

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

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

Rapid long‐distance migration in Norwegian Lesser Black‐backed Gulls Larus fuscus fuscus along their eastern flyway

We studied the long‐distance migration of Lesser Black‐backed Gulls Larus fuscus fuscus breeding in northern Norway along their eastern flyway using geolocators in 2009 and 2010. The majority of birds wintered in lakes in East Africa and the southeast Mediterranean was the most important stopover area. Larus f. fuscus along the eastern flyway travelled at a net travel speed of 399 and 177 km/day during the autumn and spring migration, respectively, higher than published travel speeds for Dutch Larus fuscus migrating along the western flyway. The results suggest that the long‐distance migratory Norwegian L. f. fuscus seek to minimize time spent in transit, whereas lower travel speed during northerly spring migration may reflect differences in wind patterns or food conditions between spring and autumn.     

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

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

Limitations and mechanisms influencing the migratory performance of soaring birds

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

Do Seabirds Differ from Other Migrants in Their Travel Arrangements? On Route Strategies of Cory’s Shearwater during Its Trans-Equatorial Journey

Long-distance migrants have developed diverse strategies to deal with the challenges imposed by their annual journeys. These are relatively well studied in some avian groups, such as passerines, shorebirds and raptors. In contrast, few studies have addressed the migratory behaviour of pelagic birds in the light of current theories of optimal migration. Using a dataset of 100 complete migratory tracks gathered along four years, we performed a detailed study on the migratory strategy of a pelagic trans-equatorial migrant, the Cory’s shearwater Calonectris diomedea. We analysed daily routines, stopover ecology and travel speed, as well as the influence of the moon on several behavioural patterns. Cory’s shearwaters adopted a “fly-and-forage” strategy when migrating, similarly to what has been observed in some raptors. However, by flying by dynamic soaring, shearwaters attained high overall migration speeds, and were able to travel thousands of kilometres without making major stopovers and, apparently, without a noticeable pre-migratory fattening period. Other major findings of this study include the ability to adapt daily schedules when crossing major ecological barriers, and the constant adjustment of migration speed implying higher rates of travel in the pre-breeding movement, with a final sprint to the nesting colony. The present study also highlights a preference of Cory’s shearwaters for starting travel at twilight and documents a strong relationship between their migratory activity and the moon phase.     

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.     

Extremely detoured migration in an inexperienced bird: interplay of transport costs and social interactions

We tagged two juvenile short‐toed eagles in southern Italian peninsula with GPS satellite transmitters. According to previous visual observations, two different migratory routes for Italian short‐toed eagles to reach Africa in autumn have been proposed: via Sicily and via Gibraltar. These routes include different over‐water distances to cross the Mediterranean Sea, and thus different proportions of flight modes (soaring–gliding vs flapping–gliding) with resulting different transport costs. Considering different scenarios of energy cost of transport, with flapping–gliding flight over water being more costly than flying over land using soaring–gliding flight, we predicted a maximum optimal detour of 1218 km. Both individuals reached Africa using the longest, detoured, route, avoiding the longest water crossing. To achieve this they began migrating northwards, keeping for ca 700 km a direction opposite to that followed by any other migrating bird from the Northern hemisphere in autumn. The comparison of optimal detour predictions with observed migratory tracks suggests that this migratory strategy prioritizes not only energy minimization, but also safety, given the mortality risk associated with the sea crossing. Finally, it is unlike that these inexperienced individuals followed such a complex route relying only on endogenous information and we therefore suggest, also on the basis of field observations, that social interactions (adult guidance) allow these individuals to learn the detoured route.