Experience does not change the importance of wind support for migratory route selection by a soaring bird.

Migration is a complex behaviour that is costly in terms of time, energy and risk of mortality. Thermal soaring birds rely on airflow, specifically wind support and uplift, to offset their energetic costs of flight. Their migratory routes are a record of movement decisions to negotiate the atmospheric environment and achieve efficiency. We expected that, regardless of age, birds use wind support to select their routes. Because thermal soaring is a complex flight behaviour that young birds need to learn, we expected that, as individuals gain more experience, their movement decisions will also increasingly favour the best thermal uplift conditions. We quantified how route choice during autumn migration of young European honey buzzards (Pernis apivorus) was adjusted to wind support and uplift over up to 4 years of migration and compared this with the choices of adult birds. We found that wind support was important in all migrations. However, we did not find an increase in the use of thermal uplifts. This could be due to the species-specific learning period and/or an artefact of the spatio-temporal scale of our uplift proxies.

The interplay of wind and uplift facilitates over-water flight in facultative soaring birds.

Flying over the open sea is energetically costly for terrestrial birds. Despite this, over-water journeys of many birds, sometimes hundreds of kilometres long, are uncovered by bio-logging technology. To understand how these birds afford their flights over the open sea, we investigated the role of atmospheric conditions, specifically wind and uplift, in subsidizing over-water flight at a global scale. We first established that ΔT, the temperature difference between sea surface and air, is a meaningful proxy for uplift over water. Using this proxy, we showed that the spatio-temporal patterns of sea-crossing in terrestrial migratory birds are associated with favourable uplift conditions. We then analysed route selection over the open sea for five facultative soaring species, representative of all major migratory flyways. The birds maximized wind support when selecting their sea-crossing routes and selected greater uplift when suitable wind support was available. They also preferred routes with low long-term uncertainty in wind conditions. Our findings suggest that, in addition to wind, uplift may play a key role in the energy seascape for bird migration that in turn determines strategies and associated costs for birds crossing ecological barriers such as the open sea.

Adaptive drift and barrier-avoidance by a fly-forage migrant along a climate-driven flyway.

BACKGROUND: Route choice and travel performance of fly-forage migrants are partly driven by large-scale habitat availability, but it remains unclear to what extent wind support through large-scale wind regimes moulds their migratory behaviour. We aimed to determine to what extent a trans-equatorial fly-forage migrant engages in adaptive drift through distinct wind regimes and biomes across Africa. The Inter-tropical Front (ITF) marks a strong and seasonally shifting climatic boundary at the thermal equator, and we assessed whether migratory detours were associated with this climatic feature. Furthermore, we sought to disentangle the influence of wind and biome on daily, regional and seasonal travel performance. METHODS: We GPS-tracked 19 adult Eleonora's falcons Falco eleonorae from the westernmost population on the Canary Islands across 39 autumn and 36 spring migrations to and from Madagascar. Tracks were annotated with wind data to assess the falcons' orientation behaviour and the wind support they achieved in each season and distinct biomes. We further tested whether falcon routes across the Sahel were correlated with the ITF position, and how realized wind support and biome affect daily travel times, distances and speeds. RESULTS: Changes in orientation behaviour across Africa's biomes were associated with changes in prevailing wind fields. Falcons realized higher wind support along their detours than was available along the shortest possible route by drifting through adverse autumn wind fields, but compromised wind support while detouring through supportive spring wind fields. Movements across the Sahel-Sudan zone were strongly associated to the ITF position in autumn, but were more individually variable in spring. Realized wind support was an important driver of daily travel speeds and distances, in conjunction with regional wind-independent variation in daily travel time budgets. CONCLUSIONS: Although daily travel time budgets of falcons vary independently from wind, their daily travel performance is strongly affected by orientation-dependent wind support. Falcons thereby tend to drift to minimize or avoid headwinds through opposing wind fields and over ecological barriers, while compensating through weak or supportive wind fields and over hospitable biomes. The ITF may offer a climatic leading line to fly-forage migrants in terms of both flight and foraging conditions.

Barrier crossings and winds shape daily travel schedules and speeds of a flight generalist.

External factors such as geography and weather strongly affect bird migration influencing daily travel schedules and flight speeds. For strictly thermal-soaring migrants, weather explains most seasonal and regional differences in speed. Flight generalists, which alternate between soaring and flapping flight, are expected to be less dependent on weather, and daily travel schedules are likely to be strongly influenced by geography and internal factors such as sex. We GPS-tracked the migration of 70 lesser kestrels (Falco naumanni) to estimate the relative importance of external factors (wind, geography), internal factors (sex) and season, and the extent to which they explain variation in travel speed, distance, and duration. Our results show that geography and tailwind are important factors in explaining variation in daily travel schedules and speeds. We found that wind explained most of the seasonal differences in travel speed. In both seasons, lesser kestrels sprinted across ecological barriers and frequently migrated during the day and night. Conversely, they travelled at a slower pace and mainly during the day over non-barriers. Our results highlighted that external factors far outweighed internal factors and season in explaining variation in migratory behaviour of a flight generalist, despite its ability to switch between flight modes.

Orientation of native versus translocated juvenile lesser spotted eagles (Clanga pomarina) on the first autumn migration.

The ontogeny of migration routines used by wild birds remains unresolved. Here we investigated the migratory orientation of juvenile lesser spotted eagles (LSE; Clanga pomarina) based on translocation and satellite tracking. Between 2004 and 2016, 85 second-hatched juveniles (Abels) were reared in captivity for release into the declining German population, including 50 birds that were translocated 940 km from Latvia. In 2009, we tracked 12 translocated juveniles, as well as eight native juveniles and nine native adults, to determine how inexperienced birds come to use strategic migration routes. Native juveniles departed around the same time as the adults and six of eight used the eastern flyway around the Mediterranean, which was used by all adults. In contrast, translocated juveniles departed on average 6 days before native LSEs, and five travelled southward and died in the central Mediterranean region. Consequently, fewer translocated juveniles (4/12) than native juveniles (7/8) reached Africa. We conclude that juvenile LSEs have a much better chance of learning the strategic southeastern flyway if they leave at an appropriate time to connect with experienced elders upon departure. It is not clear why translocated juveniles departed so early. Regardless, by the end of the year, most juveniles had perished, whether they were translocated (10/12) or not (6/8). The small number of surviving translocated juveniles thus still represents a significant increase in the annual productivity of the German LSE population in 2009.

Atmospheric conditions create freeways, detours and tailbacks for migrating birds.

The extraordinary adaptations of birds to contend with atmospheric conditions during their migratory flights have captivated ecologists for decades. During the 21st century technological advances have sparked a revival of research into the influence of weather on migrating birds. Using biologging technology, flight behaviour is measured across entire flyways, weather radar networks quantify large-scale migratory fluxes, citizen scientists gather observations of migrant birds and mechanistic models are used to simulate migration in dynamic aerial environments. In this review, we first introduce the most relevant microscale, mesoscale and synoptic scale atmospheric phenomena from the point of view of a migrating bird. We then provide an overview of the individual responses of migrant birds (when, where and how to fly) in relation to these phenomena. We explore the cumulative impact of individual responses to weather during migration, and the consequences thereof for populations and migratory systems. In general, individual birds seem to have a much more flexible response to weather than previously thought, but we also note similarities in migratory behaviour across taxa. We propose various avenues for future research through which we expect to derive more fundamental insights into the influence of weather on the evolution of migratory behaviour and the life-history, population dynamics and species distributions of migrant birds.

Seasonal detours by soaring migrants shaped by wind regimes along the East Atlantic Flyway.

Avian migrants often make substantial detours between their seasonal destinations. It is likely some species do this to make the most of predictable wind regimes along their respective flyways. We test this hypothesis by studying orientation behaviour of a long-distance soaring migrant in relation to prevailing winds along the East Atlantic Flyway. We tracked 62 migratory journeys of 12 adult European Honey Buzzards Pernis apivorus with GPS loggers. Hourly fixes were annotated with local wind vectors from a global atmospheric model to determine orientation behaviours with respect to the buzzards' seasonal goal destinations. This enabled us to determine hot spots where buzzards overdrifted and overcompensated for side winds. We then determined whether winds along the buzzards' detours differed from winds prevailing elsewhere in the flyway. Honey Buzzards cross western Africa using different routes in autumn and spring. In autumn, they overcompensated for westward winds to circumvent the Atlas Mountains on the eastern side and then overdrifted with south-westward winds while crossing the Sahara. In spring, however, they frequently overcompensated for eastward winds to initiate a westward detour at the start of their journey. They later overdrifted with side winds north-westward over the Sahel and north-eastward over the Sahara, avoiding adverse winds over the central Sahara. We conclude that Honey Buzzards make seasonal detours to utilize more supportive winds further en route and thereby expend less energy while crossing the desert. Lifelong tracking studies will be helpful to elucidate how honey buzzards and other migrants learn complex routes to exploit atmospheric circulation patterns from local to synoptic scales.