Risk-sensitive response of soaring birds to crosswind over dangerous sea highlights age-specific differences in migratory performance.

Challenges imposed by geographical barriers during migration are selective agents for animals. Juvenile soaring landbirds often cross large water bodies along their migratory path, where they lack updraft support and are vulnerable to harsh weather. However, the consequences of inexperience in accomplishing these water crossings remain largely unquantified. To address this knowledge gap, we tracked the movements of juvenile and adult black kites Milvus migrans over the Strait of Gibraltar using high-frequency tracking devices in variable crosswind conditions. We found that juveniles crossed under higher crosswind speeds and at wider sections of the strait compared with adults during easterly winds, which represent a high risk owing to their high speed and steady direction towards the Atlantic Ocean. Juveniles also drifted extensively with easterly winds, contrasting with adults who strongly compensated for lateral displacement through flapping. Age differences were inconspicuous during winds with a west crosswind speed component, as well as for airspeed modulation in all wind conditions. We suggest that the suboptimal sea-crossing behaviour of juvenile black kites may impact their survival rates, either by increasing chances of drowning owing to exhaustion or by depleting critical energy reserves needed to accomplish their first migration.

Time constraints may pace the ontogeny of movement behaviour.

During early development, juvenile animals need to acquire a diverse behavioural repertoire to interact with their environment. The ontogeny of animal behaviour, is paced by the motivation to improve, e.g. internal clocks, and limited by external constraints, e.g. weather conditions. We here evaluate how naive Egyptian vultures (Neophron percnopterus) improve in locomotor performance, measured as daily maximum displacement, prior to their first migration under three different time constraint regimes: we compared wild hatched vultures, migrating one month after fledging, with captive-hatched vultures, released in spring four months or in winter nine months before migration. We found that the time until migration paced the development of movement behaviour: wild birds rapidly increased displacement distances within the first two weeks after fledging, while spring and winter released vultures delayed movement increases by two and four months, respectively. Under relaxed time constraints captive-hatched vultures displayed diverse functional forms of performance enhancements and therefore great variability in individual ontogeny of movement behaviour. While weather conditions in winter could limit flight movements, some birds indeed moved immediately after their release, indicating that weather may not be limiting. Our findings promote the idea that relaxed ecological constraints could uncover hidden phenotypic flexibility in ontogeny, which could present a greater potential for adaptability under environmental change than currently expected.

The gateway to Africa: What determines sea crossing performance of a migratory soaring bird at the Strait of Gibraltar?

Large bodies of water represent major obstacles for the migration of soaring birds because thermal updrafts are absent or weak over water. Soaring birds are known to time their water crossings with favourable weather conditions and there are records of birds falling into the water and drowning in large numbers. However, it is still unclear how environmental factors, individual traits and trajectory choices affect their water crossing performance, this being important to understand the fitness consequences of water barriers for this group of birds. We addressed this problem using the black kite Milvus migrans as model species at a major migration bottleneck, the Strait of Gibraltar. We recorded high-resolution GPS and triaxial accelerometer data for 73 birds while crossing the Strait of Gibraltar, allowing the determination of sea crossing duration, length, altitude, speed and tortuosity, the flapping behaviour of birds and their failed crossing attempts. These parameters were modelled against wind speed and direction, time of the day, solar irradiance (proxy of thermal uplift), starting altitude and distance to Morocco, and age and sex of birds. We found that sea crossing performance of black kites is driven by their age, the wind conditions, the starting altitude and distance to Morocco. Young birds made longer sea crossings and reached lower altitude above the sea than adults. Crosswinds promoted longer sea crossings, with birds reaching lower altitudes and with higher flapping effort. Birds starting at lower altitudes were more likely to quit or made higher flapping effort to complete the crossing. The location where birds started the sea crossings impacted crossing distance and duration. We present evidence that explains why migrating soaring birds accumulate at sea passages during adverse weather conditions. Strong crosswinds during sea crossings force birds to extended flap-powered flight at low altitude, which may increase their chances of falling in the water. We also showed that juvenile birds assume more risks than adults. Finally, the way in which birds start the sea crossing is crucial for their success, particularly the starting altitude, which dictates how far birds can reach with reduced flapping effort.

The gliding speed of migrating birds: slow and safe or fast and risky?

Aerodynamic theory postulates that gliding airspeed, a major flight performance component for soaring avian migrants, scales with bird size and wing morphology. We tested this prediction, and the role of gliding altitude and soaring conditions, using atmospheric simulations and radar tracks of 1346 birds from 12 species. Gliding airspeed did not scale with bird size and wing morphology, and unexpectedly converged to a narrow range. To explain this discrepancy, we propose that soaring-gliding birds adjust their gliding airspeed according to the risk of grounding or switching to costly flapping flight. Introducing the Risk Aversion Flight Index (RAFI, the ratio of actual to theoretical risk-averse gliding airspeed), we found that inter- and intraspecific variation in RAFI positively correlated with wing loading, and negatively correlated with convective thermal conditions and gliding altitude, respectively. We propose that risk-sensitive behaviour modulates the evolution (morphology) and ecology (response to environmental conditions) of bird soaring flight.