Mikael Hake

When and where does mortality occur in migratory birds? Direct evidence from long‐term satellite tracking of raptors

Information about when and where animals die is important to understand population regulation. In migratory animals, mortality might occur not only during the stationary periods (e.g. breeding and wintering) but also during the migration seasons. However, the relative importance of population limiting factors during different periods of the year remains poorly understood, and previous studies mainly relied on indirect evidence. Here, we provide direct evidence about when and where migrants die by identifying cases of confirmed and probable deaths in three species of long-distance migratory raptors tracked by satellite telemetry. We show that mortality rate was about six times higher during migration seasons than during stationary periods. However, total mortality was surprisingly similar between periods, which can be explained by the fact that risky migration periods are shorter than safer stationary periods. Nevertheless, more than half of the annual mortality occurred during migration. We also found spatiotemporal patterns in mortality: spring mortality occurred mainly in Africa in association with the crossing of the Sahara desert, while most mortality during autumn took place in Europe. Our results strongly suggest that events during the migration seasons have an important impact on the population dynamics of long-distance migrants. We speculate that mortality during spring migration may account for short-term annual variation in survival and population sizes, while mortality during autumn migration may be more important for long-term population regulation (through density-dependent effects).

Bird orientation: compensation for wind drift in migrating raptors is age dependent

Despite the potentially strong effect of wind on bird orientation, our understanding of how wind drift affects migrating birds is still very limited. Using data from satellite-based radio telemetry, we analysed the effect of changing winds on the variation of the track direction of individual birds. We studied adults and juveniles of two raptor species, osprey Pandion haliaetus and honey buzzard Pernis apivorus, on autumn migration between North Europe and Africa, and demonstrate an important difference between the age categories of both species in the extent of wind drift. For juveniles, side- and following-wind components affected the rates of movement perpendicular to and along the mean direction, respectively, to a similar degree, suggesting full wind drift. By contrast, for adults the rate of crosswind displacement was significantly smaller than the effect of wind on forward movement, showing much reduced wind drift (29%). This indicates that adults have acquired a more sophisticated orientation system, permitting detection of and compensation for wind drift, than juveniles. These drift effects are likely to reduce the ability of juveniles to locate species-specific wintering areas in case of rapid climatic wind change.

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.

Geographical and temporal flexibility in the response to crosswinds by migrating raptors

Wind and ocean currents may potentially have important effects on travelling animals, as an animal which does not respond to lateral flow will be drifted from its intended direction of movement. By analysing daily movements of migrating ospreys Pandion haliaetus and marsh harriers Circus aeruginosus, as recorded by satellite telemetry, in relation to global wind data, we showed that these raptors allow on average 47 per cent drift. Furthermore, our analyses revealed significant geographical and temporal variation in the response to crosswinds. During some parts of the migration, the birds drifted and in other parts they compensated or even overcompensated. In some regions, the response of marsh harriers depended on the wind direction. They drifted when the wind came from one side and (over)compensated when the wind came from the opposite side, and this flexible response was different in different geographical regions. These results suggest that migrating raptors modulate their response to crosswinds at different places and times during their travels and show that individual birds use a much more varied repertoire of behavioural responses to wind than hitherto assumed. Our results may also explain why contrasting and variable results have been obtained in previous studies of the effect of wind on bird migration.

Complex timing of Marsh Harrier Circus aeruginosus migration due to pre- and post-migratory movements

We tracked three juvenile and 14 adult Marsh Harriers Circus aeruginosus from southern Sweden via satellite to investigate migration strategies. Four individuals were tracked for at least two years. All three juveniles and four of the adults made west-oriented pre-migratory movements well before the onset of autumn migration, and trans-Saharan migrants visited post-migratory stopover areas in tropical Africa. By these movements, the harriers presumably exploit short-term regional variation in food abundance. Autumn and spring migration occurred in a relatively narrow corridor, without distinct differences between sexes in timing, speed, distance, and duration of migration, except that females tended to migrate faster in spring than did males. Juveniles migrated shorter distances than adults, and migration speeds were lower. Spring migration was similar to autumn migration in terms of speed and duration. Juveniles did not cross the Sahara Desert and three birds, one female and two juveniles, wintered in Europe, which is in accordance with a recent increase in the number of (juvenile) Marsh Harriers wintering in northwestern Europe. All birds that crossed the Sahara wintered in tropical West Africa. Harriers showed site fidelity to breeding, wintering and stopover areas. The overall migration speed of Marsh Harriers was similar to that of Ospreys Pandion haliaetus and Honey Buzzards Pernis apivorus, two other trans-Saharan migrants. Ospreys use fly-and-forage migration to promote resulting speed, whereas Honey Buzzards are particularly apt to exploit thermal soaring. How Marsh Harriers balance foraging versus travelling to accomplish their rapid migration speeds remains to be resolved.

Fattening strategies in dominance-structured greenfinch (Carduelis chloris) flocks in winter

Abstract Relationships between social status and levels of body reserves stored by members of greenfinch (Carduelis chloris) flocks in winter were investigated. In addition, the adjustment of reserves by birds of different rank to experimental changes in food predictability and to changed weather conditions were examined. Birds with low social status carried overall larger body reserves than high-ranked birds. The results of the experiments suggest that this was mainly because subordinates, due to their low priority to food access, perceived future foraging success as less predictable than dominants. In response to severe weather, which probably increased the risk of starvation for birds, dominants temporarily increased their reserves more than subordinates. This response also indicated that birds with low social status carried larger reserves than high-ranked birds, and demonstrated that dominants could increase food intake when there was a risk of energetic shortfall. The results suggest that fattening strategies in greenfinches depend on social status. In winter, subordinates may be forced to carry larger reserves than dominants to safeguard against social constraints in access to food during critical times. As body reserves may be costly to carry and acquire, this should reduce the probability of surviving the winter for subordinates compared to dominants.

Converging migration routes of Eurasian hobbies Falco subbuteo crossing the African equatorial rain forest

Autumn migration of adult Eurasian hobbies Falco subbuteo from Europe to southern Africa was recorded by satellite telemetry and observed routes were compared with randomly simulated routes. Two non-random features of observed routes were revealed: (i) shifts to more westerly longitudes than straight paths to destinations and (ii) strong route convergence towards a restricted area close to the equator (1° S, 15° E). The birds migrated south or southwest to approximately 10° N, where they changed to south-easterly courses. The maximal spread between routes at 10° N (2134 km) rapidly decreased to a minimum (67 km) close to the equator. We found a striking relationship between the route convergence and the distribution of continuous rainforest, suggesting that hobbies minimize flight distance across the forest, concentrating in a corridor where habitat may be more suitable for travelling and foraging. With rainforest forming a possible ecological barrier, many migrants may cross the equator either at 15° E, similar to the hobbies, or at 30–40° E, east of the rainforest where large-scale migration is well documented. Much remains to be understood about the role of the rainforest for the evolution and future of the trans-equatorial Palaearctic-African bird migration systems.

Movements of immature European Honey Buzzards Pernis apivorus in tropical Africa

Immature European Honey Buzzards Pernis apivorus are believed to remain in tropical Africa during the first years of their lives. We studied their movements during this period with the aid of satellite telemetry. After crossing the Sahara Desert on autumn migration, all six tracked young buzzards stopped at relatively northerly latitudes, between 9.9–13.6°N. Of the five individuals that continued transmitting, four made south-directed movements, mainly in November, to areas located further south or east within latitudes 1.7–9.8°N. Three young buzzards were tracked for more than three months in tropical Africa, and these individuals continued to perform extensive movements within the tropics throughout the tracking period. They travelled between 2,430 and 3,990 km (minimum distances) during 13 to 14 months, in which they visited several sites. In contrast, adult birds migrate directly to their wintering sites where they remain stationary within restricted territories. The mobile life of young Honey Buzzards during the period prior to their first northbound migration may be associated with responses to seasonal weather changes in the tropics and prospecting behaviour. These movements may also reflect intraspecific competition which might be catalyzed by forest degradation and fragmentation.