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).

Given that soaring birds travel faster with supportive winds or in good thermal soaring conditions, we expect weather conditions en route of migration to explain commonly observed regional and seasonal patterns in the performance of soaring migrants. We used GPS-loggers to track 13 honey buzzards and four Montagu's harriers for two to six migrations each. We determined how tailwinds, crosswinds, boundary layer height (a proxy for thermal convection) and precipitation aff ected hourly speeds, daily distances and daily mean speeds with linear regression models. Honey buzzards mostly travel by soaring while Montagu's harriers supplement soaring with fl apping. Th erefore, we expect that performance of harriers will be less aff ected by weather than for buzzards. Weather conditions explained between 30 and 50% of variation in migration performance of both species. Tailwind had the largest eff ect on hourly speeds, daily mean speeds and daily travel distances. Honey buzzards travelled signifi cantly faster and farther, and Montagu ' s harriers non-signifi cantly faster, under better convective conditions. Honey buzzards travelled at slower speeds and shorter distances in crosswinds, whereas harriers maintained high speeds in crosswinds. Weather conditions varied between regions and seasons, and this variation accounted for nearly all regional and seasonal variation in fl ight performance. Hourly performance was higher than predicted at times when we suspect birds had switched to intermittent or continuous fl apping fl ight, for example during sea-crossings. Th e daily travel distance of Montagu ' s harriers was determined to a signifi cant extent by their daily travel time, which diff ered between regions, possibly also due to weather conditions. We conclude with the implications of our work for studies on migration phenology and we suggest an important role for high-resolution telemetry in understanding migratory behavior across entire migratory journeys.

Summary1. Mid-winter movements of up to several hundreds of kilometres are typical for many migratory bird species wintering in Africa. Unpredictable temporary food concentrations are thought to result in random movements of such birds, whereas resightings and recoveries of marked birds suggest some degree of site fidelity. Only detailed (e.g. satellite) tracking of individual migrants can reveal the relative importance and the causes of site choice flexibility and fidelity. The present study investigates how mid-winter movements of a Palaearctic-African migratory raptor, Montagu's harrier Circus pygargus, in the Sahel of West Africa are related to the availability of food resources. 2. Thirty harriers breeding or hatched in northern Europe were satellite tracked (2005)(2006)(2007)(2008)(2009)). On average, four home ranges, each separated by c. 200 km, were visited during one overwinter stay in the Sahel. Wintering home ranges were similar in size to breeding season home ranges (average over wintering and breeding home range size c. 200 km 2 ), and harriers showed high site fidelity between years. 3. Most preferred habitat types in the Sahel were mosaics of grass-and cropland, indicating similar habitat preferences in both the breeding-and wintering seasons. 4. The main prey of Montagu's harriers in the Sahel were grasshoppers Acrididae. Highest grasshopper numbers in the field occurred at relatively low vegetation greenness [normalized difference vegetation index (NDVI) values 0Á17-0Á27]. We used NDVI as a proxy of food availability for harriers. 5. During their overwinter stay, Montagu's harriers moved in a South-South-western direction between consecutive home ranges. The birds selected areas within the range of NDVI values associated with high grasshopper numbers, thus tracking a 'green belt' of predictable changes in highest grasshopper availability. 6. Contrary to earlier hypotheses of random movements in the Sahelian-wintering quarters, the present study shows that Montagu's harriers visited distinct home ranges, they were sitefaithful and tracked seasonal changes in food availability related to previous rainfall patterns, caused by the shifting Intertropical Convergence Zone. Itinerancy may be the rule rather than an exception among insectivorous birds wintering in African savannahs.

Knowledge about migratory connectivity, the degree to which individuals from the same breeding site migrate to the same wintering site, is essential to understand processes affecting populations of migrants throughout the annual cycle. Here, we study the migration system of a long-distance migratory bird, the Montagu's harrier Circus pygargus, by tracking individuals from different breeding populations throughout northern Europe. We identified three main migration routes towards wintering areas in subSaharan Africa. Wintering areas and migration routes of different breeding populations overlapped, a pattern best described by 'weak (diffuse) connectivity'. Migratory performance, i.e. timing, duration, distance and speed of migration, was surprisingly similar for the three routes despite differences in habitat characteristics. This study provides, to our knowledge, a first comprehensive overview of the migration system of a Palaearctic-African long-distance migrant. We emphasize the importance of spatial scale (e.g. distances between breeding populations) in defining patterns of connectivity and suggest that knowledge about fundamental aspects determining distribution patterns, such as the among-individual variation in mean migration directions, is required to ultimately understand migratory connectivity. Furthermore, we stress that for conservation purposes it is pivotal to consider wintering areas as well as migration routes and in particular stopover sites.

Summary 1.Hundreds of millions of Afro-Palaearctic migrants winter in the Sahel, a semi-arid belt south of the Sahara desert, where they experience deteriorating ecological conditions during their overwintering stay and have to prepare for spring migration when conditions are worst. This well-known phenomenon was first described by R.E. Moreau and is known ever since as Moreau's Paradox. However, empirical evidence of the deteriorating seasonal ecological conditions is limited and little is known on how birds respond. 2. Montagu's Harriers Circus pygargus spend 6 months of the year in their wintering areas in the Sahel. Within the wintering season, birds move gradually to the south, visiting several distinct sites to which they are site-faithful in consecutive years. At the last wintering site, birds find themselves at the southern edge of the Sahelian zone and have no other options than facing deteriorating conditions. 3. We tracked 36 Montagu's Harriers with GPS trackers to study their habitat use and behaviour during winter and collected data on the abundance of their main prey, grasshoppers, in Senegal. Since grasshopper abundance was positively related to vegetation greenness (measured as normalized difference vegetation index, NDVI), we used NDVI values as a proxy for prey abundance in areas where no field data were collected. Prey abundance (grasshopper counts and vegetation greenness) at wintering sites of Montagu's Harriers decreased during the wintering period. 4. Montagu's Harriers responded to decreasing food availability by increasing their flight time during the second half of the winter. Individuals increased flight time more in areas with stronger declines in NDVI values, suggesting that lower food abundance required more intense foraging to achieve energy requirements. The apparent consequence was that Montagu's Harriers departed later in spring when their final wintering site had lower NDVI values and presumably lower food abundance and consequently arrived later at their breeding site. 5. Our results confirmed the suggestions Moreau made 40 years ago: the late wintering period might be a bottleneck during the annual cycle with possible carry-over effects to the breeding season. Ongoing climate change with less rainfall in the Sahel region paired with increased human pressure on natural and agricultural habitats resulting in degradation and desertification is likely to make this period more demanding, which may negatively impact populations of migratory birds using the Sahel.