1. Light-level geolocator tags use ambient light recordings to estimate the whereabouts of an individual over the time it carried the device. Over the past decade, these tags have emerged as an important tool and have been used extensively for tracking animal migrations, most commonly small birds.

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Understanding what drives or prevents long-distance migrants to respond to environmental change requires basic knowledge about the wintering and breeding grounds, and the timing of movements between them. Both strong and weak migratory connectivity have been reported for Palearctic passerines wintering in Africa, but this remains unknown for most species. We investigated whether pied flycatchers Ficedula hypoleuca from different breeding populations also differ in wintering locations in west-Africa. Light-level geolocator data revealed that flycatchers from different breeding populations travelled to different wintering sites, despite similarity in routes during most of the autumn migration. We found support for strong migratory connectivity showing an unexpected pattern: individuals breeding in Fennoscandia (S-Finland and S-Norway) wintered further west compared to individuals breeding at more southern latitudes in the Netherlands and SW-United Kingdom. The same pattern was found in ring recovery data from sub-Saharan Africa of individuals with confirmed breeding origin. Furthermore, population-specific migratory connectivity was associated with geographical variation in breeding and migration phenology: birds from populations which breed and migrate earlier wintered further east than birds from 'late' populations. There was no indication that wintering locations were affected by geolocation deployment, as we found high repeatability and consistency in d 13 C and d 15 N stable isotope ratios of winter grown feathers of individuals with and without a geolocator. We discuss the potential ecological factors causing such an unexpected pattern of migratory connectivity. We hypothesise that population differences in wintering longitudes of pied flycatchers result from geographical variation in breeding phenology and the timing of fuelling for spring migration at the wintering grounds. Future research should aim at describing how temporal dynamics in food availability across the wintering range affects migration, wintering distribution and populations' capacity to respond to environmental changes.

Summary1. Properly timed spring migration enhances reproduction and survival. Climate change requires organisms to respond to changes such as advanced spring phenology. Pied flycatchers Ficedula hypoleuca have become a model species to study such phenological adaptations of long-distance migratory songbirds to climate change, but data on individuals' time schedules outside the breeding season are still lacking. 2. Using light-level geolocators, we studied variation in migration schedules across the year in a pied flycatcher population in the Netherlands, which sheds light on the ability for individual adjustments in spring arrival timing to track environmental changes at their breeding grounds. 3. We show that variation in arrival dates to breeding sites in 2014 was caused by variation in departure date from sub-Saharan Africa and not by environmental conditions encountered en route. Spring migration duration was short for all individuals, on average 2 weeks. Males migrated ahead of females in spring, while migration schedules in autumn were flexibly adjusted according to breeding duties. Individuals were therefore not consistently early or late throughout the year. 4. In fast migrants like our Dutch pied flycatchers, advancement of arrival to climate change likely requires changes in spring departure dates. Adaptation for earlier arrival may be slowed down by harsh circumstances in winter, or years with high costs associated with early migration.

1. Currently, the deployment of tracking devices is one of the most frequently used approaches to study movement ecology of birds. Recent miniaturization of lightlevel geolocators enabled studying small bird species whose migratory patterns were widely unknown. However, geolocators may reduce vital rates in tagged birds and may bias obtained movement data.2. There is a need for a thorough assessment of the potential tag effects on small birds, as previous meta-analyses did not evaluate unpublished data and impact of multiple life-history traits, focused mainly on large species and the number of published studies tagging small birds has increased substantially.

Each year more than two billion songbirds cross the Sahara, but how they perform this formidable task is largely unknown. Using geolocation tracks from 27 pied flycatchers, a nocturnally migrating passerine, we show that most birds made diurnal flights in both autumn and spring. These diurnal flights were estimated to be part of non-stop flights of mostly 40 -60 h. In spring, birds flew across the Sahara, while autumn migration probably circumpassed part of the desert, through a long oversea flight. Our data contradict claims that passerines cross the Sahara by intermittent flight and daytime resting. The frequent occurrence of long non-stop flights to cross the desert shows migrants' physiological abilities and poses the question why this would not be the general migration strategy to cross the Sahara.

Avian breeding populations have been shown to be regulated by territorial behaviour, often creating a surplus of non-breeding individuals. However, most evidence is of a male non-breeder surplus, whereas for a surplus to actually buffer a population both non-breeding males and females should be present. Here, we provide descriptive and experimental evidence for the existence of a population buffer consisting of mostly male and potentially also female Pied Flycatchers using nest box areas. First we show that local recruits often do not breed in their first year, with 23% of all recruiting males observed breeding in their first year, and 51% of females. When accounting for mortality in the years prior to observed first breeding, we estimate that only 9% of all first-year males breed locally, and 29% of first-year females. Similar percentages of first-year flycatchers skipping breeding have been observed in other study populations. We show that in the year of new establishment of our nest box plots, most known-aged flycatchers were first-year birds (77%), whereas after establishment, recruiting immigrants from the same source population were mostly older (28% first-year birds). An experimental removal of paired flycatchers from one study plot in two years (19 and 58 individuals removed) resulted in complete replacement by males and females. Male but not female replacements were younger than removed individuals. These results imply that a non-breeding surplus is present in Pied Flycatcher populations. The average later age at firstbreeding in males compared to females, suggests that this non-breeding surplus is strongly male-biased. Skipping breeding in the first year(s) is not just caused by shortage of suitable nesting sites, as we observed on average 12% of males defending a nest box without pairing up with a female. Using stable isotopes ratios, we show that non-breeding first-year individuals do not stay at their African wintering grounds. Competition for nest sites is one cause for refraining from breeding, as shown by our experiments, but cannot be the sole cause, as many nest boxes remain unused in a season, and up to 20% of territorial males defend a nest box without pairing up with a female. We hypothesize that many young flycatchers arrive too late for breeding and are therefore not seen in their first year. Indeed first-year Pied Flycatchers that do breed/defend a nest box arrive on average later at the breeding grounds, and we argue that the nonobserved group arrives even later. The causes of their later arrival could be the need for learning, lower quality wintering sites resulting in later departure, and/or a trade-off between low breeding success and the costs of early arrival. These could be general factors in long-distance migrants, and this pleads for a better understanding of how migration develops during ontogeny.

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