BackgroundHigh-latitude bird migration has evolved after the last glaciation, in less than 10,000–15,000 years. Migrating songbirds rely on an endogenous migratory program, encoding timing, fueling, and routes, but it is still unknown which compass mechanism they use on migration. We used geolocators to track the migration of willow warblers (Phylloscopus trochilus yakutensis) from their eastern part of the range in Russia to wintering areas in sub-Saharan Africa. Our aim was to investigate if the autumn migration route can be explained by a simple compass mechanism, based on celestial or geomagnetic information, or whether migration is undertaken as a sequence of differential migratory paths possibly involving a map sense. We compared the recorded migratory routes for our tracked birds with simulated routes obtained from different compass mechanisms.ResultsThe three tracked males were very similar in the routes they took to their final wintering sites in southern Tanzania or northern Mozambique, in their use of stopover sites and in the overall timing of migration. None of the tested compass mechanisms could explain the birds’ routes to the first stopover area in southwest Asia or to the destination in Southeast Africa without modifications. Our compass mechanism simulations suggest that the simplest scenarios congruent with the observed routes are based on either an inclination or a sun compass, assuming two sequential steps.ConclusionsThe birds may follow a magnetoclinic route coinciding closely with the tracks by first moving west, i.e. closer to the goal, and thereafter follow a constant apparent angle of inclination to the stopover site. An alternative would be to use the sun compass, but with time-adjustments along the initial part of the migration to the first stopover, and thereafter depart along a new course to the winter destination. A combination of the two mechanisms cannot be ruled out, but needs to be confirmed in future studies.Electronic supplementary materialThe online version of this article (10.1186/s40462-018-0138-0) contains supplementary material, which is available to authorized users.

Migratory routes and remote wintering quarters in birds are often species and even population specific. It has been known for decades that songbirds mainly migrate solitarily, and that the migration direction is genetically controlled. Yet, the underlying genetic mechanisms remain unknown. To investigate the genetic basis of migration direction, we track genotyped willow warblers Phylloscopus trochilus from a migratory divide in Sweden, where South-West migrating, and South-East migrating subspecies form a hybrid swarm. We find evidence that migration direction follows a dominant inheritance pattern with epistatic interaction between two loci explaining 74% of variation. Consequently, most hybrids migrate similarly to one of the parental subspecies, and therefore do not suffer from the cost of following an inferior, intermediate route. This has significant implications for understanding the selection processes that maintain narrow migratory divides.

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Backgrounds: Geographic regions, where two closely related taxa with different migration routes come into contact, are known as migratory divides. Hybrids originating from migratory divides are hypothesized to migrate intermediately relative to the parental populations. Few studies have tested this hypothesis in wild birds, and only in hybrids that have completed the migration back to the breeding grounds. Here, we make use of the wellestablished migration routes of willow warblers (Phylloscopus trochilus), for which the subspecies trochilus and acredula have migration-associated genetic markers on chromosomes 1 and 5. The genetic approach enabled us to analyze the geographic distribution of juveniles during their first autumn migration, predicting that hybrids should be more frequent in the central flyway over Italy than along the typical SW routes of trochilus and SE routes of acredula. Methods: Blood and feather samples were collected from wintering birds in Africa (n = 69), and from juveniles during autumn migration in Portugal (n = 33), Italy (n = 38) and Bulgaria (n = 32). Genotyping was carried out by qPCR SNP assays, on one SNP each on chromosome 1 (SNP 65) and chromosome 5 (SNP 285). Both these SNPs have alternative alleles that are highly fixed (> 97%) in each of the subspecies. Results: The observed combined genotypes of the two SNPs were associated with the known migration routes and wintering distributions of trochilus and acredula, respectively. We found hybrids (HH) among the juveniles in Italy (5/38) and in Portugal (2/33). The proportion of hybrids in Italy was significantly higher than expected from a background rate of hybrid genotypes (1.5%) in allopatric populations of the subspecies.

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The return of individual birds to a specific area in successional years, i.e. philopatry, is a remarkable behavioural trait. Here we report on the remarkably reversed: the complete absence of returning individuals of a migratory passerine with otherwise pronounced philopatry. At a high latitude study site in Abisko (68°32ʹN, 18°80ʹE) in northern Sweden none of the banded adult willow warblers Phylloscopus trochilus returned to breed 2011–2014. This is in stark contrast to all other reports in the literature and also to our two southern study sites (at 56°56ʹN, 18°10ʹE and at 58°94ʹN, 17°14ʹE) where 18–38% of adults returned. We investigated this aberrant pattern found in Abisko by analysing three parameters known to influence philopatry; nest predation, breeding success and breeding density, and predicted that absence of philopatry should co‐occur with low breeding success, low breeding density and/or high nest predation. The results did not corroborate this, except that breeding density was lower at Abisko (49–71 pairs km–2) than at the southern sites (106 pairs km–2, 101 pairs km–2). Instead, we suggest the hypothesis that the absence of philopatry is caused by an influx of southern, dispersal‐prone individuals deploying another breeding strategy and that this intra‐specific range expansion is enabled by milder climate and low population density.

Large gulls are generalist predators that play an important role in Arctic food webs. Describing the migratory patterns and phenology of these predators is essential to understanding how Arctic ecosystems function. However, from all six large Arctic gull taxa, including three long-distance migrants, to date seasonal movements have been studied only in three and with small sample sizes. To document the flyways and migratory behaviour of the Vega gull, a widespread but little-studied Siberian migrant, we monitored 28 individuals with GPS loggers over a mean period of 383 days. Birds used similar routes in spring and autumn, preferring coastal to inland or offshore routes, and travelled 4000–5500 km between their breeding (Siberia) and wintering grounds (mainly the Republic of Korea and Japan). Spring migration mainly occurred in May, and was twice as fast and more synchronized among individuals than autumn migration. Migration bouts mainly occurred during the day and twilight, but rates of travel were always higher during the few night flights. Flight altitudes were nearly always higher during migration bouts than during other bouts, and lower during twilight than during night or day. Altitudes above 2000m were recorded during migrations, when birds made non-stop inland flights over mountain ranges and vast stretches of the boreal forest. Individuals showed high inter-annual consistency in their movements in winter and summer, indicating strong site fidelity to their breeding and wintering sites. Within-individual variation was similar in spring and autumn, but between individual variation was higher in autumn than in spring. Compared to previous studies, our results suggest that the timing of spring migration in large Arctic gulls is likely constrained by snowmelt at breeding grounds, while the duration of migration windows could be related to the proportion of inland versus coastal habitats found along their flyways (‘fly-and-forage’ strategy). Ongoing environmental changes are hence likely in short term to alter the timing of their migration, and in long term possibly affect the duration if e.g. the resource availability along the route changes in the future.

Long-distance migrants with transcontinental breeding ranges are of particular interest for the study of local adaptation and geographic differentiation in birds. We compared phenotypes and genotypes between Far East Siberian Willow Warblers Phylloscopus trochilus yakutensis Ticehurst, 1935 with the European subspecies P. t. trochilus Linnaeus, 1758 and P. t. acredula Linnaeus, 1758. We found significant differences in mean body size and plumage colour, but intra-population variation overlapped extensively between the European and Siberian populations. We used stable isotope composition in winter-grown flight feathers as a proxy for wintering sites and found differences between all three subspecies, indicating different wintering grounds. Out of four nuclear loci analyzed (three of which are known to be substantially divergent between the European subspecies), none allowed to seperate East Siberian yakutensis from North Scandinavian acredula. Hence, neither phenotypic traits nor the currently available genetic resources provide diagnostic criteria for confidently assigning individual Willow Warblers to a particular subspecies. Despite extensive overlap in phenotypes and genotypes, we propose that the subspecies names can still be used as biogeographical references to the three Willow Warbler populations that differ in migration strategies. We propose to use yakutensis for Willow Warblers breeding east of the Ural Mountains that presumably initiate autumn migration towards the southwest or west, in contrast to the genetically most similar acredula that start autumn migration towards the southeast or south. Future field studies are needed to elucidate whether the longitudinal variation in phenotype is a cline, or whether a clear contact zone between these subspecies can be identified. Keywords Willow Warbler • Phylloscopus trochilus yakutensis • Subspecies • Genetic structure • Phenotypic variation • Clock gene • Stable isotopes Zusammenfassung Vergleichende phänotypische und genetische Charakterisierung der ostsibirischen (Phylloscopus trochilus yakutensis Ticehurst, 1935) und europäischen Fitis-Unterarten Langstreckenzieher mit trans-kontinentalem Verbreitungsgebiet eigenen sich hervorragend zur Untersuchung lokaler Anpassung und geographische Differenzierung. Hier vergleichen wir Phänotyp und Genotyp der ostsibirischem Fitis-Unterart (Phylloscopus trochilus yakutensis Ticehurst, 1935) mit den europäischen Unterarten P. t. trochilus Linnaeus, 1758 und P. t. acredula Linnaeus, 1758. Unsere Ergebnisse zeigen signifikante Unterschiede der mittleren Körpergröße, sowie Gefiederfärbung, wobei die Bandbreite beider Messgrößen zwischen den Unterarten stark überlappen. Die zur Bestimmung der jeweiligen Überwinterungsgebiete genutzte stabile Isotopenkomposition der im Winter gemauserten Communicated by M. Wink.