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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