The evolutionary history of "suboptimal" migration routes.

Migratoriness in birds is evolutionary labile, with many examples of increasing or decreasing migration distances on the timescale of modern ornithology. In contrast, shifts of migration to more nearby wintering grounds seem to be a slow process. We examine the history of how Palearctic migratory landbirds have expanded their wintering ranges to include both tropical Africa and Asia, a process that has involved major shifts in migratory routes. We found that species with shorter migration distances and with resident populations in the Palearctic more often winter in both Africa and Asia. Our results suggest that changes in wintering grounds are not by long-distance migrant populations per se, but through historic intermediate populations that were less migratory from which long-distance migration evolved secondarily. The failure of long-distance migrants to shift migration direction to more nearby winter quarters indicates that major modifications to the migratory program may be difficult to evolve.

Diurnal migration patterns in willow warblers differ between the western and eastern flyways.

It is a long-standing view that the main mechanism maintaining narrow migratory divides in passerines is the selection against intermediate and suboptimal migratory direction, but empirical proof of this is still lacking. We present novel results from a willow warbler migratory divide in central Sweden from where birds take the typical SW and SE as well as intermediate routes to winter quarters in Africa. We hypothesized that individuals that take the intermediate route are forced to migrate in daytime more often when crossing wide ecological barriers than birds that follow the typical western or eastern flyways. Analyses of geolocator tracks of willow warblers breeding across the entire Sweden, including the migratory divide, provided no support for our hypothesis. Instead, birds that migrated along the western flyway were the most likely to undertake full day flights. The probability of migrating for a full day when crossing major barriers declined linearly from west to east. We speculate that this difference is possibly caused by more challenging conditions in the western part of the Sahara Desert, such as the lack of suitable day-time roost sites. However, it may equally likely be that willow warblers benefit from migrating in daytime if favorable tailwinds offer assistance.

Flyways and migratory behaviour of the Vega gull (Larus vegae), a little-known Arctic endemic.

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.

Migration direction in a songbird explained by two loci.

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.

Transposable elements mark a repeat-rich region associated with migratory phenotypes of willow warblers (Phylloscopus trochilus).

The genetic basis of bird migration has been the focus of several studies. Two willow warbler subspecies (Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) follow different migratory routes to wintering grounds in Africa. Their breeding populations overlap in contact areas or "migratory divides" located in central Scandinavia and in eastern Poland. Earlier analyses demonstrated that the genetic differences between these two migratory phenotypes are few and cluster on chromosomes 1 and 5. In addition, an amplified fragment length polymorphism-derived biallelic marker (known as WW2) presents steep clines across both migratory divides but failed to be mapped in the genome. Here, we characterize the WW2 marker and describe its two variants (WW2 ancestral and WW2 derived) as portions of long terminal repeat retrotransposons originating from an ancient infection by an endogenous retrovirus. We used quantitative polymerase chain reaction techniques to quantify copy numbers of the WW2 derived variant in the two subspecies and their hybrids. This, together with genome analyses revealed that WW2 derived variants are much more abundant in P. t. acredula and appear embedded in a large repeat-rich region (>12 Mbp), not associated with the divergent regions of chromosomes 1 or 5. However, it might interact with genetic elements controlling migration direction. Testing this hypothesis further will require knowing the exact location of this region, such as by obtaining more complete genome assemblies preferably in combination with techniques like fluorescence in situ hybridization applied to a willow warbler karyotype, and finally to investigate the copy number of this marker in hybrids with known migratory tracks.

Autumn migration direction of juvenile willow warblers (Phylloscopus t. trochilus and P. t. acredula) and their hybrids assessed by qPCR SNP genotyping.

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 well-established 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. CONCLUSIONS: Our genetic approach to assign individuals to subspecies and hybrids allowed us to investigate migration direction in juvenile birds on their first migration, which should better reflect the innate migratory direction than studies restricted to successful migrants. The excess of hybrids in Italy, suggests that they employ an intermediate route relative to the parental populations. Our qPCR SNP genotyping method is efficient for processing large sample sizes, and will therefore be useful in migration research of species with known population genetic structure.

Dominance relationships and coalitionary aggression against conspecifics in female carrion crows.

Cooperation is a prevailing feature of many animal systems. Coalitionary aggression, where a group of individuals engages in coordinated behaviour to the detriment of conspecific targets, is a form of cooperation involving complex social interactions. To date, evidence has been dominated by studies in humans and other primates with a clear bias towards studies of male-male coalitions. We here characterize coalitionary aggression behaviour in a group of female carrion crows consisting of recruitment, coordinated chase, and attack. The individual of highest social rank liaised with the second most dominant individual to engage in coordinated chase and attack of a lower ranked crow on several occasions. Despite active intervention by the third most highly ranked individual opposing the offenders, the attack finally resulted in the death of the victim. All individuals were unrelated, of the same sex, and naïve to the behaviour excluding kinship, reproduction, and social learning as possible drivers. Instead, the coalition may reflect a strategy of the dominant individual to secure long-term social benefits. Overall, the study provides evidence that members of the crow family engage in coordinated alliances directed against conspecifics as a possible means to manipulate their social environment.

Ten grams and 13,000 km on the wing - route choice in willow warblers Phylloscopus trochilus yakutensis migrating from Far East Russia to East Africa.

BACKGROUND: High-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. RESULTS: The 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. CONCLUSIONS: The 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.