Susanne Åkesson

Wind selectivity of migratory flight departures in birds

Abstract Optimal migration theory predicts that birds minimizing the overall time of migration should adjust stopover duration with respect to the rate of fuel accumulation. Recent theoretical developments also take into account the wind situation and predict that there is a time window (a set of days) during which birds should depart when assisted by winds but will not do so if there are head winds. There is also a final day when birds will depart irrespective of wind conditions. Hence, the wind model of optimal migration theory predicts that birds should be sensitive to winds and that there should be a correlation between departures and winds blowing towards the intended migration direction. We tested this assumption by tracking the departures of radio-tagged passerines during autumn migration in southern Sweden. Our birds were moderately to very fat when released and therefore energetically ready for departure. There was a significant correlation between direction of departure and wind direction. We also found that during days when birds departed there was a significantly larger tail wind component than during days when birds were present but did not depart. Our results show that passerines do take the current wind situation into account when departing on migratory flights. We also briefly discuss possible clues that birds use when estimating wind direction and strength. The inclusion of wind is an important amendment to optimal migration theory of birds and should be explored further.

How Migrants Get There: Migratory Performance and Orientation

ABSTRACT Migratory animals show a suite of adaptations to cope with their journeys. These include not only morphological features for efficient locomotion and storage of energy but also behavioral adjustments to exploit winds and currents or to avoid drift caused by moving fluids. Migration strategies across locomotory modes can be analyzed in the context of optimality models, using some general principles concerning migration range and selection criteria. Comparisons of model predictions with natural behavior help researchers understand the selection pressures that underlie migration strategies. We give examples of typical migration speeds and distances for animals using different locomotion models. Successful migration also requires accurate orientation and/or navigation between distant areas for reproduction and survival. Animals can use a suite of different compasses, which may be cross-calibrated or integrated for direction finding, depending on the geographical and ecological situation, and may be used with an endogenous clock for time compensation.

MORPHOLOGICAL AND MOLECULAR VARIATION ACROSS A MIGRATORY DIVIDE IN WILLOW WARBLERS, PHYLLOSCOPUS TROCHILUS

A migratory divide is a narrow region in which two populations showing different migratory directions meet arid presumably also mate and hybridize. Banding of willow warblers, Phylloscopus trochilus, in Europe has demonstrated a migratory divide latitudinally across central Scandinavia. In autumn, southern birds migrate southwest to tropical West Africa, whereas northern birds migrate southeast to East and South Africa. The migratory divide is associated with concordant differences in size and plumage coloration. Based on morphology, we estimate the width of the transition zone between northern and southern willow warblers to be less than 350 km. We found indication of linkage disequilibria around the migratory divide, in that measures of body size were correlated with plumage coloration within the contact zone, but uncorrelated within the populations south or north of the contact zone. The presence of linkage disequilibria and the fact that several morphological clines occur together suggest that the hybrid zone is a result of secondary contact between populations that have differentiated in allopatry. This interpretation is in accord with the knowledge of the recolonization pattern of the Scandinavian peninsula after the last glaciation; animals and plants appeared to have colonized either from the south or from the north around the northern bay of the Baltic Sea. If northern and southern willow warblers resided in allopatric populations during late Pleistocene glaciations and the hybrid zone is a result of postglacial range expansions, we would expect some degree of genetic differentiation accumulated during the period in isolation. In contrast, northern and southern willow warblers are near panmictic in the frequencies of alleles of mitochondrial DNA and at two microsatellite loci. The observed pattern, clear morphological and behavioral differentiation without genetic differentiation at neutral loci, suggests either that the differences are maintained by strong selection on the expressed genes in combination with high levels of current gene flow or, in the case of weak gene flow, that the divergence in morphology and behavior is very recent.

Stable isotopes examined across a migratory divide in Scandinavian willow warblers (Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) reflect their African winter quarters

The C and N isotopes of feathers from two subspecies of willow warblers (Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) are isotopically distinct. Our analysis of 138 adult males from sites distributed across Sweden shows that the mean δ15N and δ13C values of subspecies acredula (from latitudes above 63° N) were significantly higher than the mean δ15N and δ13C values of subspecies trochilus (from latitudes below 61° N). The analysed willow warbler feathers had been moulted in the winter quarters and the observed isotopic signatures should thus reflect the isotopic pattern of food assimilated in Africa. The isotopic data observed in Sweden match the cline in morphology, both showing abrupt changes around 62° N.This result agrees with data from ringing recoveries indicating that the two subspecies occupy geographically and isotopically distinct wintering grounds in Africa. Our isotopic data suggest that analysis of stable isotopes of C and N is a promising method to track wintering quarters of European birds that migrate to Africa.

The genetics of migration on the move.

Across a range of organisms, related species or even populations of the same species exhibit strikingly different scales and patterns of movement. A significant proportion of the phenotypic variance in migratory traits is genetic, but the genes involved in shaping these phenotypes are still unknown. Although recent achievements in genomics will evolve migratory genetics research from a phenotypic to a molecular approach, fully sequenced and annotated genomes of migratory species are still lacking. Consequently, many of the genes involved in migration are unavailable as candidates. Migration is central to the life-history adaptations of many animals. Here, we review current understanding of the genetic architecture of migratory traits and discuss the significant implications this will have for other areas of biology, including population responses to climate change, speciation and conservation management.

Grand Challenges in Migration Biology

Abstract Billions of animals migrate each year. To successfully reach their destination, migrants must have evolved an appropriate genetic program and suitable developmental, morphological, physiological, biomechanical, behavioral, and life-history traits. Moreover, they must interact successfully with biotic and abiotic factors in their environment. Migration therefore provides an excellent model system in which to address several of the “grand challenges” in organismal biology. Previous research on migration, however, has often focused on a single aspect of the phenomenon, largely due to methodological, geographical, or financial constraints. Integrative migration biology asks ‘big questions’ such as how, when, where, and why animals migrate, which can be answered by examining the process from multiple ecological and evolutionary perspectives, incorporating multifaceted knowledge from various other scientific disciplines, and using new technologies and modeling approaches, all within the context of an annual cycle. Adopting an integrative research strategy will provide a better understanding of the interactions between biological levels of organization, of what role migrants play in disease transmission, and of how to conserve migrants and the habitats upon which they depend.

Assessing accuracy and utility of satellite-tracking data using Argos-linked Fastloc-GPS

Centre for Ecology and Conservation, School of Biosciences, University of Exeter Department of Animal Ecology, Lund University, Sweden c SEATURTLE.org, U.S.A. Ascension Island Turtle Group, Ascension Island, South Atlantic Wildlife Conservation Society, Gabon f Institute of Environmental Sustainability, Swansea University, U.K. Dipartimento di Biologia, University of Pisa, Italy Ascension Island Conservation, Ascension Island, South Atlantic

The use of AFLP to find an informative SNP: genetic differences across a migratory divide in willow warblers

We used the amplified fragment length polymorphism (AFLP) method to obtain genetic markers distinguishing two subspecies of willow warblers Phylloscopus trochilus that have different migratory behaviours but are not differentiated in mitochondrial DNA or at several microsatellite loci. With the inverse‐polymerase chain reaction (PCR) approach we converted a dominant AFLP‐marker to a codominant single nucleotide polymorphism (SNP). Across Scandinavia we typed 621 birds at the SNP locus AFLP‐WW1 and we found a sigmoid change in allele frequencies centred around 62 degrees latitude. North of the latitudinal cline was a west‐east cline. Both clines are narrower than one would expect from dispersal distances in willow warblers, which suggests that these are maintained by selection. The latitudinal cline at the locus AFLP‐WW1 is paralleled by changes in several other traits, all of which might be maintained by a single selective force. The most plausible selection factor that we have identified is selection against hybrids because of inferior migratory behaviour. The selective force maintaining the east–west cline is less obvious. We discuss alternatives to the selection scenario, involving colonization history and asymmetric gene flow.

Bimodal orientation and the occurrence of temporary reverse bird migration during autumn in south Scandinavia

Abstract Extensive ringing data from a coastal site (Falsterbo Bird Observatory) in southwesternmost Sweden were used to investigate the occurrence of reverse autumn migration among 20 passerine bird species of widely different migration categories. The data demonstrate that reverse migration is a widespread and regular phenomenon among nocturnal as well as diurnal migrants and among irruptive migrants, temperate zone migrants, and long-distance migrants destined for tropical winter quarters. The reoriented movements were directed approximately opposite to the normal migration direction, i.e. between NNW and ENE from the coast and towards inland. Median distances of reverse movements varied between 9 and 65 km. Some individuals of irruptive and partial migrants settled to winter in the reverse direction. Bird species with relatively small fat reserves at capture were more likely to perform reverse migratory movements than species with larger fat deposits. In two species birds performing forward migration were significantly heavier within 10 days after capture than individuals performing reverse movements. The reoriented movements probably are of adaptive significance for birds confronted with the sea and pre-disposed to refuelling during migration. A bimodal orientation mechanism will bring the birds from an area with high competition for food and high predation risk to more suitable resting and feeding grounds before resuming migration in the forward direction and crossing the barrier.

Conflicting patterns of mitochondrial and nuclear DNA diversity in Phylloscopus warblers

Molecular variation is often used to infer the demographic history of species, but sometimes the complexity of species history can make such inference difficult. The willow warbler, Phylloscopus trochilus, shows substantially less geographical variation than the chiffchaff, Phylloscopus collybita, both in morphology and in mitochondrial DNA (mtDNA) divergence. We therefore predicted that the willow warbler should harbour less nuclear DNA diversity than the chiffchaff. We analysed sequence data obtained from multiple samples of willow warblers and chiffchaffs for the mtDNA cytochrome b gene and four nuclear genes. We confirmed that the mtDNA diversity among willow warblers is low (π = 0.0021). Sequence data from three nuclear genes (CHD‐Z, AFLP‐WW1 and MC1R) not linked to the mitochondria demonstrated unexpectedly high nucleotide diversity (π values of 0.0172, 0.0141 and 0.0038) in the willow warbler, on average higher than the nucleotide diversity for the chiffchaff (π values of 0.0025, 0.0017 and 0.0139). In willow warblers, Tajimas D analyses showed that the mtDNA diversity, but not the nuclear DNA diversity, has been reduced relative to the neutral expectation of molecular evolution, suggesting the action of a selective sweep affecting the maternally inherited genes. The large nuclear diversity seen within willow warblers is not compatible with processes of neutral evolution occurring in a population with a constant population size, unless the long‐term effective population size has been very large (Ne > 106). We suggest that the contrasting patterns of genetic diversity in the willow warbler may reflect a more complex evolutionary history, possibly including historical demographic fluctuations or historical male‐biased introgression of nuclear genes from a differentiated population of Phylloscopus warblers.