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.