Sympatric divergence and clinal variation in multiple coloration traits of Ficedula flycatchers.

Geographic variation in phenotypes plays a key role in fundamental evolutionary processes such as local adaptation, population differentiation and speciation, but the selective forces behind it are rarely known. We found support for the hypothesis that geographic variation in plumage traits of the pied flycatcher Ficedula hypoleuca is explained by character displacement with the collared flycatcher Ficedula albicollis in the contact zone. The plumage traits of the pied flycatcher differed strongly from the more conspicuous collared flycatcher in a sympatric area but increased in conspicuousness with increasing distance to there. Phenotypic differentiation (PST ) was higher than that in neutral genetic markers (FST ), and the effect of geographic distance remained when statistically controlling for neutral genetic differentiation. This suggests that a cline created by character displacement and gene flow explains phenotypic variation across the distribution of this species. The different plumage traits of the pied flycatcher are strongly to moderately correlated, indicating that they evolve non-independently from each other. The flycatchers provide an example of plumage patterns diverging in two species that differ in several aspects of appearance. The divergence in sympatry and convergence in allopatry in these birds provide a possibility to study the evolutionary mechanisms behind the highly divergent avian plumage patterns.

Natal dispersal based on past and present environmental phenology in the pied flycatcher (Ficedula hypoleuca).

Natal dispersal allows individuals to reach suitable breeding sites. The effect of present plant phenology as a cue for dispersal into areas with favourable stages of development has been well established across avian and mammalian taxa. However, the effect of past experience is less understood. We studied the effect of past and present phenology of the environment on the direction and distance of natal dispersal in a passerine bird, the pied flycatcher (Ficedula hypoleuca). We monitored spring settlement of local recruits in six nest box plots along a 10-km stretch of a south-north gradient of plant and caterpillar food development. We found that males used both past experience of caterpillar phenology from early life and actual plant phenology during the recruitment season as independent cues for breeding settlement. Males that had experienced a mismatch with the caterpillar food peak as a nestling, and/or those that arrived late in the spring in the recruitment year, moved north of their natal site, whereas males that had experienced a better match with the caterpillars as a nestling, and/or those that migrated earlier in the spring, settled at a similar site or more to the south. In females, no such effects were found, suggesting that the usage of phenological cues is sex specific. In summary, tracking environmental phenology by natal dispersal may represent an effective mechanism for settling in new favourable areas, and may thus potentially cause rapid change of a species' geographical breeding range in response to climate change.

Flycatcher song in allopatry and sympatry--convergence, divergence and reinforcement.

The theory of reinforcement predicts that natural selection against the production of unfit hybrids favours traits that increase assortative mating. Whether culturally inherited traits, such as bird song, can increase assortative mating by reinforcement is largely unknown. We compared songs of pied (Ficedula hypoleuca) and collared flycatchers (F. albicollis) from two hybrid zones of different ages with songs from allopatric populations. Previously, a character divergence in male plumage traits has been shown to reinforce premating isolation in sympatric flycatchers. In contrast, we find that the song of the pied flycatcher has converged towards that of the collared flycatcher (mixed singing). However, a corresponding divergence in the collared flycatcher shows that the species differences in song characters are maintained in sympatry. Genetic analyses suggest that mixed song is not caused by introgression from the collared flycatcher, but rather due to heterospecific copying. Circumstantial evidence suggests that mixed song may increase the rate of maladaptive hybridization. In the oldest hybrid zone where reinforcement on plumage traits is most pronounced, the frequency of mixed singing and hybridization is also lowest. Thus, we suggest that reinforcement has reduced the frequency of mixed singing in the pied flycatcher and caused a divergence in the song of the collared flycatcher. Whether a culturally inherited trait promotes or opposes speciation in sympatry may depend on its plasticity. The degree of plasticity may be genetically determined and accordingly under selection by reinforcement.