Optimizing the trade-off between offspring number and quality in unpredictable environments: testing the role of differential androgen transfer to collared flycatcher eggs.

According to the brood reduction hypothesis, parents adjust their brood size in response to current environmental conditions. When resources are abundant, parents can successfully raise all hatched offspring, but when resources are scarce, brood reduction, i.e., the sacrifice of some siblings to secure the quality of a subset of offspring, may maximize fitness. Differential transfer of maternal androgens is one potential proximate mechanism through which female birds may facilitate brood reduction because it may alter the relative competitive ability of sibling nestlings. We tested the hypothesis that female collared flycatchers (Ficedula albicollis) manipulate sibling competition by transferring less androgens to eggs late in the laying sequence. We experimentally elevated androgen levels in i) whole clutches and ii) only the two last laid eggs, and compared growth and begging behavior of offspring from these treatments with a control treatment. By using three treatments and video assessment of begging, we examined the effects of within-clutch patterns of yolk androgen transfer on levels of sibling competition in situ. When androgens were elevated in only the two last laid eggs, begging was more even among siblings compared to control nests. We also found that female nestlings receiving additional yolk androgens showed higher mass gain later in the breeding season, while their male counterparts did not. Our results suggest that females may improve reproductive success in unpredictable environments by altering within-clutch patterns of yolk androgen transfer. We discuss the possibility that life-history divergence between the co-occurring collared and pied flycatcher (Ficedula hypoleuca) is amplified by patterns of yolk androgen transfer.

Life-history divergence facilitates regional coexistence of competing Ficedula flycatchers.

Regional coexistence of ecologically similar species is facilitated when fluctuations in environmental conditions favor different species at different times or places. However, why species with similar ecology should vary in their response to environmental change is unclear. In this study, we explore the role of a life-history divergence in causing changes in relative fitness across environmental conditions experienced by populations of two closely related Ficedula flycatchers on the Baltic island of Oland, Sweden. We compared patterns of nestling survival between Pied (Ficedula hypoleuca) and Collared (F. albicollis) Flycatchers in relation to two factors known to influence the environment experienced by nestlings: natural variation in their parents' onset of breeding and artificial manipulation of the brood size. Possible differences in the location of the nests (i.e., microhabitat differences) or in habitat use (i.e., feeding patterns) by the adult birds were controlled for by partial cross-fostering of young between the two species. We found that nestling mortality was relatively higher among Collared Flycatchers and that this difference increased with later breeding. Mass gain, which predicted survival probability, of nestling Collared Flycatchers did not respond to the seasonal decline in environmental conditions when they were raised in nests with reduced brood size (i.e., where sibling competition was experimentally reduced). This latter result suggests that the smaller clutch size of Collared Flycatchers reflects an adaptive adjustment to their offspring's higher sensitivity to environmental change. We discuss the possibility that the divergence in life-history traits between the two species represents adaptation to different environments experienced during their recent evolutionary history. We conclude that the survival of nestling Collared Flycatchers is more sensitive to harsh environment and that this is likely to limit where and when the more aggressive Collared Flycatchers are able to displace Pied Flycatchers. Our results provide support for models of species coexistence that emphasize the importance of spatial or temporal heterogeneity in relative fitness or life-history divergence. More precisely, our results demonstrate that variation in life-history adaptations may result in changes in relative fitness of species across environments despite their use of similar resources.

Competition-driven build-up of habitat isolation and selection favoring modified dispersal patterns in a young avian hybrid zone.

Competition-driven evolution of habitat isolation is an important mechanism of ecological speciation but empirical support for this process is often indirect. We examined how an on-going displacement of pied flycatchers from their preferred breeding habitat by collared flycatchers in a young secondary contact zone is associated with (a) access to an important food resource (caterpillar larvae), (b) immigration of pied flycatchers in relation to habitat quality, and (c) the risk of hybridization in relation to habitat quality. Over the past 12 years, the estimated access to caterpillar larvae biomass in the habitat surrounding the nests of pied flycatchers has decreased by a fifth due to shifted establishment possibilities, especially for immigrants. However, breeding in the high quality habitat has become associated with such a high risk of hybridization for pied flycatchers that overall selection currently favors pied flycatchers that were forced to immigrate into the poorer habitats (despite lower access to preferred food items). Our results show that competition-driven habitat segregation can lead to fast habitat isolation, which per se caused an opportunity for selection to act in favor of future "voluntarily" altered immigration patterns and possibly strengthened habitat isolation through reinforcement.

Relative performance of hybrid nestlings in Ficedula flycatchers: a translocation experiment.

Ecological speciation predicts that hybrids should experience relatively low fitness in the local environments of their parental species. In this study, we performed a translocation experiment of nestling hybrids between collared and pied flycatchers into the nests of conspecific pairs of their parental species. Our aim was to compare the performance of hybrids with purebred nestlings. Nestling collared flycatchers are known to beg and grow faster than nestling pied flycatchers under favorable conditions, but to experience higher mortality than nestling pied flycatchers under food limitation. The experiment was performed relatively late in the breeding season when food is limited. If hybrid nestlings have an intermediate growth potential and begging intensity, we expected them to beg and grow faster, but also to experience lower survival than pied flycatchers. In comparison with nestling collared flycatchers, we expected them to beg and grow slower, but to survive better. We found that nestling collared flycatchers indeed begged significantly faster and experienced higher mortality than nestling hybrids. Moreover, nestling hybrids had higher weight and tended to beg faster than nestling pied flycatchers, but we did not detect a difference in survival between the latter two groups of nestlings. We conclude that hybrid Ficedula nestlings appear to have a better intrinsic adaptation to food limitation late in the breeding season compared with nestling collared flycatchers. We discuss possible implications for gene flow between the two species.

Positive feedback between ecological and reproductive character displacement in a young avian hybrid zone.

Character displacement can reduce costly interspecific interactions between young species. We investigated the mechanisms behind divergence in three key traits-breeding habitat choice, timing of breeding, and plumage coloration-in Ficedula flycatchers. We found that male pied flycatchers became expelled from the preferred deciduous habitat into mixed forest as the superior competitor, collared flycatchers, increased in numbers. The peak in food abundance differs between habitats, and the spatial segregation was paralleled by an increased divergence in timing of breeding between the two species. Male pied flycatchers vary from brown to black with brown coloration being more frequent in sympatry with collared flycatchers, a pattern often proposed to result from selection against hybridization, that is, reinforcement. In contrast to this view, we show that brown male pied flycatchers more often hybridize than black males. Male pied flycatcher plumage coloration influenced the territory obtained in areas of co-occurrence with collared flycatchers, and brown male pied flycatchers experienced higher relative fitness than black males when faced with heterospecific competition. We suggest that allopatric divergence in resource defense ability causes a feedback loop at secondary contact where male pied flycatchers with the most divergent strategy compared to collared flycatchers are favored by selection.