Competition, Coexistence and Character Displacement : In a Young Avian Hybrid Zone

Abstract: This thesis investigates the ecological and evolutionary implications of a recent secondary contact between two closely related bird species: collared (Ficedula albicollis) and pied (F. hypoleuca) flycatchers. Collared flycatchers started to colonize the Swedish island of Öland, where pied flycatchers were already present, in the late 1950s-early1960s. My major aims were to investigate which factors are acting against versus for long-term coexistence between the two species. Specifically, I investigated the relative importance of allopatric divergence, interspecific competition, hybridization and learning in promoting or inhibiting coexistence. The combined effects of interspecific competition and hybridization drives pied flycatchers towards local extinction in their preferred deciduous habitat. However, my results also show that pied flycatchers are better able to tolerate harsh environmental conditions. This trade-off between competitive ability and resilience in the face of harsh conditions facilitates a regional coexistence between the species. Coexistence is furthermore favoured by competition-mediated divergence in breeding habitat choice, timing of breeding and male breeding plumage colouration. Due to interspecific competition, male pied flycatchers are forced to breed in a more mixed forest type with a later peak in food abundance, which is accompanied by a divergence in breeding time between the two species. In areas shared with collared flycatchers, male pied flycatchers with brown plumage coloration, most divergent from that of collared flycatchers, are favoured by selection. In addition to facilitating coexistence, the observed shift in habitat occupancy increases reproductive isolation between the two species. By using cross-fostering experiments I demonstrate that natal habitat imprinting has the potential to additionally speed up habitat segregation. Finally I show that hybrid nestlings express an intermediate response to harsh environments, indicating that another aspect of ecological-based selection may be important in reproductive isolation between the species. In summary, my results show that adaptations during historic allopatry are important both in facilitating coexistence as well as in providing a foundation for further ecological divergence at secondary contact. This is of relevance today as many species are shifting their distributions in response to habitat disturbance and global warming.