Competition for mates can play a critical role in determining reproductive success, shaping phenotypic variation within populations, and influencing divergence. Yet, studies of the role of sexual selection in divergence and speciation have focused disproportionately on mate choice. Here, we synthesize the literature on how mate competition may contribute to speciation and integrate concepts from work on sexual selection within populations—mating systems, ecology, and mate choice. Using this synthesis, we generate testable predictions for how mate competition may contribute to divergence. Then, we identify the extent of existing support for these predictions in the literature with a systematic review of the consequences of mate competition for population divergence across a range of evolutionary, ecological, and geographic contexts. We broadly evaluate current evidence, identify gaps in available data and hypotheses that need testing, and outline promising directions for future work. A major finding is that mate competition may commonly facilitate further divergence after initial divergence has occurred, e.g., upon secondary contact and between allopatric populations. Importantly, current hypotheses for how mate competition contributes to divergence do not fully explain observed patterns. While results from many studies fit predictions of negative frequency-dependent selection, agonistic character displacement, and ecological selection, results from ~30% of studies did not fit existing conceptual models. This review identifies future research aims for scenarios in which mate competition is likely important but has been understudied, including how ecological context and interactions between mate choice and mate competition can facilitate or hinder divergence and speciation.
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Abstract Phenotypic differentiation plays an important role in the formation and maintenance of reproductive barriers. In some cases, variation in a few key aspects of phenotype can promote and maintain divergence; hence, the identification of these traits and their associations with patterns of genomic divergence is crucial for understanding the patterns and processes of population differentiation. We studied hybridization between the
alba andpersonata subspecies of the white wagtail (Motacilla alba ), and quantified divergence and introgression of multiple morphological traits and 19,437SNP loci on a 3,000 km transect. Our goal was to identify traits that may contribute to reproductive barriers and to assess how variation in these traits corresponds to patterns of genome‐wide divergence. Variation in only one trait—head plumage patterning—was consistent with reproductive isolation. Transitions in head plumage were steep and occurred over otherwise morphologically and genetically homogeneous populations, whereas cline centres for other traits and genomic ancestry were displaced over 100 km from the head cline. Field observational data show that social pairs mated assortatively by head plumage, suggesting that these phenotypes are maintained by divergent mating preferences. In contrast, variation in all other traits and genetic markers could be explained by neutral diffusion, although weak ecological selection cannot be ruled out. Our results emphasize that assortative mating may maintain phenotypic differences independent of other processes shaping genome‐wide variation, consistent with other recent findings that raise questions about the relative importance of mate choice, ecological selection and selectively neutral processes for divergent evolution. -
Abstract Hybrid zones are geographic regions where isolating barriers between divergent populations are challenged by admixture. Identifying factors that facilitate or inhibit hybridization in sympatry can illuminate the processes that maintain those reproductive barriers. We analysed patterns of hybridization and phenotypic variation across two newly discovered hybrid zones between three subspecies of barn swallow (
Hirundo rustica ). These subspecies differ in ventral coloration and wing length, traits that are targets of sexual and natural selection, respectively, and are associated with genome‐wide differentiation in allopatry. We tested the hypothesis that the degree of divergence in these traits is associated with the extent of hybridization in secondary contact. We applied measures of population structure based on >23,000SNP s to confirm that named subspecies correspond to distinct genomic clusters, and assessed coincidence between geographic clines for ancestry and phenotype. Although gene flow was ongoing across both hybrid zones and pairwiseF STbetween subspecies was extremely low, we found striking differences in the extent of hybridization. In the more phenotypically differentiated subspecies pair, clines for ancestry, wing length and ventral coloration were steep and coincident, suggestive of strong isolation and, potentially, selection associated with phenotype. In the less phenotypically differentiated pair, gene flow and phenotypic variation occurred over a wide geographic span, indicative of weaker isolation. Traits associated with genome‐wide differentiation in allopatry may thus also contribute to isolation in sympatry. We discuss potentially important additional roles for evolutionary history and ecology in shaping variation in the extent hybridization between closely related pairs of subspecies. -
Abstract Migratory divides are proposed to be catalysts for speciation across a diversity of taxa. However, it is difficult to test the relative contributions of migratory behaviour vs. other divergent traits to reproductive isolation. Comparing hybrid zones with and without migratory divides offers a rare opportunity to directly examine the contribution of divergent migratory behaviour to reproductive barriers. We show that across replicate sampling transects of two pairs of barn swallow (
Hirundo rustica ) subspecies, strong reproductive isolation coincided with a migratory divide spanning 20 degrees of latitude. A third subspecies pair exhibited no evidence for a migratory divide and hybridised extensively. Within migratory divides, overwintering habitats were associated with assortative mating, implicating a central contribution of divergent migratory behaviour to reproductive barriers. The remarkable geographic coincidence between migratory divides and genetic breaks supports a long‐standing hypothesis that the Tibetan Plateau is a substantial barrier contributing to the diversity of Siberian avifauna.