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A major challenge for studies assessing drivers of phenotypic divergence is the statistical comparison of taxa with unique, often unknown, evolutionary histories, and for which there are no clear expected trait values. Because many traits are fundamentally constrained by energy availability, we suggest that trait values predicted by scaling theories such as the metabolic theory of ecology (MTE) can provide baseline expectations. Here, we introduce a metabolic scaling‐based approach to test theory involving the direction and magnitude of ecological and sexual selection, using vocal frequency as an example target of selection. First, we demonstrate that MTE predicts the relationship between the natural log of body size and natural log of vocal frequency across 795 bird species, controlling for phylogeny. Family‐wide deviations in slope and intercepts from MTE estimates reveal taxa with potentially important differences in physiology or natural history. Further, species‐level frequency deviations from MTE expectations are predicted by factors related to ecological and sexual selection and, in some cases, provide evidence that differs from current understanding of the direction of selection and identity of ecological selective agents. For example, our approach lends additional support to the findings from many cross‐habitat studies that suggest that dense vegetation selects for lower frequency signals. However, our analysis also suggests that birds in non‐forested environments vocalize at frequencies higher than expected based on MTE, prompting intriguing questions about the selective forces in non‐forest environments that may act on vocal frequency. Additionally, vocal frequency deviates more strongly from MTE expectations among species with smaller repertoires and those with low levels of sexual dichromatism, complicating the use of these common sexual selection surrogates. Broad application of our metabolic scaling approach might provide an important complementary approach to understanding how selection shapes phenotypic evolution by offering a common baseline across studies and taxa and providing the basis to explore evolutionary trade‐offs within and among multicomponent and multimodal traits.

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 >23,000SNPs 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.

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.