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Abstract PremiseEvolutionary theory predicts polymorphism should be rare; however, intraspecific variation in floral color is common and can be attributed to genetic drift, plasticity, or variable selection. Examining floral color polymorphism both within contact zones and across a species' range can reveal the mechanisms maintaining this variation. Here, we used a multistep approach to investigate spatially heterogeneous variation in floral bract color inCastilleja coccinea. MethodsWe compared frequencies of color morphs, floral morphology, fitness, and genetic structure in regional populations and in a common garden. Next, we examined habitat differences, including edaphic factors, as potential drivers of variation. Lastly, we leveraged herbarium and iNaturalist occurrence data to investigate whether patterns were consistent at the landscape scale. ResultsBract color inC. coccineais genetically heritable, with yellow dominant over red, and is under selection. Populations are predominantly monomorphic, with color distance showing no correlation to genetic or geographic distance, despite significant genetic isolation by distance. Yellow morphs were associated with open wetlands, while red morphs occurred at drier sites with nearby tree cover. Red morphs demonstrated lower fitness in a common garden, suggesting trade‐offs associated with pleiotropic effects of floral color. ConclusionsDifferences in floral color between morphs are consistent with diversification associated with a shift in ecological niche. We identified variation in edaphic and habitat conditions as probable drivers of divergence in floral color. Additionally, variation in other floral traits suggests a combined role of pollinators and habitat differences acting in concert to maintain distinct floral color morphs. 

Abstract Oenothera sect. Calylophus is a North American group of 13 recognized taxa in the evening primrose family (Onagraceae) with an evolutionary history that may include independent origins of bee pollination, edaphic endemism, and permanent translocation heterozygosity. Like other groups that radiated relatively recently and rapidly, taxon boundaries within Oenothera sect. Calylophus have remained challenging to circumscribe. In this study, we used target enrichment, flanking noncoding regions, gene tree/species tree methods, tests for gene flow modified for target-enrichment data, and morphometric analysis to reconstruct phylogenetic hypotheses, evaluate current taxon circumscriptions, and examine character evolution in Oenothera sect. Calylophus. Because sect. Calylophus comprises a clade with a relatively restricted geographic range, we were able to extensively sample across the range of geographic, edaphic, and morphological diversity in the group. We found that the combination of exons and flanking noncoding regions led to improved support for species relationships. We reconstructed potential hybrid origins of some accessions and note that if processes such as hybridization are not taken into account, the number of inferred evolutionary transitions may be artificially inflated. We recovered strong evidence for multiple evolutionary origins of bee pollination from ancestral hawkmoth pollination, edaphic specialization on gypsum, and permanent translocation heterozygosity. This study applies newly emerging techniques alongside dense infraspecific sampling and morphological analyses to effectively reconstruct the recalcitrant history of a rapid radiation. [Gypsum endemism; Oenothera sect. Calylophus; Onagraceae; phylogenomics; pollinator shift; recent radiation; target enrichment.]