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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 Understanding how evolution proceeds from molecules to organisms to interactions requires integrative studies spanning biological levels. Linking phenotypes with associated genes and fitness illuminates how adaptive walks move organisms between fitness peaks. Floral evolution can confer rapid reproductive isolation, often converging in association with pollinator guilds. Within the monkeyflowers (Mimulussect.Erythranthe), yellow flowers within red hummingbird-pollinated species have arisen at least twice, suggesting possible pollinator shifts. We compare two yellow-flowered forms ofM. cardinalisandM. verbenaceusto their red counterparts in floral phenotypes, biochemistry, transcriptomic and genomic variation, and pollinator interactions. We find convergence in ongoing adaptive walks of both yellow morphs, with consistent changes in traits of large effect (floral pigments, associated gene expression), resulting in strong preference for yellow flowers by bumblebees. Shifts in scent emission and floral opening size also favor bee adaptation, suggesting smaller-effect steps from hummingbird to bee pollination. By examining intraspecific, incipient pollinator shifts in two related species, we elucidate adaptive walks at early stages, revealing how convergent large effect mutations (floral color) may drive pollinator attraction, followed by smaller effect changes for mechanical fit and reward access. Thus, ongoing adaptive walks may impact reproductive isolation and incipient speciation via convergent evolution.