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The flower is a key evolutionary innovation, one that has led directly to the numerical dominance of angiosperm species among land plants (Regal 1977, Specht & Bartlett 2009). The staggering floral diversity exhibited by angiosperms has long been of interest to botanists and evolutionary biologists, and long attributed to selection by and coevolution with animal pollinators (Sprengel 1793, Darwin 1862, Knuth & Müller 1908, Stebbins 1970). Despite its central role in angiosperm reproduction, pollination is not the only process that shapes floral trait evolution. Many other aspects of the biotic and abiotic environment have been demonstrated to drive variation in flower size and shape, as have whole-plant life history and phenology (Bazzaz et al. 1987, Galen 1999, Teixido et al. 2016). In many cases such forces impose contrasting selective pressures to the typical selection for larger, showier flowers that often arises through competition for pollinator attraction (Bell 1985, Anton et al. 2013). The production of flowers incurs direct energetic costs in the form of carbon, water, and nutrient resources used for construction, as well as ongoing costs for maintenance of what are typically nonphotosynthetic structures. Accordingly, the ‘resource-cost hypothesis’ suggests that smaller, less conspicuous flowers should be favoured in low-resource habitats by virtue of reducing physiological stress from water and nutrient limitation (Galen 1999, Teixido et al. 2016). Indeed, in many systems smaller-flowered species and populations occur in hotter, drier, less fertile environments than those with larger flowers (e.g. Galen 2000, Herrera 2005, Teixido & Valladares 2013). Along with selection imposed by the abiotic environment, interactions with natural enemies such as flo-

Beyond pollinators: evolution of floral architecture with environment across the wild sunflowers (Helianthus, Asteraceae)