Abstract
Pollination syndromes reflect the convergence of floral traits among plants sharing the same pollinator guild. However, bee-pollinated orchids exhibit striking variation in color and size. This diversity reflects the multiple reward strategies that evolved within the family, each interacting differently with bee sensory biases. Here, we tested whether the complex floral visual displays of orchids differ in signal identity and intensity among reward systems. We also considered intrafloral modularity, measured as the color differentiation among flower parts, and color-size integration. For this, we measured and modeled floral morphometric and reflectance data from sepals, petals, lip tips, and lip bases under bee vision from 95 tropical Epidendroid species to compare chromatic and achromatic contrasts, spectral purity, and mean reflectance across wavebands, plus flower and display size, among reward systems. Reward types included 19 food-deceptive, 8 nectar-offering, 10 oil-offering, 11 fragrance-offering, and 47 orchid species of unknown reward strategy. Principal component analyses on 34 color and 9 size variables summarized major gradients of visual trait variation: first component (19.1%) represented overall green-red reflectance and achromatic contrasts, whereas the second (16.5%) captured chromatic contrast-size covariation. Reward systems differed mostly in signal identity rather than signal intensity. Flower chromatic contrasts presented strong integration with flower size, while achromatic contrasts were negatively associated with display size. While deceptive and nectar-offering orchids tend toward larger solitary flowers with bluer and spectrally purer displays, oil- and fragrance-offering orchids tend toward smaller, brownish, or yellow to green flowers, with larger inflorescences. Rewardless orchids presented more achromatically conspicuous signals than rewarding orchids, but smaller displays. Orchid species clustered by reward both in PCA spaces and in bee hexagon color space. Deceptive orchids were typically associated with UV + White colors, oil orchids with UV + Yellow lip tips, and fragrance orchids with UV-Black lip bases and UV-Green lip tips. Together, these results indicate that orchid reward systems promote qualitative rather than quantitative differentiation in visual signals, integrating display color and size. These long-evolved distinct signals potentially enable foraging bees to discriminate among resource types within the community floral market. Our results demonstrate that color and flower display size are important predictors of reward strategy, likely used by foraging bees for phenotype-reward associations, thus mediating the evolution of floral signals.