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Virtually all plants employ direct and indirect defenses against herbivores. While it is known that plant defenses can be affected by belowground symbiotic microbes under controlled conditions, studies showing these multitrophic interactions in nature are surprisingly scarce. Here we tested for effects of rhizobia on insect attraction and direct defense (cyanogenesis) in wild lima bean (Phaseolus lunatus) plants in Costa Rica. We performed bioassays with rhizobia-inoculated (R+) and rhizobiafree (R-) potted plants distributed among native lima bean communities at two spatially separated field sites (450 km apart) and in two field seasons. Without affecting overall plant size, rhizobia altered leaf chemistry (cyanogenesis and soluble leaf nitrogen) and ultimately insect communities visiting the plants. Natural herbivorous chrysomelid beetles were strongly attracted to R+ plants, while natural enemies, ants and parasitoid wasps, preferred R- plants resulting in a particularly high herbivore:carnivore ratio on R + plants. This suggests that symbiotic microbes mediate trophic interactions by influencing both direct and indirect plant defenses against herbivores. Our results show that rhizobia affect the plant defensive phenotype and have cascading effects on plant-insect interactions in nature. 

Abstract The diversity of specialized molecules produced by plants radiating along ecological gradients is thought to arise from plants' adaptations to local conditions. Therefore, closely related species growing in similar habitats should phylogenetically converge, or diverge, in response to similar climates, or similar interacting animal communities. We here asked whether closely related species in the genusHaplopappus(Asteraceae) growing within the same elevation bands in the Andes, converged to produce similar floral odors. To do so, we combine untargeted analysis of floral volatile organic compounds with insect olfactory bioassay in congenericHaplopappus(Asteraceae) species growing within the same elevation bands along the Andean elevational gradient. We then asked whether the outcome of biotic interactions (i.e., pollination vs. seed predation) would also converge across species within the same elevation. We found that flower odors grouped according to their elevational band and that the main floral visitor preferred floral heads from low‐elevation band species. Furthermore, the cost–benefit ratio of predated versus fertilized seeds was consistent within elevation bands, but increased with elevation, from 6:1 at low to 8:1 at high elevations. In the light of our findings, we propose that climate and insect community changes along elevation molded a common floral odor blend, best adapted for the local conditions. Moreover, we suggest that at low elevation where floral resources are abundant, the per capita cost of attracting seed predators is diluted, while at high elevation, sparse plants incur a higher herbivory cost per capita. Together, our results suggest that phytochemical convergence may be an important factor driving plant–insect interactions and their ecological outcomes along ecological gradients.