Heavy metals in nectar modify behaviors of pollinators and nectar robbers: Consequences for plant fitness.

Plants growing in heavy-metal-rich soils can accumulate metals into their nectar. Nectar chemical composition can alter foraging behavior of floral visitors (including pollinators and floral antagonists) and further affect plant reproductive fitness. The role of nectar heavy metals in deterring pollinators (e.g., shortening foraging time) has been recently studied, but their effects on plant reproduction via changes in behaviors of both pollinators and floral antagonists (e.g., nectar robbers) are less understood. We experimentally manipulated four nectar heavy metals (Zn, Cu, Ni, and Pb) in a native ornamental plant, Hosta ensata F. Maekawa, to investigate the effect of nectar metals on plant reproductive success. We also recorded nectar robbing as well as foraging time and visitation rate of pollinators to assess whether nectar metals could alter the behavior of antagonists and mutualists. Although metals in nectar had no significant direct effects on plant reproduction via hand-pollination, we detected their positive indirect effects on components of female fitness mediated by pollinators and nectar robbers. Matching effects on female plant fitness, nectar robbers responded negatively to the presence of metals in nectar, robbing metal-treated flowers less often. Pollinators spent less time foraging on metal-treated flowers, but their visitation rate to metal-treated flowers was significantly higher than to control flowers. Moreover, pollinators removed less nectar from flowers treated with metals. Our results provide the first direct evidence to date that heavy metals in nectar are capable of deterring nectar robbers and modifying pollinator foraging behavior to enhance plant reproductive fitness.