Dear neighbor: Trees with extrafloral nectaries facilitate defense and growth of adjacent undefended trees

Plant diversity can increase productivity. One mechanism behind this biodiversity effect is facilitation, which is when one species increases the performance of another species. Plants with extrafloral nectaries (EFNs) establish defense mutualisms with ants. However, whether EFN plants facilitate defense of neighboring non-EFN plants is unknown. Synthesizing data on ants, herbivores, leaf damage, and defense traits from a forest biodiversity experiment, we show that trees growing adjacent to EFN trees had higher ant biomass and species richness and lower caterpillar biomass than conspecific controls without EFN-bearing neighbors. Concurrently, the composition of defense traits in non-EFN trees changed. Thus, when non-EFN trees benefit from lower herbivore loads as a result of ants spilling over from EFN tree neighbors, this may allow relatively reduced resource allocation to defense in the former, potentially explaining the higher growth of those trees. Via this mutualist-mediated facilitation, promoting EFN trees in tropical reforestation could foster carbon capture and multiple other ecosystem functions.