High intermediary mutualist density provides consistent biological control in a tripartite mutualism

Understanding the ecology of mutualisms becomes a particularly important task when considering agroecosystems, as many ecosystem services are associated with mutualistic interactions. Here we report on experiments associated with an indirect pest control mutualism between the arboreal nesting ant Azteca sericeasur and coffee. This system is particularly interesting because the indirect Azteca-Coffea mutualism emerges from an Azteca-scale insect mutualism that takes place on the coffee plant. We describe this interaction structure as a mutualism-dependent mutualism and ask whether the density of intermediary mutualist (scale insects, Coccus) that benefits Azteca also influences the benefits provided to coffee plants. We found that indeed Azteca's benefit to Coffea is consistent when Coccus density is high. Furthermore, we also found that at low Coccus density Azteca only benefits Coffea in the beginning of the rainy season, and this effect is likely due to the fact that Coccus produces less sugars with higher precipitation. We suggest a framework for thinking about context-dependency in agroecosystem mutualisms that may provide a more mechanistic way to tease apart the prevalent context-dependent results in ecological literature. Finally, we address some past recommendations as it pertains to the management of the Azteca-Coffea-Coccus complex in coffee agroecosystems.