1. The abiotic environment drives species abundances and distributions both directly and indirectly through effects on multi-trophic species interactions. However, few studies have documented the individual and combined consequences of these direct and indirect effects. 2. We studied an ant-tended a

1. Plants interact simultaneously with both mutualists and antagonists. While webs of plant-animal interactions in natural systems can be highly complex, most interactions can be simplified into those that are either direct (mediated through pairwise interactions) or indirect (mediated through third