Ecosystem engineers on plants: indirect facilitation of arthropod communities by leaf-rollers at different scales.

Ecosystem engineering is a process by which organisms change the distribution of resources and create new habitats for other species via non-trophic interactions. Leaf-rolling caterpillars can act as ecosystem engineers because they provide shelter to secondary users. In this study, we report the influence of leaf-rolling caterpillars on speciose tropical arthropod communities along both spatial scales (leaf-level and plant-level effects) and temporal scales (dry and rainy seasons). We predict that rolled leaves can amplify arthropod diversity at both the leaf and plant levels and that this effect is stronger in dry seasons, when arthropods are prone to desiccation. Our results show that the abundance, richness, and biomass of arthropods within several guilds increased up to 22-fold in naturally and artificially created leaf shelters relative to unaltered leaves. These effects were observed at similar magnitudes at both the leaf and plant scales. Variation in the shelter architecture (funnel, cylinders) did not influence arthropod parameters, as diversity, abundance, orbiomass, but rolled leaves had distinct species composition if compared with unaltered leaves. As expected, these arthropod parameters on the plants with rolled leaves were on average approximately twofold higher in the dry season. Empty leaf rolls and whole plants were rapidly recolonized by arthropods over time, implying a fast replacement of individuals; within 15-day intervals the rolls and plants reached a species saturation. This study is the first to examine the extended effects of engineering caterpillars as diversity amplifiers at different temporal and spatial scales. Because shelter-building caterpillars are ubiquitous organisms in tropical and temperate forests, they can be considered key structuring elements for arthropod communities on plants.