Do the impacts of an ecosystem engineer vary along an environmental gradient? Tests with the leaf-cutter ant Atta laevigata in a Neotropical savanna
Background. Species that transform habitats in ways that influence other species are known as Ecosystem Engineers. While the impacts of many engineers have been well described, our understanding of how these impacts vary along or alter environmental gradients remains limited. While disentangling the effects of gradients and engineers on biodiversity is complicated – the gradients themselves can be altered by engineers – doing so is necessary to advance models of ecosystem engineering. We used leaf-cutter ants (Atta spp.), a major engineer in the Neotropics, as a model system with which to investigate the relative influence of gradients and engineers on the abundance and diversity of seedlings. Methods. We conducted our research South Americas Cerrado (savanna woodlands). We used a survey of seedlings along a canopy cover gradient and data on environmental conditions previously shown to influence seedling establishment that were collected at different distances from ant colonies to fit statistical models addressing the following questions: Do the environmental conditions previously shown to influence seedling establishment vary with canopy cover? Does A. laevigata alter canopy cover and related environmental conditions influencing seedling establishment? What is the spatial extent of Atta impact on the gradient and environmental conditions, and how does this vary along the canopy cover gradient? Do A. laevigata and canopy cover act independently or in synergy to influence seedling abundance and species richness? Results. We found that the environmental conditions varied in concert with canopy cover, but that ants are not modifying canopy cover. However, ants are modifying environmental conditions, with the magnitude and spatial extent of changes consistent across the gradient. In contrast to prior studies, we found that seedling abundance and diversity were independent of canopy-related environmental conditions. Proximity to leaf-cutter nests had the strongest effect on diversity and richness. Discussion. Atta laevigata engineers environmental conditions in the Cerrado, with strong but spatially restricted consequences for seedling communities. We hypothesize that ants indirectly reduce seedling establishment by increasing rates of seed dessication and reduce seedling growth and survivorship. As such the indirect reduction of seedling abundance due to engineering could exacerbate the direct reduction due to herbivory. The effects of Atta appear restricted to the nest mound, but they could be long-lasting because mounds persist long after a colony has died or migrated. These results underscore the hypothesis that Atta plays a dominant role in Cerrado plant demography, and that its footprint may increase dramatically in coming decades due to ongoing transformations of the Cerrado.