Comparison of direct, indirect, and ecosystem engineering effects of an earthworm on the red-backed salamander.

In addition to creating or modifying habitat, ecosystem engineers interact with other species as predators, prey, or competitors. The earthworm, Lumbricus terrestris, interacts with the common woodland salamander, Plethodon cinereus, via: (1) ecosystem engineering, by providing burrows that are used as a refuge, (2) direct effects as a prey item, and (3) indirectly, by competing with microinvertebrates, another prey item for P. cinereus. Using enclosures in the forest, I examined the relative strengths of these component pathways between seasons and salamander age classes. I found that the relative strength (partial eta2) of the positive direct (trophic) effect of L. terrestris on the change in mass of P. cineresus was greater than that of the negative indirect effect, but only in summer. Positive effects of ecosystem engineering were only evident over the winter as increased adult survival. This research has implications for how habitat provisioning complements more well-studied species interactions, such as competition and predation, within communities.