Regional extent of an ecosystem engineer: Earthworm invasion in northern hardwood forests

The invasion of exotic earthworms into northern temperate and boreal forests previously devoid of earthworms is an important driver of ecosystem change. Earthworm invasion can cause significant changes in soil structure and communities, nutrient cycles, and the diversity and abundance of herbaceous plants. However, the regional extent and patterns of this invasion are poorly known. We conducted a regional survey in the Chippewa and Chequamegon National Forests, in Minnesota and Wisconsin, USA, respectively, to measure the extent and patterns of earthworm invasion and their relationship to potential earthworm introduction sites. We sampled earthworms, soils, and vegetation in 20 mature, sugar maple-dominated forest stands in each national forest and analyzed the relationship between the presence of five earthworm taxonomic groups, habitat variables, and distance to the nearest potential introduction site. Earthworm invasion was extensive but incomplete in the two national forests. Four of the six earthworm taxonomic groups occurred in 55-95% of transects; however 20% of all transects were invaded by only one taxonomic group that has relatively minor ecological effects. Earthworm taxonomic groups exhibited a similar sequence of invasion found in other studies: Dendrobaena > Aporrectodea = Lumbricus juveniles > L. rubellus > L. terrestris. Distance to the nearest road was the best predictor of earthworm invasion in Wisconsin while distance to the nearest cabin was the best predictor in Minnesota. These data allow us to make preliminary assessments of landscape patterns of earthworm invasion. As an example, we estimate that 82% of upland mesic hardwood stands in the Wisconsin region are likely invaded by most taxonomic groups while only 3% are unlikely to be invaded at present. Distance to roads and cabins provides a coarse-scale predictor of earthworm invasion to focus stand-level assessments that will help forest managers better understand current and potential forest conditions and identify uninvaded areas that could serve as important refugia for plant species threatened by earthworm invasion.