Effects of earthworm invasion on plant species richness in northern hardwood forests

The invasion of non-native earthworms (Lumbricus spp.) into a small number of intensively studied stands of northern hardwood forest has been linked to declines in plant diversity and the local extirpation of one threatened species. It is unknown, however whether these changes have occurred across larger regions of hardwood forests, which plant species are most vulnerable, or with which earthworm species such changes are associated most closely. To address these issues we conducted a regional survey in the Chippewa and Chequamegon national forests in Minnesota and Wisconsin (U.S.A.), respectively. We sampled earthworms, soils, and vegetation, examined deer browse in 20 mature, sugar-maple-dominated forest stands in each national forest, and analyzed the relationship between invasive earthworms and vascular plant species richness and composition. Invasion by Lumbricus was a strong indicator of reduced plant richness in both national forests. The mass of Lumbricus juveniles was significantly and negatively related to plant-species richness in both forests. In addition, Lumbricus was a significant factor affecting plant richness in a full model that included multiple variables. In the Chequamegon National Forest earthworm mass was associated with higher sedge cover and lower cover of sugar maple seedlings and several forb species. The trends were similar but not as pronounced in Chippewa, perhaps due to lower deer densities and different earthworm species composition. Our results provide regional evidence that invasion by Lumbricus species may be an important mechanism in reduced plant-species richness and changes in plant communities in mature forests dominated by sugar maples.