Exotic earthworms of great lakes forests: A search for indicator plant species in maple forests

The invasion of exotic earthworms in previously earthworm-free northern deciduous forests has been linked to the disappearance of forest floor litter, declines in plant species richness, and the development of monotypic stands of Carex pensylvanica. However, the impact of exotic earthworms on the regeneration of trees and understory plants is largely unknown. We examined the relationships between earthworm density, plant species richness, leaf litter accumulation, number of tree (Acer) seedlings, and cover of C. pensylvanica at 14 sites in mesic hardwood forests of the Chequamegon-Nicolet National Forest in northern Wisconsin. Earthworm biomass, especially of the genera Lumbricus, Aporrectodea, and Octolasion was negatively associated with leaf litter mass and number of Acer seedlings. Nine plant species, including two spring ephemerals (Dicentra cucullaria and Osmorhiza claytonii), a sedge (Carexdeweyana), and an invasive mint (Galeopsis tetrahit) were negatively correlated with exotic earthworm densities. Dryopteris intermedia and Allium tricoccum, on the other hand, were associated with high earthworm densities. The activities of exotic earthworms appear to have significant impacts on Acer regeneration and the persistence of many herbaceous plants. Our findings suggest that the effects of exotic earthworms on litter mass are strongly related to the observed effects on species composition. Sensitive species can be used as indicators of high and low earthworm densities and might be useful for identifying forests of high conservation value where future invasions of exotic earthworms should be prevented.