Changes in hardwood forest understory plant communities in response to European earthworm invasions.

European earthworms are colonizing earthworm-free northern hardwood forests across North America. Leading edges of earthworm invasion provide an opportunity to investigate the response of understory plant communities to earthworm invasion and whether the species composition of the earthworm community influences that response. Four sugar maple-dominated forest sites with active earthworm invasions were identified in the Chippewa National Forest in north central Minnesota, USA. In each site, we established a 30 x 150 m sample grid that spanned a visible leading edge of earthworm invasion and sampled earthworm populations and understory vegetation over four years. Across leading edges of earthworm invasion, increasing total earthworm biomass was associated with decreasing diversity and abundance of herbaceous plants in two of four study sites, and the abundance and density of tree seedlings decreased in three of four study sites. Sample points with the most diverse earthworm species assemblage, independent of biomass, had the lowest plant diversity. Changes in understory plant community composition were most affected by increasing biomass of the earthworm species Lumbricus rubellus. Where L. rubellus was absent there was a diverse community of native herbaceous plants, but where L. rubellus biomass reached its maximum, the herbaceous-plant community was dominated by Carex pensylvanica and Arisaema triphyllum and, in some cases, was completely absent. Evidence from these forest sites suggests that earthworm invasion can lead to dramatic changes in the understory community and that the nature of these changes is influenced by the species composition of the invading earthworm community.