RESUMEN

Nonnative European earthworms are invading hardwood forests of the Chippewa National Forest, MN. While effects on plant communities at the leading edge of invasion have been studied, little is known about longer-term effects of invasive earthworms. We applied a model using historic O-horizon soil thickness and a chronosequence approach to classify 41 hardwood sites in the Chippewa National Forest as "long-term wormed" (wormed >2 decades), "short-term wormed" or "unwormed/lightly wormed." Graminoids, especially Carex pensylvanica, had the greatest mean percent cover in sites that had been wormed for over two decades. The families with the greatest negative change in mean percent cover after over two decades of earthworm invasion were Asteraceae, Violaceae, and Sapindaceae (specifically Acer species). Across all diversity metrics measured, long-term wormed sites had the lowest understory plant species diversity, short-term wormed sites had intermediate diversity, and unwormed/lightly wormed sites exhibited the highest diversity. Long-term wormed sites had the lowest mean species richness across all sample scales (1-1024 m2). The greatest within-group compositional dissimilarity occurred at sites that had been wormed for over two decades, suggesting that sites that had been wormed for over two decades have not reached a compositionally similar end-state "wormed" community type. Our study suggests that understory diversity will decrease as hardwood forest stands become wormed over time. While our results support other findings that exotic earthworm invasion is associated with lower understory plant diversity in hardwood forests, our study was the first to use space-for-time substitution to document the effects after multiple decades of earthworm invasion.