Hardwood forests in eastern North America are being colonized by multiple nonnative plant and animal species. Colonization rates can be affected by stand structure and distance from edge. We sampled earthworm densities and understory plant species cover in transects located in paired old- and second-growth forests in Indiana. Two 100-m transects were established within each forest stand during late April to early May in each year. One transect was placed parallel to and within 5 m of a south- or west-facing edge. The second transect was placed parallel to the first. but at no less than 100 m from any edge. Nonnative earthworms and plants were found in forest edge and interior regardless of structural stage (second-growth vs. old-growth). The number of native plant species decreased linearly as the densities of adult Lumbricus and Aporrectodea earthworms and the percent cover of multiflora rose (an invasive plant species) increased. Densities of L. terrestris and Aporrectodea earthworms and percent cover of multiflora rose cumulatively explained 39% of the variation in the number of native plant species found in transects across the state. However, multivariate analyses suggested that the species composition of Indiana understory plant communities was affected more by geography than by earthworm densities. Our results suggest that nonnative earthworms and plants are ubiquitous in Indiana hardwood forests and that they may reduce the number of native plant species.

Two factors that can degrade native plant community composition and structure, and hinder restoration efforts, are invasive species and chronic overbrowsing by ungulates such as white-tailed deer. Beginning in 2007, the effectiveness, costs, and impacts of Japanese barberry control treatments and herbivory on nonnative and native plant communities was examined at eight study areas over 4 to 5 yr. Prescribed burning and mechanical mowing by wood shredder or brush saw were utilized as initial treatments to reduce the aboveground portion of established barberry and were equally effective. Without a follow-up treatment, barberry had recovered to 56 to 81% of pretreatment levels 50 to 62 mo after initial treatment. Follow-up treatments in mid-summer to kill new sprouts included directed heating and foliar herbicide applications. Relative to untreated controls, follow-up treatments lowered barberry cover 50 to 62 mo after initial treatment by at least 72%. Although all follow-up treatments were equally effective, the labor cost of directed heating was four times higher than for herbicide applications. Follow-up treatment type (directed heating vs. herbicide) had minimal impact on species other than barberry. White-tailed deer herbivory had a larger impact on other species than did barberry control treatments. Native grass and fern cover was higher outside of exclosures. Areas inside exclosures had higher cover of Oriental bittersweet and multiflora rose, but not Japanese barberry. Thus, recovery of native communities will require more than simply removing the dominant invasive species where deer densities are high. Excellent reduction of Japanese barberry cover can be achieved using either directed heating or herbicides as follow-up treatments in a two-step process, but other invasive plants may become a problem when barberry is removed if deer populations are low.