“…In our study, upslope locations were consistently closer to potential sources of invasion, such as trails, roads, and homes, corroborating previous observations that earthworm invasions are frequently facilitated by human activity and development (Suárez, Tierney, et al ., 2006; Paudel et al ., 2016; Bellitürk et al ., 2015). Asplenium scolopendrium is also associated with rocky and steep terrain, and thus might be less vulnerable to earthworm invasions and subsequent impacts, as earthworms are less abundant and more likely to be absent from areas with shallower soil and steeper slopes (Fleri and Arcese, 2021; Suárez, Tierney, et al ., 2006). However, earthworms can be carried downslope to A. scolopendrium populations and ravine bottoms by water, especially during storm events, suggesting ferns might experience episodic earthworm dispersal events.…”