Direct and indirect effects of allelopathy in the soil legacy of an exotic plant invasion

Invasive species may leave behind legacies that persist even after removal, inhibiting subsequent restoration efforts. We examined the soil legacy of Cytisus scoparius, a nitrogen-fixing, putatively allelopathic shrub invading the western US. We tested the hypothesis that allelopathy plays a critical role in the depressive effect of Cytisus on the key native Douglas-fir, both directly on tree growth and indirectly via effects on its ectomycorrhizal fungi (EMF). In a greenhouse factorial experiment, we used activated carbon to inhibit Cytisus-produced allelochemicals and sucrose to reduce elevated nitrogen (N). We found that: (1) Cytisus-invaded soils depressed Douglas-fir growth compared to uninvaded forest soils. The effect of adding Cytisus litter was positive (possibly reflecting an N fertilization effect) only in the presence of activated carbon, providing evidence for a role of allelopathic compounds. Activated carbon did not increase growth in the absence of Cytisus litter. Finally, sucrose addition provided weak support for a nitrogen effect of Cytisus litter. (2) Seedlings grown in Cytisus soils had lower EMF abundance compared to those in uninvaded forest soils. In forest soil from one site, adding Cytisus litter also decreased EMF abundance. Douglas-fir growth increased significantly with EMF across sites and soils suggesting that changes in EMF were linked to tree growth. The fungal taxon Cenococcum geophilum was significantly depressed in Cytisus soils compared to forest soils, while Rhizopogon rogersii abundance was similar across soil types. These results together suggest an overall negative effect of Cytisus on the growth of a dominant native tree and its fungal symbionts. Our study suggests how the role of allelopathy in ecological restoration may play out on two time scales: through immediate, direct impacts on native plants as well as through long-term, persistent impacts mediated by the collapse or transformation of microbial communities.