Effects of tree sapling diversity and nutrient addition on herb-layer invasibility in communities of subtropical species

Exotic species are assumed to alter ecosystem functioning. However, little is known of the relationships within vertically structured plant communities such as forests, where tree saplings interact with herbaceous species, especially in the early phases of succession. This relationship was tested in a common garden experiment which assessed the impacts on tree saplings and herbaceous species following nutrient addition and the introduction of exotic herb species. The experiment was established in South-East China using four broad-leaved tree species (Elaeocarpus decipiens, Schima superba, Castanea henryi and Quercus serrata) to study the relationships between tree sapling diversity, herb-layer productivity and invasibility. Tree saplings were planted in monoculture and in mixtures of two and four species. A full factorial design was applied, within which species composition was crossed with nutrient and exotic seed-addition treatments. The seed-addition treatment included mixtures of seeds from eight exotic herb species, and herb community attributes were assessed after a four month growing season. Results indicate that certain tree species negatively affect native as well as exotic herbs; however, the high productivity of native herbs had a stronger negative impact on exotic species than tree saplings. Nutrient addition increased the productivity of exotic herbs but had no effect on native herbs. Remarkably, exotic species introduction had a negative feedback effect on the growth of tree saplings, which highlights the potential of exotic herbs to diminish tree recruitment. Although tree saplings reduced invasive effects on the herb-layer duriing the earliest phase of forest succession, nutrient addition had a more profound and op- posite effect on these invaders.