Soil heterogeneity tends to promote the growth of naturalized aliens when competing with native plant communities

1. Elton's diversity-invasibility hypothesis predicts that diverse communities are more resistant against alien invaders. However, observational studies frequently find positive relationships between the numbers of alien and native species. It has been suggested, but rarely tested, that environmental heterogeneity may cause such positive relationships.2. Here, we experimentally tested the effects of soil heterogeneity and diversity (species richness) on the invasibility of native communities. We first filled mesocosm pots with either a heterogeneous soil, consisting of one patch of sand, one patch of peat and one patch of an equal mixture of both substrates, or a homogeneous soil, consisting of the mixture only. Then, we planted those pots with 29 native communities consisting of one, four or eight species, and invaded them with populations of four individuals of one of five alien species.3. In the heterogeneous soils, individual alien plants benefited more strongly from the resource-rich peat soil than the native communities did. Moreover, in the mixture soil of the heterogeneous treatment, individual alien plant over-proportionally produced more biomass than in the mixture soil of the homogeneous treatment. Consequently, the populations of naturalized alien plants in each pot benefited from heterogeneous soil conditions, and this tended to be particularly the case when a native community was present. The native communities did not respond to soil heterogeneity, but they had a negative effect on the naturalized plants, irrespective of their diversity.4. Synthesis. Our results indicate that soil heterogeneity might alleviate the competitive effects of native communities on the alien invaders, as the latter took more advantage of the high resource patches than the natives did. The beneficial effect of heterogeneity on invasion success could thus explain why observational studies usually find positive relationships between the numbers of alien and native species.publishe