Using a nested plot design in oak forests in Minnesota, USA we measured the percent cover of young individuals of an exotic invading shrub, Rhamnus cathartica ( common buckthorn), to assess the relationships at two scales between invasibility, propagule and light availability, and richness and cover of native species. Comparing patches (1 m(2)) within 17 Quercus-dominated stands (each 1 ha, within a 100 km by 150 km area), cover of young R. cathartica was negatively related to both species richness and cover of native species. In 1 m(2) patches, native cover was positively associated with native richness and thus cover-related competition was a likely mechanism by which richness influenced R. cathartica. At the landscape scale (comparing the aggregate stand-scale metrics among the 17 stands), native cover and richness were still positively related, but had opposite relationships with R. cathartica cover. R. cathartica cover was positively related to species richness and negatively related to native species cover. The observed switch at different scales from a positive to a negative relationship between R. cathartica cover and native richness supported the hypothesized scale dependence of these relations. Propagule pressure, which we estimated by measuring the size of nearby mature R. cathartica shrubs, had a large positive effect on R. cathartica seedling cover at the landscape scale. These results suggest that landscape patterns of invasion may be best understood in light of the combination of many factors including native diversity, native cover, and propagule pressure.
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