Synergy between pathogen release and resource availability in plant invasion

Abstract
Why do some exotic plant species become invasive? Two common hypotheses, increased resource availability and enemy release, may more effectively explain invasion if they favor the same species, and therefore act in concert. This would be expected if plant species adapted to high levels of available resources in their native range are particularly susceptible to enemies, and therefore benefit most from a paucity of enemies in their new range. We tested this possibility by examining how resource adaptations influence pathogen richness and release among 243 European plant species naturalized in the United States. Plant species adapted to higher resource availability hosted more pathogen species in their native range. Plants from mesic environments hosted more fungi than plants from xeric environments, and plants from nitrogen-rich environments hosted more viruses than plants from nitrogen-poor environments. Furthermore, plants classified as competitors hosted more than 4 times as many fungi and viruses as did stress tolerators. Patterns of enemy release mirrored those of pathogen richness: competitors and species from mesic and nitrogen-rich environments were released from many pathogen species, while stress tolerators and species from xeric and nitrogen-poor environments were released from relatively few pathogen species. These results suggest that enemy release contributes most to invasion by fast-growing species adapted to resource-rich environments. Consequently, enemy release and increases in resource availability may act synergistically to favor exotic over native species.