Loss of native herbaceous species due to woody plant encroachment facilitates the establishment of an invasive grass

Although negative relationships between diversity (frequently measured as species richness) and invasibility at neighborhood or community scales have often been reported, realistic natural diversity gradients have rarely been studied at this scale. We recreated a naturally occurring gradient in species richness to test the effects of species richness on community invasibility. In central Texas savannas, as the proportion of woody plants increases (a process known as woody plant encroachment), herbaceous habitat is both lost and fragmented, and native herbaceous species richness declines. We examined the effects of these species losses on invasibility in situ by removing species that occur less frequently in herbaceous patches as woody plant encroachment advances. This realistic species removal was accompanied by a parallel and equivalent removal of biomass with no changes in species richness. Over two springs, the nonnative bunchgrass Bothriochloa ischaemum germinated significantly more often in the biomass-removal treatment than in unmanipulated control plots, suggesting an effect of native plant density independent of diversity. Additionally, significantly more germination occurred in the species-removal treatment than in the biomass-removal treatment. Changes in species richness had a stronger effect on B. ischaemum germination than changes in plant density, demonstrating that niche-related processes contributed more to biotic resistance in this system than did species-neutral competitive interactions. Similar treatment effects were found on transplant growth. Thus we show that woody plant encroachment indirectly facilitates the establishment of an invasive grass by reducing native diversity. Although we found a negative relationship between species richness and invasibility at the scale of plots with similar composition and environmental conditions, we found a positive relationship between species richness and invasibility at larger scales. This apparent paradox is consistent with reports from other systems and may be the result of variation in environmental factors at larger scales similarly influencing both invasibility and richness. The habitat loss and fragmentation associated with woody plant encroachment are two of many processes that commonly threaten biodiversity, including climate change. Many of these processes are similarly likely to increase invasibility via their negative effects on native diversity.