Complex facilitation and competition in a temperate grassland: loss of plant diversity and elevated CO2 have divergent and opposite effects on oak establishment.

Encroachment of woody vegetation into grasslands is a widespread phenomenon that alters plant community composition and ecosystem function. Woody encroachment is often the result of fire suppression, but it may also be related to changes in resource availability associated with global environmental change. We tested the relative strength of three important global change factors (CO(2) enrichment, nitrogen deposition, and loss of herbaceous plant diversity) on the first 3 years of bur oak (Quercus macrocarpa) seedling performance in a field experiment in central Minnesota, USA. We found that loss of plant diversity decreased initial oak survival but increased overall oak growth. Conversely, elevated CO(2) increased initial oak seedling survival and reduced overall growth, especially at low levels of diversity. Nitrogen deposition surprisingly had no net effect on survival or growth. The magnitude of these effects indicates that long-term woody encroachment trends may be most strongly associated with those few individuals that survive, but grow much larger in lower diversity patches. Further, while the CO(2) results and the species richness results appear to describe opposing trends, this is due only to the fact that the natural drivers are moving in opposite directions (decreasing species richness and increasing CO(2)). Interestingly, the mechanisms that underlie both patterns are very similar, increased CO(2) and increased species richness both increase herbaceous biomass which (1) increases belowground competition for resources and (2) increases facilitation of early plant survival under a more diverse plant canopy; in other words, both competition and facilitation help determine community composition in these grasslands.