Differential invasion of a wetland grass explained by tests of nutrients and light availability on establishment and clonal growth.

Phalaris arundinacea (Poaceae) is aggressively invading wetlands across North America. We tested the hypotheses that open canopies and increased nutrients facilitate vegetative establishment in the field, using a phytometer (6 rhizome fragments/plot, 24 plots/wetland). In each of three wetlands, phytometers received three levels of an NPK fertilizer or served as controls. Emergence and survival differed among sites (P=0.0005), but not due to NPK addition. P. arundinacea survival was highest in a wet prairie with a late-developing canopy, but limited by prolonged flooding in one sedge meadow and by an early-growing, dense plant canopy in a second. These patterns were explained in greenhouse experiments, where both flooding (P<0.0001) and heavy shade (P=0.0002) decreased P. arundinacea aboveground biomass by up to 73% and 97%, respectively. Rhizome fragment survival was reduced by 30% under flooded conditions and 25% under heavy shade. We then tested the hypothesis that a clonal subsidy facilitates vegetative expansion into heavy shade. Established clones were allowed access to bare soil under four levels of shade and two levels of NPK fertilizer in a two-factor greenhouse experiment. Young ramets attached to parent clones readily grew into heavy shade, and the high nutrient treatment increased aboveground growth (P<0.0001) and distance of ramet spread (P=0.0051) by nearly 50%. Under low nutrient conditions, root biomass increased by 30% (P<0.0001). P. arundinacea's rapid expansion into a variety of wetland types is likely a function of clonal subsidy, morphological plasticity, and nutrient availability: young ramets that emerge under shaded conditions are supported by parental subsidies; where nutrients are plentiful, P. arundinacea can maximize aboveground growth to capture more light; and where nutrients are scarce, it can increase belowground foraging.