Disturbance-mediated competition and the spread of Phragmites australis in a coastal marsh

In recent decades the grass Phragmites australis has been aggressively invading coastal, tidal marshes of North America, and in many areas it is now considered a nuisance species. While P. australis has historically been restricted to the relatively benign upper border of brackish and salt marshes, it has been expanding seaward into more physiologically stressful regions. Here we test a leading hypothesis that the spread of P. australis is due to anthropogenic modification of coastal marshes. We did a field experiment along natural borders between stands of P. australis and the other dominant grasses and rushes (i.e., matrix vegetation) in a brackish marsh in Rhode Island, USA. We applied a pulse disturbance in one year by removing or not removing neighboring matrix vegetation and adding three levels of nutrients (specifically nitrogen) in a factorial design, and then we monitored the aboveground performance of P. australis and the matrix vegetation. Both disturbances increased the density, height, and biomass of shoots of P. australis, and the effects of fertilization were more pronounced where matrix vegetation was removed. Clearing competing matrix vegetation also increased the distance that shoots expanded and their reproductive output, both indicators of the potential for P. australis to spread within and among local marshes. In contrast, the biomass of the. matrix vegetation decreased with increasing severity of disturbance. Disturbance increased the total aboveground production of plants in the marsh as matrix vegetation was displaced by P. australis. A greenhouse experiment showed that, with increasing nutrient levels, P. australis allocates proportionally more of its biomass to aboveground structures used for spread than to belowground structures used for nutrient acquisition. Therefore, disturbances that enrich nutrients or remove competitors promote the spread of P. australis by reducing belowground competition for nutrients between P. australis and the matrix vegetation, thus allowing P. australis, the largest plant in the marsh, to expand and displace the matrix vegetation. Reducing nutrient load and maintaining buffers of matrix vegetation along the terrestrial-marsh ecotone will, therefore, be important methods of control for this nuisance species.