Spatially variable propagule pressure and herbivory influence invasion of chaparral shrubland by an exotic grass.

Although numerous studies have identified mechanisms that either resist or facilitate biological invasions, few studies have explicitly tested how resisting and facilitating mechanisms interact to drive invasion success. In California, USA, undisturbed Mediterranean-type shrublands have resisted invasion by the perennial tussock grass Cortaderia jubata. In some cases, however, this resistance has been spectacularly breached even in the absence of large-scale disturbance. I tested the hypothesis that these invasions are facilitated by local reductions in the strength of biotic resistance. I evaluated invasive success using C. jubata seed and seedling additions at different microhabitats: the edge of a chaparral stand, under shrub canopy at different distances from the stand edge, and in canopy gaps within the stand. When left exposed to mammalian herbivores, seedling survivorship decreased sharply from nearly 40% on the stand edge to zero just 10 m into the stand. When transplants were protected from herbivory, however, distance from the edge had no significant influence on transplant survivorship. Seedling emergence was also greater on the edge and in canopy gaps than under the canopy, but these differences were not caused by differences in herbivory. The flux of invasive propagules reaching the soil surface was immense and greater along the edge and within gaps than under the stand canopy. Mirroring these patterns, naturally occurring seedling abundance declined dramatically with distance from the stand edge, and seedlings were far more common within stand gaps than would be expected given gap frequency within the stand. Despite strong biotic resistance to invasion within the stand, the cover of C. jubata has increased 20% over the last 9 years. These results suggest that the relative amount of susceptible edge habitat and the supply of invasive propagules can facilitate invasion even in the face of strong local biotic resistance.