Resilience of beach grasses along a biogeomorphic successive gradient: resource availability vs. clonal integration

Coastal ecosystems are often formed through two-way interactions between plants and their physical landscape. By expanding clonally, landscape-forming plants can colonize bare unmodified environments and stimulate vegetation-landform feedback interactions. Yet, to what degree these plants rely on clonal integration for overcoming physical stress during biogeomorphological succession remains unknown. Here, we investigated the importance of clonal integration and resource availability on the resilience of two European beach grasses (i.e. Elytrigia juncea and Ammophila arenaria) over a natural biogeomorphic dune gradient from beach (unmodified system) to foredune (biologically modified system). We found plant resilience, as measured by its ability to recover and expand following disturbance (i.e. plant clipping), to be independent on the presence of rhizomal connections between plant parts. Instead, resource availability over the gradient largely determined plant resilience. The pioneer species, Elytrigia, demonstrated a high resilience to physical stress, independent of its position on the biogeomorphic gradient (beach or embryonic dune). In contrast, the later successional species (Ammophila) proved to be highly resilient on the lower end of its distribution (embryonic dune), but it did not fully recover on the foredunes, most likely as a result of nutrient deprivation. We argue that in homogenously resource-poor environments as our beach system, overall resource availability, instead of translocation through a clonal network, determines the resilience of plant species. Hence, the formation of high coastal dunes may increase the resistance of beach grasses to the physical stresses of coastal flooding, but the reduced marine nutrient input may negatively affect the resilience of plants.