Scale-dependent interactions in coastal biogeomorphic landscapes: pioneer both inhibits and facilitates primary foredune builder across spatial scales Version 8

Ecosystem engineers often drive landscape formation and vegetation succession by modifying environmental conditions. Along the Northwestern European coast, dune formation is classically believed to be pioneered by sand couch (Elytrigia juncea), followed by the primary foredune builder marram grass (Calamagrostis arenaria, formerly Ammophila arenaria) once sufficiently large dunes have formed. However, these ideas lack experimental validation, leaving the specific relationship between sand couch, the landscape they form, and marram grass establishment unknown. Here, we experimentally investigated this relationship by planting 975 young marram grass plants in two sand couch dominated, embryonic dune systems. Using structural equation models, we examined the effect of sand couch and the landscape attributes (i.e., elevation, distance to sea and elevation change) on marram grass establishment. Results reveal indirect local inhibition by sand couch via landscape modifications on survival of establishing marram grass. Specifically, sand couch elevated the environment, and elevated areas in turn eroded more in winter, which was the key factor reducing marram grass survival. At the landscape scale, however, we observed the highest survival in relatively stable and sheltered microsites formed because of sand couch dune building. While the indirect inhibition by sand couch through elevation was found at both locations, the direct effects of sand couch on survival (i.e., neutral in exposed and positive in sheltered conditions) and shoot formation (i.e., negative in exposed and neutral in sheltered conditions) were location and thus context dependent. Similar to scale-dependent interactions found in other biogeomorphic landscapes (e.g., salt marshes, seagrass, mussel beds), we suggest that interspecific facilitation acts on larger spatial scales rather than the commonly suggested small-scale facilitation through created elevation. More specifically, we suggest that dune formation by pioneer species leads to the creation of stable and sheltered microsites beneficial for establishment of later successional species at the landscape-scale.