The pervasive impact of invasive species has motivated considerable research to understand how characteristics of invaded communities, such as native species diversity, affect the establishment of invasive species. Efforts to identify general mechanisms that limit invasion success, however, have been frustrated by disagreement between landscape-scale observations that generally find a positive relationship between native diversity and invasibility and smaller-scale experiments that consistently reveal competitive interactions that generate the opposite relationship. Here we experimentally elucidate the mechanism explaining the large-scale positive associations between invasion success and native intertidal diversity revealed in our landscape-scale surveys of New England shorelines. Experimental manipulations revealed this large-scale pattern is driven by a facilitation cascade where ecosystem-engineering species interact nonlinearly to enhance native diversity and invasion success by alleviating thermal stress and substrate instability. Our findings reveal that largescale diversity-invasion relationships can be explained by small-scale positive interactions that commonly occur across multiple trophic levels and functional groups. We argue that facilitation has played an important but unrecognized role in the invasion of other well studied systems, and will be of increasing importance with anticipated climate change.

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Thorough understanding of the potential for threshold dynamics and catastrophic shifts to occur in natural systems is of great importance for ecosystem conservation and restoration. However, verifying the presence of alternative stable states, one of the theoretical explanations for sudden shifts in natural systems, has proven to be a major challenge. We examine processes on local and landscape scales in salt-marsh pioneer zones, to assess the presence of alternative stable states in this system. To that end, we investigated the presence of typical characteristics of alternative stable states: bimodality and threshold dynamics. We also studied whether vegetation patches remained stable over long time periods. Analysis of false-color aerial photographs revealed clear bimodality in plant biomass distribution. By transplanting Spartina anglica plants of three different biomass classes on three geographically different marshes, we showed that a biomass threshold limits the establishment of Spartina patches, potentially explaining their patchy distribution. The presence of bimodality and biomass thresholds points to the presence of alternative stable states and the potential for sudden shifts, at small, within-patch scales and on short time scales. However, overlay analysis of aerial photographs from a salt marsh in The Netherlands, covering a time span of 22 years, revealed that there was little long-term stability of patches, as vegetation cover in this area is slowly increasing. Our results suggest that the concept of alternative stable states is applicable to the salt-marsh pioneer vegetation on small spatio-temporal scales. However, the concept does not apply to long-term dynamics of decades or centuries of heterogeneous salt-marsh pioneer zones, as landscape-scale processes may determine the large-scale dynamics of salt marshes. Hence, our results provide the interesting perspective that threshold dynamics may occur in systems with, on the long term, only a single stable state.@en

Though species interactions across local environmental gradients are well studied, the way in which species interactions shift between different habitats on a landscape scale has received less attention. We hypothesised that interactions among a suite of shoreline plant species shift across a hydrodynamic-exposure gradient, leading to generation of apparently distinct habitat types (e.g. bare cobble beaches, vegetated cobble beaches, fringing marshes and salt marshes). We examined hydrodynamic forcing and found that it was strongly correlated with shoreline habitat type. A transplant experiment revealed that all plants were rapidly crushed and abraded in bare cobble areas with high hydrodynamic energy and were out-competed by grasses in the low-energy salt marshes. Vegetated cobble beaches are inhabited by the largest amount of plant species under intermediate conditions, where hydrodynamic energy is sufficiently low to allow the establishment of the ecosystem engineer Spartina and sufficiently high to prevent the monopolisation of space by competitively dominant, but stress intolerant grasses. Experimentally reducing physical and biotic stresses (hydrodynamics and interspecific competition) on bare cobble beaches and salt marshes, respectively, enabled forbs to persist across the whole gradient. These results demonstrate that the outcome of interspecific interactions at landscape scales is driven by background physical conditions, and that this can result in the development of what are considered distinct habitats.@en