Lifeforms ranging from bacteria to humans employ specialized random movement patterns. Although effective as optimization strategies in many scientific fields, random walk application in biology has remained focused on search optimization by mobile organisms. Here, we report on the discovery that heavy-tailed random walks underlie the ability of clonally expanding plants to self-organize and dictate the formation of biogeomorphic landscapes. Using cross-Atlantic surveys, we show that congeneric beach grasses adopt distinct heavy-tailed clonal expansion strategies. Next, we demonstrate with a spatially explicit model and a field experiment that the Lévy-type strategy of the species building the highest dunes worldwide generates a clonal network with a patchy shoot organization that optimizes sand trapping efficiency. Our findings demonstrate Lévy-like movement in plants, and emphasize the role of species-specific expansion strategies in landscape formation. This mechanistic understanding paves the way for tailor-made planting designs to successfully construct and restore biogeomorphic landscapes and their services.

Mounting evidence shows that the functioning and stability of coastal ecosystems often depends critically on habitat‐forming foundation species such as seagrasses, mangroves and saltmarsh grasses that engage in facultative mutualistic interactions. However, although restoration science is now gradually expanding its long‐standing paradigm of minimizing competition to including intraspecific, or within species, facilitation in its designs, the potential of harnessing mutualistic interactions between species for restoration purposes remains uninvestigated. Here, we experimentally tested whether a previously documented mutualism between marsh‐forming Spartina alterniflora (cordgrass) and Geukensia demissa (mussels) can increase restoration success in degraded US saltmarshes. We found that co‐transplanted mussels locally increased nutrients and reduced sulphide stress, thereby increasing cordgrass growth and clonal expansion by 50%. We then removed above‐ground vegetation and mussels to simulate a disturbance event and discovered that cordgrass co‐transplanted with mussels experienced three times greater survival than control transplants. Synthesis and applications. Our findings indicate that mussels amplify cordgrass re‐colonization and resilience over spatial and temporal scales that exceed those of their actual mutualistic interaction. By experimentally demonstrating that mutualistic partners can enable foundation species to overcome stress barriers to establish and persist, we highlight that coastal restoration needs to evolve beyond the sole inclusion of intraspecific‐positive interactions. In particular, we suggest that integrating mutualisms in restoration designs may powerfully enhance long‐term restoration success and ecosystem resilience in the many coastal ecosystems where mutualisms involving foundation species are important ecosystem‐structuring interactions.

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Food webs are an integral part of every ecosystem on the planet, yet understanding the mechanisms shaping these complex networks remains a major challenge. Recently, several studies suggested that non-trophic species interactions such as habitat modification and mutualisms can be important determinants of food web structure. However, it remains unclear whether these findings generalize across ecosystems, and whether non-trophic interactions affect food webs randomly, or affect specific trophic levels or functional groups. Here, we combine analyses of 58 food webs from seven terrestrial, freshwater and coastal systems to test (1) the general hypothesis that non-trophic facilitation by habitat-forming foundation species enhances food web complexity, and (2) whether these enhancements have either random or targeted effects on particular trophic levels, functional groups, and linkages throughout the food web. Our empirical results demonstrate that foundation species consistently enhance food web complexity in all seven ecosystems. Further analyses reveal that 15 out of 19 food web properties can be well-approximated by assuming that foundation species randomly facilitate species throughout the trophic network. However, basal species are less strongly, and carnivores are more strongly facilitated in foundation species' food webs than predicted based on random facilitation, resulting in a higher mean trophic level and a longer average chain length. Overall, we conclude that foundation species strongly enhance food web complexity through non-trophic facilitation of species across the entire trophic network. We therefore suggest that the structure and stability of food webs often depends critically on non-trophic facilitation by foundation species.

Salt marshes suffered large‐scale degradation in recent decades. Extreme events such as hot and dry spells contributed significantly to this, and are predicted to increase not only in intensity, but also in frequency under future climate scenarios. Such repetitive extreme events may generate cumulative effects on ecosystem resilience. It is therefore important to elucidate how marsh vegetation responds to repetitive stress, and whether changes in key species interactions can modulate vegetation resilience. In this study, we investigated how moderate but repetitive desiccation events, caused by the combined effects of drought and high temperatures, affect cordgrass (Spartina alterniflora), the dominant habitat‐forming grass in southeastern US salt marshes. In a 4‐month field experiment, we simulated four consecutive desiccation events by periodically excluding tidal flooding and rainfall, while raising temperature. We crossed this desiccation treatment with the presence/absence of ribbed mussels (Geukensia demissa) – a mutualist of cordgrass known to enhance its desiccation resilience – and with grazing pressure by the marsh periwinkle (Littoraria irrorata) that is known to suppress cordgrass’ desiccation resilience. We found that each subsequent desiccation event deteriorated sediment porewater conditions, resulting in high salinity (53 ppt), low pH‐levels (3.7) and increased porewater Al and Fe concentrations (≈800 μmol/L and ≈1,500 μmol/L) upon rewetting. No effects on porewater chemistry were found as a result of snail grazing, while ribbed mussels strongly mitigated desiccation effects almost to control levels and increased cordgrass biomass by approximately 128%. Importantly, although cordgrass generally appeared healthy above‐ground at the end of the experiment, we found clear negative responses of the repetitive desiccation treatment on cordgrass below‐ground biomass, on proline (osmolyte) levels in shoots and on the number of tillers (−40%), regardless of mussel and/or snail presence. Synthesis. Even though the mutualism with mussels strongly mitigated chemical effects in the sediment porewater throughout the experiment, mussels could not buffer the adverse ecophysiological effects observed in cordgrass tissue. Our results therefore suggest that although mussels may alleviate desiccation stress, the predicted increased frequency and intensity of hot dry spells may eventually affect saltmarsh resilience by stressing the mutualism beyond its buffering capacity.