Although there is mounting evidence that biodiversity is an important and widespread driver of ecosystem multifunctionality, much of this research has focused on small-scale biodiversity manipulations. Hence, which mechanisms maintain patches of enhanced biodiversity in natural systems and if these patches elevate ecosystem multifunctionality at both local and landscape scales remain outstanding questions. In a 17 month experiment conducted within southeastern United States salt marshes, we found that patches of enhanced biodiversity and multifunctionality arise only where habitat-forming foundation species overlap-i.e. where aggregations of ribbed mussels (Geukensia demissa) form around cordgrass (Spartina alterniflora) stems. By empirically scaling up our experimental results to the marsh platform at 12 sites, we further show that mussels-despite covering only approximately 1% of the marsh surface-strongly enhance five distinct ecosystem functions, including decomposition, primary production and water infiltration rate, at the landscape scale. Thus, mussels create conditions that support the co-occurrence of high densities of functionally distinct organisms within cordgrass and, in doing so, elevate salt marsh multifunctionality from the patch to landscape scale. Collectively, these findings suggest that patterns in foundation species' overlap drive variation in biodiversity and ecosystem functioning within and across natural ecosystems. We therefore argue that foundation species should be integrated in our conceptual understanding of forces that moderate biodiversity -ecosystem functioning relationships, approaches for conserving species diversity and strategies to improve the multifunctionality of degraded ecosystems.

Droughts are increasing in severity and frequency, yet the mechanisms that strengthen ecosystem resilience to this stress remain poorly understood. Here, we test whether positive interactions in the form of a mutualism between mussels and dominant cordgrass in salt marshes enhance ecosystem resistance to and recovery from drought. Surveys spanning 250 km of southeastern US coastline reveal spatially dispersed mussel mounds increased cordgrass survival during severe drought by 5- to 25-times. Surveys and mussel addition experiments indicate this positive effect of mussels on cordgrass was due to mounds enhancing water storage and reducing soil salinity stress. Observations and models then demonstrate that surviving cordgrass patches associated with mussels function as nuclei for vegetative re-growth and, despite covering only 0.1–12% of die-offs, markedly shorten marsh recovery periods. These results indicate that mutualisms, in supporting stress-resistant patches, can play a disproportionately large, keystone role in enhancing ecosystem resilience to climatic extremes.

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.

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.

Rates of dark respiration and net photosynthesis were measured for six replicate clonal fragments of the stony coral Galaxea fascicularis (Linnaeus 1767), which were incubated under 12 different combinations of dissolved oxygen (20%, 100% and 150% saturation), dissolved carbon dioxide (9.5 and 19.1 µmol l −1 ) and water flow (1-1.6 versus 4-13 cm s) in a repeated measures design. Dark respiration was enhanced by increased flow and increased oxygen saturation in an interactive way, which relates to improved oxygen influx into the coral tissue. Oxygen saturation did not influence net photosynthesis: neither hypoxia nor hyperoxia affected net photosynthesis, irrespective of flow and pH, which suggests that hyperoxia does not induce high rates of photorespiration in this coral. Flow and pH had a synergistic effect on net photosynthesis: at high flow, a decrease in pH stimulated net photosynthesis by 14%. These results indicate that for this individual of G. fascicularis, increased uptake of carbon dioxide rather than increased efflux of oxygen explains the beneficial effect of water flow on photosynthesis. Rates of net photosynthesis measured in this study are among the highest ever recorded for scleractinian corals and confirm a strong scope for growth.

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.

No abstract

Foundation species are typically suggested to enhance community diversity non‐trophically by increasing habitat structure and mitigating physical stress, while their trophic role is considered of minor importance. Yet, there is little experimental evidence on the relative importance of trophic and non‐trophic effects and the interaction with patch size. Here, we transplanted different festoon sizes of living Tillandsia usneoides (Spanish moss) and structural mimics assessing the trophic and non‐trophic roles of this habitat‐forming epiphyte in mediating the invertebrate community. Compared to bare branches, mimics enhanced species and feeding guild richness and abundances, but living festoons even more so, demonstrating that trophic and non‐trophic effects jointly stimulated the community. Specifically, our results show that, independent of patch size, 40% of the total species richness and 46% of total guild richness increase could be contributed to habitat structure alone, while Spanish moss trophically stimulated these metrics by another 60% and 54%. As detritivores were particularly enhanced in living festoons, our findings suggest that trophic stimulation occurred primarily through the provisioning of Spanish moss detritus. Our results highlight that foundation species can facilitate their associated communities through both trophic and non‐trophic pathways, calling for studies addressing their indirect trophic role via the brown food web.