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There is substantial evidence that biodiversity underpins ecosystem functioning, but it is unclear how these relationships change with multiple stressors in complex real-world settings. Coastal zones are affected by numerous stressors (for example, sediment input and nutrient runoff from land) and the cumulative effects of these stressors may result in pronounced and unexpected changes in the functioning of ecosystems. To investigate the cumulative effects of turbidity and elevated nutrients on coastal biodiversity-ecosystem functioning relationships, we performed a large-scale field experiment manipulating in situ sediment porewater ammonium concentrations and measured multiple ecosystem functions related to carbon fixation and mineralisation in 15 estuaries with varying levels of turbidity. The results indicated that the benthic macrofauna diversity (species richness, abundance, and functional richness) declined with increased porewater ammonium concentrations and there were clear thresholds in light levels at the seafloor in relation to the biodiversity-ecosystem function relationships. Multifunctionality indices (an integrated index of all measured functions) in moderately turbid and turbid estuaries (daily mean seafloor PAR −2 s−1) decreased with the loss of macrofauna biodiversity. Functioning in low-turbidity estuaries (daily mean PAR > 420 µmol m−2 s−1) however remained relatively constant, suggesting that they were more resilient against the nutrient-induced biodiversity loss. Our results demonstrate that ecosystems already stressed by stressors that alter functional performance (turbidity) may be more prone to loss of overall functioning if biodiversity is reduced by another stressor (nutrient enrichment), highlighting the potential snowballing effects of cumulative change.

The biodiversity crisis has increased interest in understanding the role of biodiversity for ecosystem functioning. Functional traits are often used to infer ecosystem functions to increase our understanding of these relationships over larger spatial scales. The links between specific traits and ecosystem functioning are, however, not always well established. We investigated how the choice of analyzing either individual species, selected modalities, or trait combinations affected the spatial patterns observed on a sandflat and how this was related to the natural variability in ecosystem functioning. A large dataset of 400 benthic macrofauna samples was used to explore distribution patterns. We hypothesized that (1) if multiple species (redundancy) represent a trait combination or a modality their spatial patterns would be smoothed out, and (2) the lost spatial variability within a trait combination or modality, due to the smoothing effect, would potentially affect their utility for predicting ecosystem functioning (tested on a dataset of 24 samples). We predicted that species would show heterogeneous small spatial patterns, while modalities and trait combinations would show larger and more homogeneous patterns because they would represent a collection of many distributions. If modalities and trait combinations are better predictors of ecosystem functioning than species, then the smoother spatial patterns of modalities and trait combinations would result in a more homogeneous landscape of ecosystem function and the number of species exhibiting specific traits would provide functional redundancy. Our results showed some smoothing of spatial patterns progressing from species through modalities to trait combinations, but generally spatial patterns reflected a few dominant key species. Moreover, some individual modalities and species explained more or equal proportions of the variance in the ecosystem functioning than the combined traits. The findings thus suggest that only some spatial variability is lost when species are combined into modalities and trait combinations and that a homogeneous landscape of ecosystem function is not likely., Functional traits are often used to infer ecosystem functioning across larger scales, but how do they reflect species distributions and how well do they actually explain the natural spatial variability in ecosystem functioning? This was examined with a rarely seen extensive dataset of benthic macrofauna samples and solute flux measurements across the sediment–water interface.

Caio Graco-Roza was funded by the Coordination for the Improvement of Higher Education Personnel (CAPES), the Carlos Chagas Filho Research Support Foundation (FAPERJ) and the Ella and Georg Erhnrooth Foundation; Jan Altman by research grants INTER-EXCELLENCE LTAUSA19137 provided by Czech Ministry of Education, Youth and Sports, 20-05840Y of the Czech Science Foundation, and long-term research development project no. RVO 67985939 of the Czech Academy of Sciences; Otso Ovaskainen was funded by Academy of Finland (grant no. 309581), Jane and Aatos Erkko Foundation, Research Council of Norway through its Centres of Excellence Funding Scheme (223257), and the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement no. 856506; ERC-synergy project LIFEPLAN); and Jianjun Wang was funded by CAS Key Research Program of Frontier Sciences (QYZDB-SSW-DQC043) and National Natural Science Foundation of China (91851117). The "sPlot" project was initiated by sDiv, the Synthesis Centre of the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, funded by the German Research Foundation (DFG FZT 118), and is now a platform of iDiv. The study was supported by the TRY initiative on plant traits (). We are also grateful to Jens Kattge and TRY database. TRY is hosted, developed and maintained at the Max Planck Institute for Biogeochemistry (MPI-BGC) in Jena, Germany, in collaboration with the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig. The CESTES database of metacommunities is also an initiative of iDiv led by Alienor Jeliazkov. We thank sDiv for supporting the open science initiative., Aim: Understanding the variation in community composition and species abundances (i.e., β-diversity) is at the heart of community ecology. A common approach to examine β-diversity is to evaluate directional variation in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distance. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 148 datasets comprising different types of organisms and environments. Location: Global. Time period: 1990 to present. Major taxa studied: From diatoms to mammals. Method: We measured the strength of the decay using ranked Mantel tests (Mantel r) and the rate of distance decay as the slope of an exponential fit using generalized linear models. We used null models to test whether functional similarity decays faster or slower than expected given the taxonomic decay along the spatial and environmental distance. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm and organismal features. Results: Taxonomic distance decay was stronger than functional distance decay along both spatial and environmental distance. Functional distance decay was random given the taxonomic distance decay. The rate of taxonomic and functional spatial distance decay was fastest in the datasets from mid-latitudes. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distance but a higher rate of decay along environmental distance. Marine ecosystems had the slowest rate of decay along environmental distances. Main conclusions: In general, taxonomic distance decay is a useful tool for biogeographical research because it reflects dispersal-related factors in addition to species responses to climatic and environmental variables. Moreover, functional distance decay might be a cost-effective option for investigating community changes in heterogeneous environments., Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) Fundacao Carlos Chagas Filho de Amparo a Pesquisa do Estado do Rio De Janeiro (FAPERJ), Ella and Georg Erhnrooth Foundation, Ministry of Education, Youth & Sports - Czech Republic LTAUSA19137 Grant Agency of the Czech Republic 20-05840Y Czech Academy of Sciences RVO 67985939, Academy of Finland 309581, Jane and Aatos Erkko Foundation, Research Council of Norway through its Centres of Excellence Funding Scheme 223257, European Research Council (ERC) 856506, CAS Key Research Program of Frontier Sciences QYZDB-SSW-DQC043, National Natural Science Foundation of China (NSFC) 91851117, German Research Foundation (DFG) DFG FZT 118, TRY initiative on plant traits

The ongoing loss of biodiversity and global environmental changes severely affect the structure of coastal ecosystems. Consequences, in terms of ecosystem functioning, are, however, difficult to predict because the context dependency of the biodiversity-ecosystem function relationships within these heterogeneous seascapes is poorly understood. To assess the effects of biological and environmental factors in mediating ecosystem functioning (nutrient cycling) in different natural habitats, intact sediment cores were collected at 18 sites on a grain size gradient from coarse sand to silt, with varying organic matter content and vegetation. To assess ecosystem functioning, solute fluxes (O-2, NH4+, PO43-, Si) across the sediment-water interface were measured. The macrofaunal communities changed along the grain size gradient with higher abundance, biomass and number of species in coarser sediments and in habitats with more vegetation. Across the whole gradient, the macrofauna cumulatively accounted for 25% of the variability in the multivariate solute fluxes, whereas environmental variables cumulatively accounted for 20%. Only the biomass and abundance of a few of the most dominant macrofauna species, not the number of species, appeared to contribute significantly to the nutrient recycling processes. Closer analyses of different sediment types (grouped into coarse, medium and fine sediment) showed that the macrofauna was an important predictor in all sediment types, but had the largest impact in fine and medium sediments. The results imply that even if the ecosystem functioning is similar in different sediment types, the underpinning mechanisms are different, which makes it challenging to generalize patterns of functioning across the heterogeneous shallow coastal zones.

The current decrease in biodiversity affects all ecosystems, and the impacts of diversity on ecosystem functioning need to be resolved. So far, marine studies about diversity?ecosystem productivity-relationships have concentrated on small-scale, controlled experiments, with often limited relevance to natural ecosystems. Here, we provide a real-world study on the effects of microorganismal diversity (measured as the diversity of benthic diatom communities) on ecosystem productivity (using chlorophyll a concentration as a surrogate) in a heterogeneous marine coastal archipelago. We collected 78 sediment cores at 17 sites in the northern Baltic Sea and found exceptionally high diatom diversity (328 observed species). We used structural equation models and quantile regression to explore relationships between diatom diversity and productivity. Previous studies have found contradictory results in the relationship between microorganismal diversity and ecosystem productivity, but we showed a linear and positive basal relationship between diatom diversity and productivity, which indicates that diatom diversity most likely forms the lowest boundary for productivity. Thus, although productivity can be high even when diatom diversity is low, high diatom diversity supports high productivity. The trait composition was more effective than taxonomical composition in showing such a relationship, which could be due to niche complementarity. Our results also indicated that environmental heterogeneity leads to substantial patchiness in the diversity of benthic diatom communities, mainly induced by the variation in sediment organic matter content. Therefore, future changes in precipitation and river runoff and associated changes in the quality and quantity of organic matter in the sea, will also affect diatom communities and, hence, ecosystem productivity. Our study suggests that benthic microorganisms are vital for ecosystem productivity, and together with the substantial heterogeneity of coastal ecosystems, they should be considered when evaluating the potential productivity of coastal areas.

Bioturbation by benthic macrofauna communities plays a significant role in the setting and maintenance of important ecosystem functions and the delivery of associated ecosystem services. We investigated the context-dependence of bioturbation performed by natural benthic communities in the coastal northern Baltic Sea by quantifying three bioturbation metrics (particle mixing intensity, surface sediment reworking and bioturbation depth) across 18 sites ranging from cohesive muddy sediments to non-cohesive coarse sands, while accounting for the complexity of natural communities and habitat characteristics. We identified two distinct patterns of bioturbation; in fine sediments bioturbation rates were highly variable and in coarse sediments bioturbation rates were less variable and characterized by lower maximal values. Using distance-based linear multiple regressions, we found that 75.5% of the variance in bioturbation rates in fine sediment could be explained by key functional groups/species abundance and/or biomass (i.e. biomass of the gallery-diffusors and abundances of biodiffusors, surface modifiers, conveyors and gallery diffusors, respectively). In coarse sediment, 47.8% of the variance in bioturbation rates could be explained by a combination of environmental factors (grain size, organic matter content, buried plant material) and faunal functional groups, although fauna alone explained only 13% of this variance. Bioturbation in fine sediments was therefore more predictable based on the composition of benthic fauna. In coarse sediment, the bioturbation activities of benthic fauna were strongly modified by habitat characteristics (including the presence of buried plant material, sediment organic content and grain size) whereas in fine sediments this was not the case. Our results therefore highlight that variability in spatial patterns of bioturbation is a result of complex relationships between macrofauna community structure, sediment type and other habitat characteristics, likely modifying bioturbation performance of individual fauna.

Coastal ecosystems are important because of the vital ecosystem functions and services they provide, but many are threatened by eutrophication and hypoxia. This results in loss of biodiversity and subsequent changes in ecosystem functioning. Consequently, the need for empirical field studies regarding biodiversity-ecosystem functioning in coastal areas has been emphasized. The present field study quantified the links between benthic macrofaunal communities (abundance, biomass, and species richness), sediment oxygen consumption, and solute fluxes (NO3 − + NO2 −, NH4 +, PO4 3−, SiO4, Fe, Mn) along a 7.5-km natural gradient of seasonal hypoxia in the coastal northern Baltic Sea. Sampling was done in late August 2010 in the middle archipelago zone of the Hanko peninsula, Finland. As predicted, the macrofaunal communities were decimated with increasing hypoxia, and the nutrient transformation processes were changed at the sediment-water interface, with notably higher effluxes of phosphate and ammonium from the sediment. Solute fluxes varied even during normoxia, which implies a high context-dependency, and could be explained by even small variations in environmental variables such as organic matter and C/N ratios. Importantly, the low diversity benthic macrofaunal communities, which were dominated by Macoma balthica and the invasive Marenzelleria spp., had a large influence on the solute fluxes, especially under normoxia, but also under hypoxia.

Seafloor ecosystems play pivotal roles in biogeochemical cycling, but hypoxia (O2 1200 km). This broad-scale sampling confirmed the strong salinity-driven diversity gradient and large spatial variations in oxygen conditions, which affected the status of the benthic communities. Benthic nutrient fluxes varied several orders of magnitude, both between sea areas and along gradients of hypoxia within areas. DistLM modeling indicated that benthic fauna can affect nutrient cycling, also under hypoxic conditions, with the invasive polychaete Marenzelleria spp. being particularly important. However, as the oxygen content decreases—even slightly—the subsequent changes in the faunal abundance and functional diversity may influence the nature and the rate of fluxes. These results imply that management targets for oxygen concentrations need to be raised to sustain healthy ecosystem functioning and to facilitate the recovery of large degraded sea areas.

Marine ecosystems world-wide are threatened by oxygen deficiency, with potential serious consequences for ecosystem functioning and the goods and services they provide. While the effects of hypoxia on benthic species diversity are well documented, the effects on ecosystem function have only rarely been assessed in real-world settings. To better understand the links between structural changes in macro- and meiofaunal communities, hypoxic stress and benthic ecosystem function (benthic nutrient fluxes, community metabolism), we sampled a total of 11 sites in Haystensfjord and Askerofjord (Swedish west coast) in late summer, coinciding with the largest extent and severity of seasonal hypoxia in the area. The sites spanned oxic to anoxic bottom water, and a corresponding gradient in faunal diversity. Intact sediment cores were incubated to measure fluxes of oxygen and nutrients (NO3-, NO2-, NH4+, PO43-, SiO4) across the sediment-water interface. Sediment profile imaging (SPI) footage was obtained from all sites to assess structural elements and the bioturbadon depth, and additional samples were collected to characterise sediment properties and macro- and meiofaunal community composition. Bottom-water O-2 concentration was the main driver of macrofauna communities, with highest abundance and biomass, as well as variability, at the sites with intermediate O-2 concentration. Meiofauna on the other hand was less sensitive to bottom-water O-2 concentration. Oxygen was the main driver of nutrient fluxes too, but macrofauna as well meiofauna were also significant predictors; DistLM analyses indicated that O-2 concentration, macrofaunal abundance or biomass, and meiofaunal abundance collectively explained 63%, 30% and 28% of the variation in sediment O-2 consumption, NH4+ flux and PO43+ flux, respectively. The study provides a step towards a more realistic understanding of the link between benthic fauna and ecosystem functioning, and the influence of disturbance on this relationship, which is important for management decisions aimed at protecting the dwindling biodiversity in the coastal zones around the world.