Ecological network analysis of traits reveals variable response capacity to stress.

Response diversity increases the potential 'options' for ecological communities to respond to stress (i.e. response capacity). An indicator of community response diversity is the diversity of different traits associated with their capacity to be resistant to stress, to recover and to regulate ecosystem functions. We conducted a network analysis of traits using benthic macroinvertebrate community data from a large-scale field experiment to explore the loss of response diversity along environmental gradients. We elevated sediment nutrient concentrations (a process that occurs with eutrophication) at 24 sites (in 15 estuaries) with varying environmental conditions (water column turbidity and sediment properties). Macroinvertebrate community response capacity to nutrient stress was dependent on the baseline trait network complexity in the ambient community (i.e. non-enriched sediments). The greater the complexity of the baseline network, the less variable the network response to nutrient stress was; in contrast, more variable responses to nutrient stress occurred with simpler networks. Thus, stressors or environmental variables that shift baseline network complexity also shift the capacity for these ecosystems to respond to additional stressors. Empirical studies that explore the mechanisms responsible for loss of resilience are essential to inform our ability to predict changes in ecological states.

Toward a network perspective in coastal ecosystem management.

Environmental management in coastal ecosystems has been challenged by the complex cumulative effects that occur when many small issues result in large ecological shifts. Current environmental management of these spaces focuses on identifying and limiting problematic stressors via a series of assessment techniques. Whilst there is a strong desire among managers to consider complexity in ecological responses to cumulative effects, current approaches for assessing risk focus on breaking down the issues into multiple cause and effect relationships. However, uncertainty arises when data and information for a place are limited, as is commonly the case, and this creates decision paralysis while more information is generated. Here, we discuss how ecological understanding of network interactions in coastal marine ecosystems can be used as a lens to bring together multiple lines of evidence and create actions. We list and describe four characteristics of marine ecosystem interaction networks including the possibility for; 1) indirect effects, 2) effects that emerge as stressor magnitude increases the number of network components implicated, 3) network interactions that amplify these indirect effects, and 4) feedbacks that reinforce or stabilise against indirect effects. We then link these four characteristics to three case studies of common coastal environmental issues to demonstrate how a general understanding of ecological interaction networks can enhance priorities for stressor management that can be applied even when specific data is limited.

Modelling interactive effects of multiple disturbances on a coastal lake ecosystem: Implications for management.

Coastal lakes, also known as temporarily open/closed estuaries or intermittently closed and open lakes and lagoons, are common worldwide, are typically sites of high biodiversity and often contain abundant macrophyte populations. Anthropogenic stressors such as increased nutrient and sediment loading have adverse effects on submerged macrophytes, and when closed, the lack of tidal flushing makes coastal lakes highly susceptible to eutrophication. Lake openings to the sea may occur naturally, but many coastal lakes are also opened artificially, often to reduce inundation of surrounding land. Here we used a coupled hydrodynamic-ecological model (DYRESM-CAEDYM), modified to include dynamic feedback between submerged macrophyte biomass and sediment resuspension, to explore the interactive effects of multiple disturbances (openings, eutrophication and climate change) on the dynamics of primary producers in a coastal lake (Waituna Lagoon) in South Island, New Zealand. Our results indicate that with exposure to high external nutrient loads, the frequent disturbances caused by artificial openings prevent sustained dominance by algae (algal biomass averaged 192 g C m-2 with artificial openings compared to 453 g C m-2 with no openings). However, under current nutrient loading, climate change is likely to enhance the effects of eutrophication on the system (algal biomass averaged 227 g C m-2 with climate change compared with 192 g C m-2 for current climate). The model provides a decision-support tool to guide lake management in setting limits for nutrient loads and managing the opening regime, in order to prevent eutrophication and the potential collapse of the macrophyte community.

Changes in the location of biodiversity-ecosystem function hot spots across the seafloor landscape with increasing sediment nutrient loading.

Declining biodiversity and loss of ecosystem function threatens the ability of habitats to contribute ecosystem services. However, the form of the relationship between biodiversity and ecosystem function (BEF) and how relationships change with environmental change is poorly understood. This limits our ability to predict the consequences of biodiversity loss on ecosystem function, particularly in real-world marine ecosystems that are species rich, and where multiple ecosystem functions are represented by multiple indicators. We investigated spatial variation in BEF relationships across a 300 000 m2 intertidal sandflat by nesting experimental manipulations of sediment pore water nitrogen concentration into sites with contrasting macrobenthic community composition. Our results highlight the significance of many different elements of biodiversity associated with environmental characteristics, community structure, functional diversity, ecological traits or particular species (ecosystem engineers) to important functions of coastal marine sediments (benthic oxygen consumption, ammonium pore water concentrations and flux across the sediment-water interface). Using the BEF relationships developed from our experiment, we demonstrate patchiness across a landscape in functional performance and the potential for changes in the location of functional hot and cold spots with increasing nutrient loading that have important implications for mapping and predicating change in functionality and the concomitant delivery of ecosystem services.

Post-settlement dispersal: the neglected link in maintenance of soft-sediment biodiversity.

Seafloor integrity is threatened by disturbances owing to human activities. The capacity of the system to recover from disturbances, as well as maintain resilience and function, depends on dispersal. In soft-sediment systems, dispersal continues after larval settlement, but there are very few measurements of how far the post-settlers disperse in nature. Spatial scales of post-settlement dispersal are, however, likely to be similar to pelagic larval dispersal because of continued, frequent, small-scale dispersal over longer periods. The consequences of this dispersal may be more important for the maintenance of biodiversity and metacommunity dynamics than is pelagic larval dispersal, because of the greater size and competency of the dispersers. We argue that an increased empirical understanding of post-settlement dispersal processes is key for predicting how benthic communities will respond to local disturbances and shrinking regional species pools, with implications for monitoring, managing and conserving biodiversity.

Effects of catchment land use on temperate mangrove forests.

Human land use changes are threatening the integrity and health of coastal ecosystems worldwide. Intensified land use for anthropogenic purposes increases sedimentation rates, pollutants, and nutrient concentrations into adjacent coastal areas, often with detrimental effects on marine life and ecosystem functioning. However, how these factors interact to influence ecosystem health in mangrove forests is poorly understood. This study investigates the effects of catchment human land use on mangrove forest architecture and sedimentary attributes at a landscape-scale. Thirty sites were selected along a gradient of human land use within a narrow latitudinal range, to minimise the effects of varying climatic conditions. Land use was quantified using spatial analysis tools with existing land use databases (LCDB5). Twenty-six forest architectural and sedimentary variables were collected from each site. The results revealed a significant effect of human land use on ten out of 26 environmental variables. Eutrophication, characterised by changes in redox potential, pH, and sediment nutrient concentrations, was strongly associated with increasing human land use. The δ15N values of sediments and leaves also indicated increased anthropogenic nitrogen input. Furthermore, the study identified a positive correlation between human land use and tree density, indicating that increased nutrient delivery from catchments contributes to enhanced mangrove growth. Propagule and seedling densities were also positively correlated with human land use, suggesting potential recruitment success mechanisms. This research underpins the complex interactions between human land use and mangrove ecosystems, revealing changes in carbon dynamics, potential alterations in ecosystem services, and a need for holistic management approaches that consider the interconnectedness of species and their environment. These findings provide essential insights for regional ecosystem models, coastal management, and restoration strategies to address the impacts of human pressures on temperate mangrove forests, even in estuaries that may be relatively healthy.

Diurnal fluctuations in seawater pH influence the response of a calcifying macroalga to ocean acidification.

Coastal ecosystems that are characterized by kelp forests encounter daily pH fluctuations, driven by photosynthesis and respiration, which are larger than pH changes owing to ocean acidification (OA) projected for surface ocean waters by 2100. We investigated whether mimicry of biologically mediated diurnal shifts in pH-based for the first time on pH time-series measurements within a kelp forest-would offset or amplify the negative effects of OA on calcifiers. In a 40-day laboratory experiment, the calcifying coralline macroalga, Arthrocardia corymbosa, was exposed to two mean pH treatments (8.05 or 7.65). For each mean, two experimental pH manipulations were applied. In one treatment, pH was held constant. In the second treatment, pH was manipulated around the mean (as a step-function), 0.4 pH units higher during daylight and 0.4 units lower during darkness to approximate diurnal fluctuations in a kelp forest. In all cases, growth rates were lower at a reduced mean pH, and fluctuations in pH acted additively to further reduce growth. Photosynthesis, recruitment and elemental composition did not change with pH, but δ(13)C increased at lower mean pH. Including environmental heterogeneity in experimental design will assist with a more accurate assessment of the responses of calcifiers to OA.

Diversity and distributional patterns of ciliates in Guaymas Basin hydrothermal vent sediments.

Little is known about protists at deep-sea hydrothermal vents. The vent sites at Guaymas Basin in the Gulf of California are characterized by dense mats of filamentous pigmented or nonpigmented Beggiatoa that serve as markers of subsurface thermochemical gradients. We constructed 18S rRNA libraries to investigate ciliate assemblages in Beggiatoa mats and from bare sediments at the Guaymas vent site. Results indicated a high diversity of ciliates, with 156 operational taxonomic units identified in 548 sequences. Comparison between mat environments demonstrated that ciliate and bacterial assemblages from pigmented mats, nonpigmented mats, and bare sediments were significantly different and highly correlated with bacterial assemblages. Neither bacterial nor ciliate assemblages were correlated with environmental factors. The most abundant ciliates at Guaymas were more likely to be represented in clone libraries from other hydrothermal, deep-sea, and/or anoxic or microaerophilic environments, supporting the hypothesis that these ciliate species are broadly distributed. The orange mat environment included a higher proportion of ciliate sequences that were more similar to those from other environmental studies than to cultured ciliate species, whereas clone libraries from bare sediments included sequences that were the most highly divergent from all other sequences and may represent species that are endemic to Guaymas.

Estimating Effects of Sea Level Rise on Benthic Biodiversity and Ecosystem Functioning in a Large Meso-Tidal Coastal Lagoon.

Estuaries are among the world's most productive ecosystems, but due to their geographic location, they are at the forefront of anthropogenic pressures. Sea level rise (SLR) is one major consequence of climate change that poses a threat to estuaries with extensive intertidal habitats. The ecological implications of intertidal habitat loss have been largely overlooked despite their likely significance. We aimed to address this knowledge gap by investigating how benthic macroinvertebrate communities and their contributions to ecosystem function are likely to respond to SLR. Based on a spatially extensive dataset (119 sites) from a large coastal lagoon, depth, sediment chlorophyll concentrations, mud content, and average current speed were identified as the main drivers of community compositional turnover. Shifts in benthic community structure and associated functional implications were then evaluated using depth as a proxy for SLR. Three main macrofaunal groups representing intertidal, shallow subtidal, and deep subtidal habitats were identified. Functional trait analysis indicated low functional redundancy for a key intertidal suspension-feeding bivalve (Austrovenus stutchburyi) and the lack of a shallow subtidal functional replacement should intertidal habitats become inundated. These findings strongly suggest SLR and the associated environmental changes will alter estuarine macroinvertebrate communities, with implications for future ecosystem function and resilience.

Mitochondrial DNA (COI) analyses reveal that amphipod diversity is associated with environmental heterogeneity in deep-sea habitats.

The relationship between species diversity and environmental parameters is poorly understood for the mobile macrofauna of deep-sea habitats due to under-sampling and subsequent lack of accurate taxonomic information. To redress this, cytochrome oxidase c subunit I (COI) DNA sequences were used to estimate species diversity and to compare phoxocephalid amphipod assemblages among 20 stations encompassing a range of environmental conditions. Two regions, east (Chatham Rise) and west (Challenger Plateau) of New Zealand were sampled to depths of 200-1200 m with an epibenthic sled. Using a comparison among identified morphospecies, we found a clear gap in sequence divergences between 6% and 13% and used a 6% threshold to designate molecular operational taxonomic units (MOTUs), as a surrogate to putative species. DNA sequences (n = 297) revealed high total diversity (n = 49 MOTUs), as well as high beta diversity (28 MOTUs found at single location only). Novel phoxocephalid MOTUs were found at most stations, especially on Challenger Plateau and the flanks of Chatham Rise. Analyses of interstation assemblages revealed a major split between regions, indicating minimal overlap in taxon distributions. A cluster of highly similar stations was identified, broadly distributed over the crest of Chatham Rise, in association with elevated food availability, probably resulting from higher surface productivity and relatively shallow depth. Accordingly, multivariate analysis revealed a strong correlation between phoxocephalid assemblages and food supply. This study highlights the value of molecular approaches, in particular COI sequences, for quantifying and comparing diversity in under-sampled and/or under-studied taxa.

Who is contributing where? Predicting ecosystem service multifunctionality for shellfish species through ecological principles.

A key challenge in environmental management is determining how to manage multiple ecosystem services (ES) simultaneously, to ensure efficient and sustainable use of the environment and its resources. In marine environments, the spatial assessment of ES is lagging as a result of data-scarcity and modelling complexity. Applying mechanistic models to link ecological processes with ecosystem functions and services to assess areas of high ES potential can bridge this gap and accommodate assessments of functional differences between service providers. Here, we applied an ecosystem principles approach to assess ES potential for food provision, water quality regulation, nitrogen removal, and sediment stabilisation, provided by two estuarine bivalves (Austrovenus stutchburyi and Paphies australis) that differ in habitat association (broad and narrow distributions), to gain insight into the utility of these models for local-scale management. Maps of individual ES displayed differing patterns related to habitat associations of the species providing them, with variation in the quantities of services being delivered and locations of importance. Areas of importance for the provision of multiple services (number of services provided and their combined intensity per species) were assessed using hotspot analyses, which suggested that areas of high shellfish density at the harbour entrances were important for ES multifunctionality. A targeted management approach that includes environmental context, rather than a focus solely on the protection of high-density shellfish areas, is required to sustain the provision of individual ES.

Altered nitrogen transformation pathways and a legacy of sediment organic matter enrichment.

Estuaries are ecologically valuable ecosystems that process nutrients through complex biogeochemical processes. Here we identify drivers and inhibitors of nitrogen removal in unvegetated intertidal sandflats at six sites in Manukau Harbour (37° 2.00'S 174° 42.00'E) to quantify the exchange of solutes across the sediment-water interface, with nitrogen removal rates (NRR) measured at two contrasting sites (PI and CB) near and far, respectively, from an historical wastewater treatment plant. Solute fluxes were paired with sediment and macrofauna community data to identify drivers of ecosystem function. Fluxes of oxygen, nitrogen and phosphorous were found to vary among sites, with differences attributed to variation in sediment characteristics (grain size, chlorophyll a, organic content) and macrofauna community structure. Mean NRR was found to vary between sites (PI = 32.2 vs CB = 217.9 µmol N2/m2/h), with bioturbating macrofauna (bivalves Austrovenus stutchburyi and Macomona liliana), microphytobenthic biomass, and exposure to nutrients likely key contributing drivers.

The impact of cumulative stressor effects on uncertainty and ecological risk.

To enable environmental management actions to be more effectively prioritized, cumulative effects between multiple stressors need to be accounted for in risk-assessment frameworks. Ecological risk and uncertainty are generally high when multiple stressors occur. In the face of high uncertainty, transparent communication is essential to inform decision-making. The impact of stressor interactions on risk and uncertainty was assessed using generalized linear models for additive and multiplicative effect of six anthropogenic stressors on the abundance of estuarine macrofauna across New Zealand. Models that accounted for multiplicative stressor interactions demonstrated that non-additive effects dominated, had increased explanatory power (6 to 73 % relative increase between models), and thereby reduced the risk of unexpected ecological responses to stress. Secondly, 3D-plots provide important insights in the direction, magnitude and gradients of change, and aid transparency and communication of complex stressor effects. Notably, small changes in a stressor can cause a disproportionally steep gradient of change for a synergistic effect where the tolerance to stressors are lost, and would invoke precautionary management. 3D-plots were able to clearly identify directional shifts where the nature of the interaction changed from antagonistic to synergistic along increasing stressor gradients. For example, increased nitrogen load and exposure caused a shift from positive to negative effect on the abundance of a deposit-feeding polychaete (Magelona). Assessments relying on model coefficient estimates, which provide one effect term, could not capture the complexities observed in 3D-plots and are at risk of mis-identifying interaction types. Finally, visualising model uncertainty demonstrated that although error terms were higher for multiplicative models, they better captured the uncertainty caused by data availability. Together, the steep gradients of change identified in 3D-plots and the higher uncertainty in model predictions in multiplicative models urges more conservative limits to be set for management that account for risk and uncertainty in multiple stressor effects.

Biogeochemical feedbacks to ocean acidification in a cohesive photosynthetic sediment.

Ecosystem feedbacks in response to ocean acidification can amplify or diminish diel pH oscillations in productive coastal waters. Benthic microalgae generate such oscillations in sediment porewater and here we ask how CO2 enrichment (acidification) of the overlying seawater alters these in the absence and presence of biogenic calcite. We placed a 1-mm layer of ground oyster shells, mimicking the arrival of dead calcifying biota (+Calcite), or sand (Control) onto intact silt sediment cores, and then gradually increased the pCO2 in the seawater above half of +Calcite and Control cores from 472 to 1216 µatm (pH 8.0 to 7.6, CO2:HCO3- from 4.8 to 9.6 × 10-4). Porewater [O2] and [H+] microprofiles measured 16 d later showed that this enrichment had decreased the O2 penetration depth (O2-pd) in +Calcite and Control, indicating a metabolic response. In CO2-enriched seawater: (1) sediment biogeochemical processes respectively added and removed more H+ to and from the sediment porewater in darkness and light, than in ambient seawater increasing the amplitude of the diel porewater [H+] oscillations, and (2) in darkness, calcite dissolution in +Calcite sediment decreased the porewater [H+] below that in overlying seawater, reversing the sediment-seawater H+ flux and decreasing the amplitude of diel [H+] oscillations. This dissolution did not, however, counter the negative effect of CO2 enrichment on O2-pd. We now hypothesise that feedback to CO2 enrichment-an increase in the microbial reoxidation of reduced solutes with O2-decreased the sediment O2-pd and contributed to the enhanced porewater acidification.

Biodiversity breakpoints along stress gradients in estuaries and associated shifts in ecosystem interactions.

Denitrification in coastal sediments can provide resilience to eutrophication in estuarine ecosystems, but this key ecosystem function is impacted directly and indirectly by increasing stressors. The erosion and loading of fine sediments from land, resulting in sedimentation and elevated sediment muddiness, presents a significant threat to coastal ecosystems worldwide. Impacts on biodiversity with increasing sediment mud content are relatively well understood, but corresponding impacts on denitrification are uncharacterised. Soft sediment ecosystems have a network of interrelated biotic and abiotic ecosystem components that contribute to microbial nitrogen cycling, but these components (especially biodiversity measures) and their relationships with ecosystem functions are sensitive to stress. With a large dataset spanning broad environmental gradients this study uses interaction network analysis to present a mechanistic view of the ecological interactions that contribute to microbial nitrogen cycling, showing significant changes above and below a stressor (mud) threshold. Our models demonstrate that positive biodiversity effects become more critical with a higher level of sedimentation stress, and show that effective ecosystem management for resilience requires different action under different scenarios.

Linking Traits across Ecological Scales Determines Functional Resilience.

Under globally accelerating rates of ecosystem degradation, maintaining ecosystem function is a priority to avoid loss of valuable ecosystem services. Two factors are important: changes to the disturbance regime (stresses imposed) and resilience of biodiversity and ecosystem functions (the ecosystem's capacity to respond to change). Various attributes at different scales of ecological organisation confer resilience (from individual species to communities at landscape scales), but it is critical to understand how these attributes interact to inform how ecosystem function changes with disturbances that vary in intensity, spatial extent, and frequency. Individual species attributes influence their resistance, while attributes at the landscape-scale influence recovery of communities and function. Understanding resilience to disturbances requires defining the characteristics of a resilient community at multiple scales.

Calcium carbonate alters the functional response of coastal sediments to eutrophication-induced acidification.

Coastal ocean acidification research is dominated by laboratory-based studies that cannot necessarily predict real-world ecosystem response given its complexity. We enriched coastal sediments with increasing quantities of organic matter in the field to identify the effects of eutrophication-induced acidification on benthic structure and function, and assess whether biogenic calcium carbonate (CaCO3) would alter the response. Along the eutrophication gradient we observed declines in macrofauna biodiversity and impaired benthic net primary productivity and sediment nutrient cycling. CaCO3 addition did not alter the macrofauna community response, but significantly dampened negative effects on function (e.g. net autotrophy occurred at higher levels of organic matter enrichment in +CaCO3 treatments than -CaCO3 (1400 vs 950 g dw m-2)). By identifying the links between eutrophication, sediment biogeochemistry and benthic ecosystem structure and function in situ, our study represents a crucial step forward in understanding the ecological effects of coastal acidification and the role of biogenic CaCO3 in moderating responses.

Effects of CO2 enrichment on benthic primary production and inorganic nitrogen fluxes in two coastal sediments.

Ocean acidification may alter the cycling of nitrogen in coastal sediment and so the sediment-seawater nitrogen flux, an important driver of pelagic productivity. To investigate how this perturbation affects the fluxes of NOX- (nitrite/nitrate), NH4+ and O2, we incubated estuarine sand and subtidal silt in recirculating seawater with a CO2-adjusted pH of 8.1 and 7.9. During a 41-day incubation, the seawater kept at pH 8.1 lost 97% of its NOX- content but the seawater kept at pH 7.9 lost only 18%. Excess CO2 increased benthic photosynthesis. In the silt, this was accompanied by a reversal of the initial NOX- efflux into influx. The estuarine sand sustained its initial NOX- influx but, by the end of the incubation, released more NH4+ at pH 7.9 than at pH 8.1. We hypothesise that these effects share a common cause; excess CO2 increased the growth of benthic microalgae and so nutrient competition with ammonia oxidising bacteria (AOB). In the silt, diatoms likely outcompeted AOB for NH4+ and photosynthesis increased the dark/light fluctuations in the pore water oxygenation inhibiting nitrification and coupled nitrification/denitrification. If this is correct, then excess CO2 may lead to retention of inorganic nitrogen adding to the pressures of increasing coastal eutrophication.

Estimating the effect of burrowing shrimp on deep-sea sediment community oxygen consumption.

Sediment community oxygen consumption (SCOC) is a proxy for organic matter processing and thus provides a useful proxy of benthic ecosystem function. Oxygen uptake in deep-sea sediments is mainly driven by bacteria, and the direct contribution of benthic macro- and mega-infauna respiration is thought to be relatively modest. However, the main contribution of infaunal organisms to benthic respiration, particularly large burrowing organisms, is likely to be indirect and mainly driven by processes such as feeding and bioturbation that stimulate bacterial metabolism and promote the chemical oxidation of reduced solutes. Here, we estimate the direct and indirect contributions of burrowing shrimp (Eucalastacus cf. torbeni) to sediment community oxygen consumption based on incubations of sediment cores from 490 m depth on the continental slope of New Zealand. Results indicate that the presence of one shrimp in the sediment is responsible for an oxygen uptake rate of about 40 µmol d-1, only 1% of which is estimated to be due to shrimp respiration. We estimate that the presence of ten burrowing shrimp m-2 of seabed would lead to an oxygen uptake comparable to current estimates of macro-infaunal community respiration on Chatham Rise based on allometric equations, and would increase total sediment community oxygen uptake by 14% compared to sediment without shrimp. Our findings suggest that oxygen consumption mediated by burrowing shrimp may be substantial in continental slope ecosystems.

Density of Key-Species Determines Efficiency of Macroalgae Detritus Uptake by Intertidal Benthic Communities.

Accumulating evidence shows that increased biodiversity has a positive effect on ecosystem functioning, but the mechanisms that underpin this positive relationship are contentious. Complete extinctions of regional species pools are comparatively rare whereas compositional changes and reductions in abundance and biomass are common, although seldom the focus of biodiversity-ecosystem functioning studies. We use natural, small-scale patchiness in the density of two species of large bivalves with contrasting feeding modes (the suspension-feeding Austrovenus stutchburyi and deposit-feeding Macomona liliana) to examine their influence on the uptake of nitrogen from macroalgae detritus (i.e. measure of ecosystem function and food web efficiency) by other infauna in a 10-d laboratory isotope-tracer experiment. We predicted that densities of these key bivalve species and functional group diversity (calculated as Shannons H, a density-independent measure of community composition) of the intact infaunal community will be critical factors explaining variance in macroalgal per capita uptake rates by the community members and hence determine total uptake by the community. Results show that only two species, M. liliana and a large orbiniid polychaete (Scoloplos cylindrifer) dominated macroalgal nitrogen taken up by the whole community due to their large biomass. However, their densities were mostly not important or negatively influenced per capita uptake by other species. Instead, the density of a head-down deposit-feeder (the capitellid Heteromastus filiformis), scavengers (mainly nemertines and nereids) and species and functional group diversity, best explained per capita uptake rates in community members. Our results demonstrate the importance of species identity, density and large body size for ecosystem functioning and highlight the complex interactions underlying loss of ecological functions with declining biodiversity and compositional changes.