Ecosystem Processes Performed by Unionid Mussels in Stream Mesocosms: Species Roles and Effects of Abundance

Unionid mussels are a guild of freshwater, sedentary filter-feeders experiencing a global decline in both species richness and abundance. To predict how these losses may impact stream ecosystems we need to quantify the effects of both overall mussel biomass and individual species on ecosystem processes. In this study we begin addressing these fundamental questions by comparing rates of ecosystem processes for two common mussel species, Amblema plicata and Actinonaias ligamentina, across a range of abundance levels and at two trophic states (low and high productivity) in stream mesocosms. At both low and high productivity, community respiration, water column ammonia, nitrate, and phosphorus concentrations, and algal clearance rates were all linearly related to overall mussel biomass. After removing the effects of biomass with ANCOVA, we found few differences between species. In a separate series of experiments, nutrient excretion (phosphorus, ammonia, and molar N:P) and biodeposition rates were only marginally different between species. For the species studied here, functional effects of unionids in streams were similar between species and linearly related to biomass, indicating the potential for strong effects when overall mussel biomass is high and hydrologic residence times are long.     

Environmental context determines multi‐trophic effects of consumer species loss

Loss of biodiversity and nutrient enrichment are two of the main human impacts on ecosystems globally, yet we understand very little about the interactive effects of multiple stressors on natural communities and how this relates to biodiversity and ecosystem functioning. Advancing our understanding requires the following: (1) incorporation of processes occurring within and among trophic levels in natural ecosystems and (2) tests of context‐dependency of species loss effects. We examined the effects of loss of a key predator and two groups of its prey on algal assemblages at both ambient and enriched nutrient conditions in a marine benthic system and tested for interactions between the loss of functional diversity and nutrient enrichment on ecosystem functioning. We found that enrichment interacted with food web structure to alter the effects of species loss in natural communities. At ambient conditions, the loss of primary consumers led to an increase in biomass of algae, whereas predator loss caused a reduction in algal biomass (i.e. a trophic cascade). However, contrary to expectations, we found that nutrient enrichment negated the cascading effect of predators on algae. Moreover, algal assemblage structure varied in distinct ways in response to mussel loss, grazer loss, predator loss and with nutrient enrichment, with compensatory shifts in algal abundance driven by variation in responses of different algal species to different environmental conditions and the presence of different consumers. We identified and characterized several context‐dependent mechanisms driving direct and indirect effects of consumers. Our findings highlight the need to consider environmental context when examining potential species redundancies in particular with regard to changing environmental conditions. Furthermore, non‐trophic interactions based on empirical evidence must be incorporated into food web‐based ecological models to improve understanding of community responses to global change.     

Biodiversity effects on productivity and stability of marine macroalgal communities: the role of environmental context

The influence of biodiversity on ecosystem functioning has been the focus of much recent research, but the role of environmental context and the mechanisms by which it may influence diversity effects on production and stability remain poorly understood. We assembled marine macroalgal communities in two mesocosm experiments that varied nutrient supply, and at four field sites that differed naturally in environmental conditions. Concordant with theory, nutrient addition promoted positive species richness effects on algal growth in the first mesocosm experiment; however, it tended to weaken the positive diversity relationship found under ambient conditions in a second experiment the next year. In the field experiments, species richness increased algal biomass production at two of four sites. Together, these experiments indicate that diversity effects on algal biomass production are strongly influenced by environmental conditions that vary over space and time. In decomposing the net biodiversity effect into its component mechanisms, seven of the eight experimental settings showed positive complementarity effects (suggesting facilitation or complementary resource use) countered by negative selection effects (i.e. enhanced growth in mixture of otherwise slow growing species) to varying degrees. Under no conditions, including nutrient enrichment, did we find evidence of positive selection effects commonly thought to drive positive diversity effects. Species richness enhanced stability of algal community biomass across a range of environmental settings in our field experiments. Hence, while species richness can increase production, enhanced stability is also an important functional outcome of maintaining diverse marine macroalgal communities.     

A cross‐system meta‐analysis reveals coupled predation effects on prey biomass and diversity

Predator diversity and abundance are under strong human pressure in all types of ecosystems. Whereas predator potentially control standing biomass and species interactions in food webs, their effects on prey biomass and especially prey biodiversity have not yet been systematically quantified. Here, we test the effects of predation in a cross‐system meta‐analysis of prey diversity and biomass responses to local manipulation of predator presence. We found 291 predator removal experiments from 87 studies assessing both diversity and biomass responses. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey across ecosystems. Predation effects were highly similar between ecosystem types, whereas previous studies had shown that herbivory or decomposition effects differed fundamentally between terrestrial and aquatic systems based on different stoichiometry of plant material. Such stoichiometric differences between systems are unlikely for carnivorous predators, where effect sizes on species richness strongly correlated to effect sizes on biomass. However, the negative predation effect on prey biomass was ameliorated significantly with increasing prey richness and increasing species richness of the manipulated predator assemblage. Moreover, with increasing richness of the predator assemblage present, the overall negative effects of predation on prey richness switched to positive effects. Our meta‐analysis revealed strong general relationships between predator diversity, prey diversity and the interaction strength between trophic levels in terms of biomass. This study indicates that anthropogenic changes in predator abundance and diversity will potentially have strong effects on trophic interactions across ecosystems. Synthesis The past centuries we have experienced a dramatic loss of top–predator abundance and diversity in most types of ecosystems. To understand the direct consequences of predator loss on a global scale, we quantitatively summarized experiments testing predation effects on prey communities in a cross‐system meta‐analysis. Across ecosystem types, predator presence significantly decreased both biomass and diversity of prey, and predation effects were highly similar. However, with increasing predator richness, the overall negative effects of predation on prey richness switched to positive ones. Anthropogenic changes in predator communities will potentially have strong effects on prey diversity, biomass, and trophic interactions across ecosystems.     

Predator diversity effects cascade across an ecosystem boundary

Food webs in different ecosystems are often connected through spatial resource subsidies. As a result, biodiversity effects in one ecosystem may cascade to adjacent ecosystems. I tested the hypothesis that aquatic predator diversity effects cascade to terrestrial food webs by altering a prey subsidy (biomass and trophic structure of emerging aquatic insects) entering terrestrial food webs, in turn altering the distribution of a terrestrial consumer (spider) that feeds on emerging aquatic insects. Fish presence, but not diversity, altered the trophic structure of emerging aquatic insects by strongly reducing the biomass of emerging predators (dragonflies) relative to non‐feeding taxa (chironomid midges). Fish diversity reduced emerging insect biomass through enhanced effects on the most common prey taxa: predatory dragonflies Pantala flavescens and non‐feeding chironomids. Terrestrial spiders (Tetragnathidae) primarily captured emerging chironomids, which were reduced in the high richness (3 spp.) treatment relative to the 1 and 2 species treatments. As a result, terrestrial spider abundance was lower above pools with high fish richness (3 species) than pools with 1 and 2 species. Synergistic predation effects were mostly limited to the high richness treatment, in which fish occupied each level of vertical microhabitat in the water‐column (benthic, middle, surface). This study demonstrates that predator diversity effects are not limited to the habitat of the predator, but can propagate to adjacent ecosystems, and demonstrates the utility of using simple predator functional traits (foraging domain) to more accurately predict the direction of predator diversity effects.     

Effects of algal diversity on the production of biomass in homogeneous and heterogeneous nutrient environments: a microcosm experiment

Background

One of the most common questions addressed by ecologists over the past decade has been-how does species richness impact the production of community biomass? Recent summaries of experiments have shown that species richness tends to enhance the production of biomass across a wide range of trophic groups and ecosystems; however, the biomass of diverse polycultures only rarely exceeds that of the single most productive species in a community (a phenomenon called ‘transgressive overyielding’). Some have hypothesized that the lack of transgressive overyielding is because experiments have generally been performed in overly-simplified, homogeneous environments where species have little opportunity to express the niche differences that lead to ‘complementary’ use of resources that can enhance biomass production. We tested this hypothesis in a laboratory experiment where we manipulated the richness of freshwater algae in homogeneous and heterogeneous nutrient environments.

Methodology/Principal Findings

Experimental units were comprised of patches containing either homogeneous nutrient ratios (16∶1 nitrogen to phosphorus (N∶P) in all patches) or heterogeneous nutrient ratios (ranging from 4∶1 to 64∶1 N∶P across patches). After allowing 6–10 generations of algal growth, we found that algal species richness had similar impacts on biomass production in both homo- and heterogeneous environments. Although four of the five algal species showed a strong response to nutrient heterogeneity, a single species dominated algal communities in both types of environments. As a result, a ‘selection effect’–where diversity maximizes the chance that a competitively superior species will be included in, and dominate the biomass of a community–was the primary mechanism by which richness influenced biomass in both homo- and heterogeneous environments.

Conclusions/Significance

Our study suggests that spatial heterogeneity, by itself, is not sufficient to generate strong effects of biodiversity on productivity. Rather, heterogeneity must be coupled with variation in the relative fitness of species across patches in order for spatial niche differentiation to generate complementary resource use.     

Ecosystem Functions across Trophic Levels Are Linked to Functional and Phylogenetic Diversity

In experimental systems, it has been shown that biodiversity indices based on traits or phylogeny can outperform species richness as predictors of plant ecosystem function. However, it is unclear whether this pattern extends to the function of food webs in natural ecosystems. Here we tested whether zooplankton functional and phylogenetic diversity explains the functioning of 23 natural pond communities. We used two measures of ecosystem function: (1) zooplankton community biomass and (2) phytoplankton abundance (Chl a). We tested for diversity-ecosystem function relationships within and across trophic levels. We found a strong correlation between zooplankton diversity and ecosystem function, whereas local environmental conditions were less important. Further, the positive diversity-ecosystem function relationships were more pronounced for measures of functional and phylogenetic diversity than for species richness. Zooplankton and phytoplankton biomass were best predicted by different indices, suggesting that the two functions are dependent upon different aspects of diversity. Zooplankton community biomass was best predicted by zooplankton trait-based functional richness, while phytoplankton abundance was best predicted by zooplankton phylogenetic diversity. Our results suggest that the positive relationship between diversity and ecosystem function can extend across trophic levels in natural environments, and that greater insight into variation in ecosystem function can be gained by combining functional and phylogenetic diversity measures.     

Patterns in Benthic Biodiversity Link Lake Trophic Status to Structure and Potential Function of Three Large,Deep Lakes

Relative to their scarcity, large, deep lakes support a large proportion of the world’s freshwater species. This biodiversity is threatened by human development and is in need of conservation. Direct comparison of biodiversity is the basis of biological monitoring for conservation but is difficult to conduct between large, insular ecosystems. The objective of our study was to conduct such a comparison of benthic biodiversity between three of the world’s largest lakes: Lake Tahoe, USA; Lake Hövsgöl, Mongolia; and Crater Lake, USA. We examined biodiversity of common benthic organism, the non-biting midges (Chironomidae) and determined lake trophic status using chironomid-based lake typology, tested whether community structure was similar between the three lakes despite geographic distance; and tested whether chironomid diversity would show significant variation within and between lakes. Typology analysis indicated that Lake Hövsgöl was ultra-oligotrophic, Crater Lake was oligotrophic, and Lake Tahoe was borderline oligotrophic/mesotrophic. These results were similar to traditional pelagic measures of lake trophic status for Lake Hövsgöl and Crater Lake but differed for Lake Tahoe, which has been designated as ultra-oligotrophic by traditional pelagic measures such as transparency found in the literature. Analysis of similarity showed that Lake Tahoe and Lake Hövsgöl chironomid communities were more similar to each other than either was to Crater Lake communities. Diversity varied between the three lakes and spatially within each lake. This research shows that chironomid communities from these large lakes were sensitive to trophic conditions. Chironomid communities were similar between the deep environments of Lake Hövsgöl and Lake Tahoe, indicating that chironomid communities from these lakes may be useful in comparing trophic state changes in large lakes. Spatial variation in Lake Tahoe’s diversity is indicative of differential response of chironomid communities to nutrient enrichment which may be an indication of changes in trophic state within and across habitats.     

Diversity and structure of macrozoobenthic communities in the highly mineralized Khara River (territory adjacent to Lake Elton)

The present paper deals with the results of the first study (April and August, 2006–2007) of the taxonomic composition of macrozoobenthos in the highly mineralized Khara River. The study revealed 35 species and taxa of the rank above species. Data were obtained on the spatial distribution of the species composition dynamics and on the quantitative development of the macrozoobenthic community. The dependence of the structure of bottom communities on the parameters of the river’s trophic status, biotopic diversity, and changes in hydrochemical parameters was determined. The environmental factors (total mineralization, ionic composition, pH, oxygen content, water temperature, overgrowth by macrophytes) influencing structural changes in macrozoobenthic community were determined. It is shown that diversity, number, and biomass of halotolerant macrozoobenthic communities increase with a rise in water mineralization from 6.9 to 14 mg/L. A new halophilic chironomid species of the family Chironominae, Tanytarsus kharaensis Zorina et Zinchenko, has been found.     

Predator Assemblage Structure and Temporal Variability of Species Richness and Abundance in Forests of High Tree Diversity

Predators significantly affect ecosystem functions, but our understanding of to what extent findings can be transferred from experiments and low‐diversity systems to highly diverse, natural ecosystems is limited. With a particular threat of biodiversity loss at higher trophic levels, however, knowledge of spatial and temporal patterns in predator assemblages and their interrelations with lower trophic levels is essential for assessing effects of trophic interactions and advancing biodiversity conservation in these ecosystems. We analyzed spatial and temporal variability of spider assemblages in tree species‐rich subtropical forests in China, across 27 study plots varying in woody plant diversity and stand age. Despite effects of woody plant richness on spider assemblage structure, neither habitat specificity nor temporal variability of spider richness and abundance were influenced. Rather, variability increased with forest age, probably related to successional changes in spider assemblages. Our results indicate that woody plant richness and theory predicting increasing predator diversity with increasing plant diversity do not necessarily play a major role for spatial and temporal dynamics of predator assemblages in such plant species‐rich forests. Diversity effects on biotic or abiotic habitat conditions might be less pronounced across our gradient from medium to high plant diversity than in previously studied less diverse systems, and bottom‐up effects might level out at high plant diversity. Instead, our study highlights the importance of overall (diversity‐independent) environmental heterogeneity in shaping spider assemblages and, as indicated by a high species turnover between plots, as a crucial factor for biodiversity conservation at a regional scale in these subtropical forests.     

The influence of a dissolved inorganic nitrogen gradient on phytoplankton community dynamics in a chain of lakes

In a chain of lakes along which nutrient availability varies in a gradient, we performed factorial nutrient enrichment experiments to determine if nitrogen limitation was the principal factor controlling the differences in phytoplankton biomass, photosynthetic productivity, diversity, and species composition among two of the lakes in the chain. In the least productive lake, East Graham Lake, P and C enrichments (in the absence of N enrichment) had no effect on biomass and diversity, whereas within two weeks the N enrichments (alone or in any combination with P and/or C) increased the biomass and decreased the diversity of East Graham Lake phytoplankton to levels similar or identical to those in more productive Shoe Lake. Short-term ¹⁴C photosynthetic rates in East Graham Lake water also responded only to N in the third week. However, photosynthesis was stimulated by P in the first week, and a few species did increase in numbers with P enrichment, suggesting that some degree of P limitation remains in addition to the strong N limitation in East Graham Lake. A number of species responded individually to the enrichments in a manner similar to that of the overall community, and a strong overlapping of discriminant analysis scores for N-enriched East Graham Lake with those of Shoe Lake was consistent with our prediction that the community structure of N-enriched East Graham Lake water would shift toward that of Shoe Lake. However, many species did not respond consistently with these results, and the nutrients tested were clearly not a major factor in the differences in abundance of those species among the two lakes. The results support the argument that overall biomass production and diversity of the phytoplankton community in a lake can be a relatively simple function of a single most-limiting nutrient. However, many of the species responses also confirm that, while nutrient availability is an important factor in the control of the species composition of the community, other factors are likely to prevent reliable predictions of all species effects on the basis of nutrient availability alone.     

Plant Diversity Impacts Decomposition and Herbivory via Changes in Aboveground Arthropods

Loss of plant diversity influences essential ecosystem processes as aboveground productivity, and can have cascading effects on the arthropod communities in adjacent trophic levels. However, few studies have examined how those changes in arthropod communities can have additional impacts on ecosystem processes caused by them (e.g. pollination, bioturbation, predation, decomposition, herbivory). Therefore, including arthropod effects in predictions of the impact of plant diversity loss on such ecosystem processes is an important but little studied piece of information. In a grassland biodiversity experiment, we addressed this gap by assessing aboveground decomposer and herbivore communities and linking their abundance and diversity to rates of decomposition and herbivory. Path analyses showed that increasing plant diversity led to higher abundance and diversity of decomposing arthropods through higher plant biomass. Higher species richness of decomposers, in turn, enhanced decomposition. Similarly, species-rich plant communities hosted a higher abundance and diversity of herbivores through elevated plant biomass and C:N ratio, leading to higher herbivory rates. Integrating trophic interactions into the study of biodiversity effects is required to understand the multiple pathways by which biodiversity affects ecosystem functioning.     

湖北四湖泊营养类型与轮虫群落的关系

对湖北梁子湖水系不同营养类型(中营养型、富营养型)4个湖泊中轮虫的群落结构和物种多样性进行了周年研究,分析比较了不同营养类型湖泊的轮虫种类组成、分布、优势种组成、密度、生物量和多样性指数。结果表明：轮虫的种类数、物种多样性与营养水平呈负相关关系,轮虫密度大体上随营养水平提高而增大,富营养化引起轮虫空间异质性降低,受污染湖泊与非污染湖泊轮虫种类数、寡污性种类数及分布差异尤为明显。用多样性指数评价湖泊营养状态与TLIc方法一致。     

Changes in Nematode Communities in Different Physiographic Sites of the Condor Seamount (North-East Atlantic Ocean) and Adjacent Sediments

Several seamounts are known as ‘oases’ of high abundances and biomass and hotspots of biodiversity in contrast to the surrounding deep-sea environments. Recent studies have indicated that each single seamount can exhibit a high intricate habitat turnover. Information on alpha and beta diversity of single seamount is needed in order to fully understand seamounts contribution to regional and global biodiversity. However, while most of the seamount research has been focused on summits, studies considering the whole seamount structure are still rather poor. In the present study we analysed abundance, biomass and diversity of nematodes collected in distinct physiographic sites and surrounding sediments of the Condor Seamount (Azores, North-East Atlantic Ocean). Our study revealed higher nematode biomass in the seamount bases and values 10 times higher in the Condor sediments than in the far-field site. Although biodiversity indices did not showed significant differences comparing seamount sites and far-field sites, significant differences were observed in term of nematode composition. The Condor summit harboured a completely different nematode community when compared to the other seamount sites, with a high number of exclusive species and important differences in term of nematode trophic diversity. The oceanographic conditions observed around the Condor Seamount and the associated sediment mixing, together with the high quality of food resources available in seamount base could explain the observed patterns. Our results support the hypothesis that seamounts maintain high biodiversity through heightened beta diversity and showed that not only summits but also seamount bases can support rich benthic community in terms of standing stocks and diversity. Furthermore functional diversity of nematodes strongly depends on environmental conditions link to the local setting and seamount structure. This finding should be considered in future studies on seamounts, especially in view of the potential impacts due to current and future anthropogenic threats.     

Contrasting effects of functionally distinct tadpole species on nutrient cycling and litter breakdown in a tropical rainforest stream

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Experimental nitrogen and phosphorus enrichment stimulates multiple trophic levels of algal and detrital-based food webs: a global meta-analysis from streams and rivers

Anthropogenic increases in nitrogen (N) and phosphorus (P) concentrations can strongly influence the structure and function of ecosystems. Even though lotic ecosystems receive cumulative inputs of nutrients applied to and deposited on land, no comprehensive assessment has quantified nutrient-enrichment effects within streams and rivers. We conducted a meta-analysis of published studies that experimentally increased concentrations of N and/or P in streams and rivers to examine how enrichment alters ecosystem structure (state: primary producer and consumer biomass and abundance) and function (rate: primary production, leaf breakdown rates, metabolism) at multiple trophic levels (primary producer, microbial heterotroph, primary and secondary consumers, and integrated ecosystem). Our synthesis included 184 studies, 885 experiments, and 3497 biotic responses to nutrient enrichment. We documented widespread increases in organismal biomass and abundance (mean response = +48%) and rates of ecosystem processes (+54%) to enrichment across multiple trophic levels, with no large differences in responses among trophic levels or between autotrophic or heterotrophic food-web pathways. Responses to nutrient enrichment varied with the nutrient added (N, P, or both) depending on rate versus state variable and experiment type, and were greater in flume and whole-stream experiments than in experiments using nutrient-diffusing substrata. Generally, nutrient-enrichment effects also increased with water temperature and light, and decreased under elevated ambient concentrations of inorganic N and/or P. Overall, increased concentrations of N and/or P altered multiple food-web pathways and trophic levels in lotic ecosystems. Our results indicate that preservation or restoration of biodiversity and ecosystem functions of streams and rivers requires management of nutrient inputs and consideration of multiple trophic pathways.     

Determining the mechanism by which fish diversity influences production

Understanding the ability of biodiversity to govern ecosystem function is essential with current pressures on natural communities from species invasions and extirpations. Changes in fish communities can be a major determinant of food web dynamics, and even small shifts in species composition or richness can translate into large effects on ecosystems. In addition, there is a large information gap in extrapolating results of small-scale biodiversity–ecosystem function experiments to natural systems with realistic environmental complexity. Thus, we tested the key mechanisms (resource complementarity and selection effect) for biodiversity to influence fish production in mesocosms and ponds. Fish diversity treatments were created by replicating species richness and species composition within each richness level. In mesocosms, increasing richness had a positive effect on fish biomass with an overyielding pattern indicating species mixtures were more productive than any individual species. Additive partitioning confirmed a positive net effect of biodiversity driven by a complementarity effect. Productivity was less affected by species diversity when species were more similar. Thus, the primary mechanism driving fish production in the mesocosms was resource complementarity. In the ponds, the mechanism driving fish production changed through time. The key mechanism was initially resource complementarity until production was influenced by the selection effect. Varying strength of intraspecific interactions resulting from differences in resource levels and heterogeneity likely caused differences in mechanisms between the mesocosm and pond experiments, as well as changes through time in the ponds. Understanding the mechanisms by which fish diversity governs ecosystem function and how environmental complexity and resource levels alter these relationships can be used to improve predictions for natural systems.     

Habitat engineering by the invasive zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis> (Pallas) in a boreal coastal lagoon: impact on biodiversity

Habitat engineering role of the invasive zebra mussel Dreissena polymorpha (Pallas) was studied in the Curonian lagoon, a shallow water body in the SE Baltic. Impacts of live zebra mussel clumps and its shell deposits on benthic biodiversity were differentiated and referred to unmodified (bare) sediments. Zebra mussel bed was distinguished from other habitat types by higher benthic invertebrate biomass, abundance, and species richness. The impact of live mussels on biodiversity was more pronounced than the effect of shell deposits. The structure of macrofaunal community in the habitats with >10³ g/m² of shell deposits devoid of live mussels was similar to that found within the zebra mussel bed. There was a continuous shift in species composition and abundance along the gradient ‘bare sediments—shell deposits—zebra mussel bed’. The engineering impact of zebra mussel on the benthic community became apparent both in individual patches and landscape-level analyses.     

Dynamics of a Boreal Lake Ecosystem during a Long-Term Manipulation of Top Predators

We assessed the long-term (16 years) effects of introducing piscivores (northern pike) into a small, boreal lake (Lake 221, Experimental Lakes Area) containing abundant populations of two planktivorous fish species. After the introduction, pearl dace were extirpated and yellow perch abundance was greatly reduced. Daphnia species shifted from D. galeata mendota to larger bodied Daphnia catawba, but the total zooplankton biomass did not increase, nor did the biomass of large grazers such as Daphnia. Phytoplankton biomass decreased after the northern pike introduction, but increased when northern pike were partially removed from the lake. Phosphorus (P) excretion by fish was ∼0.18 mg P m⁻² d⁻¹ before pike addition, declined rapidly to approximately 0.03–0.10 as planktivorous perch and dace populations were reduced by pike, and increased back to premanipulation levels after the pike were partially removed and the perch population recovered. When perch were abundant, P excretion by fish supported about 30% of the P demand by primary producers, decreasing to 6–14% when pike were abundant. Changes in phytoplankton abundance in Lake 221 appear to be driven by changes in P cycling by yellow perch, whose abundance was controlled by the addition and removal of pike. These results confirm the role of nutrient cycling in mediating trophic cascades and are consistent with previous enclosure experiments conducted in the same lake.     

A general biodiversity–function relationship is mediated by trophic level

Species diversity affects the functioning of ecosystems, including the efficiency by which communities capture limited resources, produce biomass, recycle and retain biologically essential nutrients. These ecological functions ultimately support the ecosystem services upon which humanity depends. Despite hundreds of experimental tests of the effect of biodiversity on ecosystem function (BEF), it remains unclear whether diversity effects are sufficiently general that we can use a single relationship to quantitatively predict how changes in species richness alter an ecosystem function across trophic levels, ecosystems and ecological conditions. Our objective here is to determine whether a general relationship exists between biodiversity and standing biomass. We used hierarchical mixed effects models, based on a power function between species richness and biomass production (Y = a × S^b), and a database of 374 published experiments to estimate the BEF relationship (the change in biomass with the addition of species), and its associated uncertainty, in the context of environmental factors. We found that the mean relationship (b = 0.26, 95% CI: 0.16, 0.37) characterized the vast majority of observations, was robust to differences in experimental design, and was independent of the range of species richness levels considered. However, the richness–biomass relationship varied by trophic level and among ecosystems; in aquatic systems b was nearly twice as large for consumers (herbivores and detritivores) compared to primary producers; in terrestrial ecosystems, b for detritivores was negative but depended on few studies. We estimated changes in biomass expected for a range of changes in species richness, highlighting that species loss has greater implications than species gains, skewing a distribution of biomass change relative to observed species richness change. When biomass provides a good proxy for processes that underpin ecosystem services, this relationship could be used as a step in modeling the production of ecosystem services and their dependence on biodiversity.