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1: Body size is a decisive functional trait in many organisms, especially for phytoplankton, which span several orders of magnitude in cell volume. Therefore, the analysis of size as a functional trait driving species’ performance has received wide attention in aquatic ecology, amended in recent decades by studies documenting changes in phytoplankton size in response to abiotic or biotic factors in the environment. 2. We performed a systematic literature review to provide an overarching, partially quantitative synthesis of cell size as a driver and sentinel of phytoplankton ecology. We found consistent and significant allometric relationships between cell sizes and the functional performance of phytoplankton species (cellular rates of carbon fixation, respiration, and exudation as well as resource affinities, uptake, and content). Size-scaling became weaker, absent or even negative when addressing C- or volume-specific rates or growth. C-specific photosynthesis and population growth rate peaked at intermediate cell sizes around 100 µm³. 4: Additionally, we found a rich literature on sizes changing in response to warming, nutrients and pollutants. Whereas small cells tended to dominate under oligotrophic and warm conditions, there are a few notable exceptions, which indicates that other environmental or biotic constraints alter this general trend. Grazing seems a likely explanation, which we reviewed to understand both how size affects edibility and how size structure changes in response to grazing. Cell size also predisposes the strength and outcome of competitive interactions between algal species. Finally, we address size in a community context, where size-abundance scaling describes community composition and thereby the biodiversity in phytoplankton assemblages. 5: We conclude that i) size is a highly predictive trait for phytoplankton metabolism at the cellular scale, with less strong and non-linear implications for growth and specific metabolism, and ii) size structure is a highly suitable sentinel of phytoplankton responses to changing environments.

Cell size is a master trait in the functional ecology of phytoplankton correlating with numerous morphological, physiological, and life‐cycle characteristics of species that constrain their nutrient use, growth, and edibility. In contrast to well‐known spatial patterns in cell size at macroecological scales or temporal changes in experimental contexts, few data sets allow testing temporal changes in cell sizes within ecosystems. To analyze the temporal changes of intraspecific and community‐wide cell size, we use the phytoplankton data derived from the Lower Saxony Wadden Sea monitoring program, which comprises sample‐ and species‐specific measurements of cell volume from 1710 samples collected over 14 yr. We find significant reductions in both the cell volume of most species and the weighted mean cell size of communities. Mainly diatoms showed this decline, whereas the size of dinoflagellates seemed to be less responsive. The magnitude of the trend indicates that cell volumes are about 30% smaller now than a decade ago. This interannual trend is overlayed by seasonal cycles with smaller cells typically observed in summer. In the subset of samples including environmental conditions, small community cell size was strongly related to high temperatures and low total phosphorus concentration. We conclude that cell size captures ongoing changes in phytoplankton communities beyond the changes in species composition. In addition, based on the changes in species biovolumes revealed by our analysis, we warn that using standard cell size values in phytoplankton assessment will not only miss temporal changes in size, but also lead to systematic errors in biomass estimates over time., Bundesministerium für Bildung und Forschung http://dx.doi.org/10.13039/501100002347, Deutsche Forschungsgemeinschaft http://dx.doi.org/10.13039/501100001659, Interreg V A program Deutschland‐Nederland of the European Union, Niedersächsisches Ministerium für Wissenschaft und Kultur http://dx.doi.org/10.13039/501100010570, https://doi.org/10.5281/zenodo.5799263

While environmental science, and ecology in particular, is working to provide better understanding to base sustainable decisions on, the way scientific understanding is developed can at times be detrimental to this cause. Locked-in debates are often unnecessarily polarized and can compromise any common goals of the opposing camps. The present paper is inspired by a resolved debate from an unrelated field of psychology where Nobel laureate David Kahneman and Garry Klein turned what seemed to be a locked-in debate into a constructive process for their fields. The present paper is also motivated by previous discourses regarding the role of thresholds in natural systems for management and governance, but its scope of analysis targets the scientific process within complex social-ecological systems in general. We identified five features of environmental science that appear to predispose for locked-in debates: 1) The strongly context dependent behaviour of ecological systems. 2) The dominant role of single hypothesis testing. 3) The high prominence given to theory demonstration compared investigation. 4) The effect of urgent demands to inform and steer policy. This fertile ground is further cultivated by human psychological aspects as well as the structure of funding and publication systems.

Increasing human impact on the environment is causing drastic changes in disturbance regimes and how they prevail over time. Of increasing relevance is to further our understanding on biological responses to pulse disturbances (short duration) and how they interact with other ongoing press disturbances (constantly present). Because the temporal and spatial contexts of single experiments often limit our ability to generalize results across space and time, we conducted a modularized mesocosm experiment replicated in space (five lakes along a latitudinal gradient in Scandinavia) and time (two seasons, spring and summer) to generate general predictions on how the functioning and composition of multitrophic plankton communities (zoo-, phyto- and bacterioplankton) respond to pulse disturbances acting either in isolation or combined with press disturbances. As pulse disturbance, we used short-term changes in fish presence, and as press disturbance, we addressed the ongoing reduction in light availability caused by increased cloudiness and lake browning in many boreal and subarctic lakes. First, our results show that the top-down pulse disturbance had the strongest effects on both functioning and composition of the three trophic levels across sites and seasons, with signs for interactive impacts with the bottom-up press disturbance on phytoplankton communities. Second, community composition responses to disturbances were highly divergent between lakes and seasons: temporal accumulated community turnover of the same trophic level either increased (destabilization) or decreased (stabilization) in response to the disturbances compared to control conditions. Third, we found functional recovery from the pulse disturbances to be frequent at the end of most experiments. In a broader context, these results demonstrate that top-down, pulse disturbances, either alone or with additional constant stress upon primary producers caused by bottom-up disturbances, can induce profound but often functionally reversible changes across multiple trophic levels, which are strongly linked to spatial and temporal context dependencies. Furthermore, the identified dichotomy of disturbance effects on the turnover in community composition demonstrates the potential of disturbances to either stabilize or destabilize biodiversity patterns over time across a wide range of environmental conditions.

For aquatic scientists mesocosm experiments are important tools for hypothesis testing as they offer a compromise between experimental control and realism. Here we present a new mesocosm infrastructure-SITES AquaNET-located in five lakes connected to field stations in Sweden that cover a similar to 760 km latitudinal gradient. SITES AquaNet overcomes major hindrances in aquatic experimental research through: (i) openness to the scientific community, (ii) the potential to implement coordinated experiments across sites and time, and (iii) high-frequency measurements (temperature, photosynthetic photon flux density, turbidity and dissolved oxygen, chlorophyll a and phycocyanin concentrations) with an autonomous sensor system. Moreover, the infrastructure provides operational guidance and sensor expertise from technical staff, and connections to a multi-layered monitoring programme ("SITES Water") for each lake. This enables ecological observations from whole lake ecosystems to be compared with experimental studies aiming at disentangling major drivers and mechanisms underlying observed changes. Here we describe the technical properties of the infrastructure along with possibilities for experimental manipulations to tackle pressing issues in aquatic ecology and global change science. As a proof of concept, we also present a first mesocosm experiment across all five field sites with a cross-factorial design to evaluate responses of the sensor measurements to press/bottom-up (constant light reduction) and pulse/top-down (temporary fish predation) disturbances. This demonstrates the suitability of the infrastructure and autonomous sensor system to host modularized experiments and exemplifies the power and advantages of the approach to integrate a network of mecsocosm facilities with manageable costs across large geographic areas.

Ecological stability encompasses multiple dimensions of functional and compositional responses to environmental change. Though no single stability dimension used in isolation can fully reflect the overall response to environmental change, a common vulnerability assessment that integrates simultaneously across multiple stability components is highly desirable for ecological risk assessment. We develop both functional and compositional counterparts of a novel, integrative metric of overall ecological vulnerability (OEV). We test the framework with data from a modularized experiment replicated in five lakes over two seasons, examining functional and compositional responses to both pulse and press disturbances across three trophic groups. OEV is measured as the area under the curve integrated over the entire observation period, with the curve delimiting the difference between the disturbance treatment and undisturbed parallel controls, expressed either as the log response ratio of biomass (functional OEV) or community dissimilarity index (compositional OEV). Both, functional and compositional OEV correlated negatively with functional and compositional ‘resistance’, ‘temporal stability’ and ‘final/extent of recovery’ following both pulse and press disturbances, though less so with ‘resilience’ following a pulse disturbance. We also found a positive correlation between functional and compositional OEV, which reveals the potential to also evaluate the intricate linkage between biodiversity and functional change. Our findings demonstrate that OEV comprises a robust framework to: (a) capture simultaneously multiple functional and compositional stability components, and (b) quantify the functional consequences of biodiversity change. Our results provide the basis for an overarching framework for quantifying the overall vulnerability of ecosystems to environmental change, opening new possibilities for ecological risk assessment and management. Synthesis. Ecological stability comprises multiple dimensions that together encapsulate how ecosystems respond to environmental change. Considering these multiple aspects of stability simultaneously often poses a problem in environmental assessments, which frequently require overarching indicators of risk or vulnerability. While an analysis of multiple dimensions allows for deeper exploration of mechanisms, here we develop and test a new univariate indicator that integrates stability aspects under a broad range of disturbance regimes. Using a modularized experiment in Swedish lakes, we show that this integrative measure captures multiple stability dimensions reflecting compositional and functional vulnerability and their relationships between them.

Whereas the anthropogenic impact on marine biodiversity is undebated, the quantification and prediction of this change are not trivial. Simple traditional measures of biodiversity (e.g. richness, diversity indices) do not capture the magnitude and direction of changes in species or functional composition. In this paper, we apply recently developed methods for measuring biodiversity turnover to time-series data of four broad taxonomic groups from two coastal regions: the southern North Sea (Germany) and the South African coast. Both areas share geomorphological features and ecosystem types, allowing for a critical assessment of the most informative metrics of biodiversity change across organism groups. We found little evidence for directional trends in univariate metrics of diversity for either the effective number of taxa or the amount of richness change. However, turnover in composition was high (on average nearly 30% of identities when addressing presence or absence of species) and even higher when taking the relative dominance of species into account. This turnover accumulated over time at similar rates across regions and organism groups. We conclude that biodiversity metrics responsive to turnover provide a more accurate reflection of community change relative to conventional metrics (absolute richness or relative abundance) and are spatially broadly applicable. This article is part of the theme issue ‘Integrative research perspectives on marine conservation’.

Aim Biodiversity dynamics comprise evolutionary and ecological changes on multiple temporal scales from millions of years to decades, but they are often interpreted within a single time frame. Planktonic foraminifera communities offer a unique opportunity for analysing the dynamics of marine biodiversity over different temporal scales. Our study aims to provide a baseline for assessments of biodiversity patterns over multiple time-scales, which is urgently needed to interpret biodiversity responses to increasing anthropogenic pressure. Location Global (26 sites). Time period Five time-scales: multi-million-year (0-7 Myr), million-year (0-0.5 Myr), multi-millennial (0-15 thousand years), millennial (0-1,100 years) and decadal (0-32 years). Major taxa studied Planktonic foraminifera. Methods We analysed community composition of planktonic foraminifera at five time-scales, combining measures of standing diversity (richness and effective number of species, ENS) with measures of temporal community turnover (presence-absence-based, dominance-based). Observed biodiversity patterns were compared with the outcome of a neutral model to separate the effects of sampling resolution (the highest in the shortest time series) from biological responses. Results Richness and ENS decreased from multi-million-year to millennial time-scales, but higher standing diversity was observed on the decadal scale. As predicted by the neutral model, turnover in species identity and dominance was strongest at the multi-million-year time-scale and decreased towards the millennial scale. However, contrary to the model predictions, modern time series show rapid decadal variation in the dominance structure of foraminifera communities, which is of comparable magnitude as over much longer time periods. Community turnover was significantly correlated with global temperature change, but not on the shortest time-scale. Main conclusions Biodiversity patterns can be to some degree predicted from the scaling effects related to different durations of time series, but changes in the dominance structure observed over the last few decades reach higher magnitude, probably forced by anthropogenic effects, than those observed over much longer durations.

Pronounced atmospheric and oceanic warming along the West Antarctic Peninsula (WAP) has resulted in abundance shifts in populations of Antarctic krill and Salpa thompsoni determined by changes in the timing of sea-ice advance, the duration of sea-ice cover and food availability. Krill and salps represent the most important macrozooplankton grazers at the WAP, but differ profoundly in their feeding biology, population dynamics and stoichiometry of excretion products with potential consequences for the relative availability of dissolved nitrogen and phosphorus. Alternation of the dissolved nutrient pool due to shifts in krill and salp densities have been hypothesized but never explicitly tested by using observational data. We therefore used the Palmer LTER dataset in order to investigate whether the dominance of either grazer is related with the observed dissolved nitrogen:phosphorus (N:P) ratios at the WAP. Across the whole sampling grid, the dominance of salps over krill was significantly correlated to higher concentrations of both N and P as well as a higher N:P ratios. Using actual long-term data, our study shows for the first time that changes in key grazer dominance may have consequences for the dynamics of dissolved nitrogen and phosphorus at the WAP.

1. A resampling of 38 small farmland ponds in Belgium after 10 years revealed a high temporal species turnover for both phytoplankton and zooplankton communities, associated with substantial changes in abiotic factors, especially a reduction in total phosphorus concentration. 2. Across ponds, phytoplankton biomass decreased while evenness and richness increased between the samplings in 2003 and 2013. By contrast, the zooplankton assemblage was characterised by lower biomass, richness and evenness in 2013. Ponds experiencing larger environmental change showed stronger changes in phytoplankton richness and evenness. 3. Resource use efficiency (RUE) of zooplankton increased with greater environmental change and zooplankton evenness, which points to a switch towards species with higher RUE or greater variety in food sources in higher trophic levels. 4. As ponds are important habitats for freshwater biodiversity and ecosystems services, the strong but predictable species turnover and the opposing effects of environmental change on different trophic levels need to be embedded in conservation and management plans.

Global concern about human impact on biological diversity has triggered an intense research agenda on drivers and consequences of biodiversity change in parallel with international policy seeking to conserve biodiversity and associated ecosystem functions. Quantifying the trends in biodiversity is far from trivial, however, as recently documented by meta-analyses, which report little if any net change in local species richness through time.Here, we summarise several limitations of species richness as a metric of biodiversity change and show that the expectation of directional species richness trends under changing conditions is invalid. Instead, we illustrate how a set of species turnover indices provide more information content regarding temporal trends in biodiversity, as they reflect how dominance and identity shift in communities over time.We apply these metrics to three monitoring datasets representing different ecosystem types. In all datasets, nearly complete species turnover occurred, but this was disconnected from any species richness trends. Instead, turnover was strongly influenced by changes in species presence (identities) and dominance (abundances). We further show that these metrics can detect phases of strong compositional shifts in monitoring data and thus identify a different aspect of biodiversity change decoupled from species richness.Synthesis and applications: Temporal trends in species richness are insufficient to capture key changes in biodiversity in changing environments. In fact, reductions in environmental quality can lead to transient increases in species richness if immigration or extinction has different temporal dynamics. Thus, biodiversity monitoring programmes need to go beyond analyses of trends in richness in favour of more meaningful assessments of biodiversity change.Synthesis and applications:Temporal trends in species richness are insufficient to capture key changes in biodiversity in changing environments. In fact, reductions in environmental quality can lead to transient increases in species richness if immigration or extinction has different temporal dynamics. Thus, biodiversity monitoring programmes need to go beyond analyses of trends in richness in favour of more meaningful assessments of biodiversity change.

International audience; To understand ecosystem responses to anthropogenic global change, a prevailing framework is the definition of threshold levels of pressure, above which response magnitudes and their variances increase disproportionately. However, we lack systematic quantitative evidence as to whether empirical data allow definition of such thresholds. Here, we summarize 36 meta-analyses measuring more than 4,600 global change impacts on natural communities. We find that threshold transgressions were rarely detectable, either within or across meta-analyses. Instead, ecological responses were characterized mostly by progressively increasing magnitude and variance when pressure increased. Sensitivity analyses with modelled data revealed that minor variances in the response are sufficient to preclude the detection of thresholds from data, even if they are present. The simulations reinforced our contention that global change biology needs to abandon the general expectation that system properties allow defining thresholds as a way to manage nature under global change. Rather, highly variable responses, even under weak pressures, suggest that ‘safe-operating spaces’ are unlikely to be quantifiable.

Summary 1.Niche complementarity and competitive disparity are driving mechanisms behind plant community assembly and productivity. Consequently, there is great interest in predicting species complementarity and their competitive differences from their functional traits as dissimilar species may compete less and result in more complete use of resources. 2.Here we assessed the role of trait dissimilarities on species complementarity and competitive disparities within an experimental gradient of plant species richness and functional trait dissimilarity. Communities were assembled using three pools of grass and forb species based on a priori knowledge of traits related to (1) above- and belowground spatial differences in resource acquisition, (2) phenological differences, or (3) both. Complementarity and competitive disparities were assessed by partitioning the overyielding in mixed species communities into species complementarity and dominance effects. 3.Community overyielding and the underlying complementarity and competitive dominance varied strongly among the three plant species pools. Overyielding and complementarity was greatest among species that were assembled based on their variation in both spatial and phenological traits. Competitive dominance was greatest when species were assembled based on spatial resource-acquisition traits alone. 4.In communities that were assembled based on species variation in only spatial or phenological traits greater competitive dominance was predicted by greater differences SLA and flowering initiation respectively, while greater complementarity was predicted by greater dissimilarity in leaf area and flowering senescence, respectively. Greater differences in leaf area could also be linked to greater species complementarity in communities assembled based on variation in both phenological and spatial traits, but trait dissimilarity was unrelated to competitive dominance in these communities. 5.Our results indicate that complementarity and competitive disparity among species are both driven by trait dissimilarities. However, the identity of the traits that drives the complementarity and competitive disparity depends on the trait variation among species that comprise the community. Moreover, we demonstrate that communities assembled with the greater variation in both spatial and phenological traits show the greatest complementarity among species. This article is protected by copyright. All rights reserved.

Climatic warming is a primary driver of change in ecosystems worldwide. Here, we synthesize responses of species richness and evenness from 187 experimental warming studies in a quantitative meta-analysis. We asked 1) whether effects of warming on diversity were detectable and consistent across terrestrial, freshwater and marine ecosystems, 2) if effects on diversity correlated with intensity, duration, and experimental unit size of temperature change manipulations, and 3) whether these experimental effects on diversity interacted with ecosystem types. Using multilevel mixed linear models and model averaging, we also tested the relative importance of variables that described uncontrolled environmental variation and attributes of experimental units. Overall, experimental warming reduced richness across ecosystems (mean log-response ratio = -0.091, 95% bootstrapped CI: -0.13, -0.05) representing an 8.9% decline relative to ambient temperature treatments. Richness did not change in response to warming in freshwater systems, but was more strongly negative in terrestrial (-11.8%) and marine (-10.5%) experiments. In contrast, warming impacts on evenness were neutral overall and in aquatic systems, but weakly negative on land (7.6%). Intensity and duration of experimental warming did not explain variation in diversity responses, but negative effects on richness were stronger in smaller experimental units, particularly in marine systems. Model-averaged parameter estimation confirmed these main effects while accounting for variation in latitude, ambient temperature at the sites of manipulations, venue (field versus lab), community trophic type, and whether experiments were open or closed to colonization. These analyses synthesize extensive experimental evidence showing declines in local richness with increased temperature, particularly in terrestrial and marine communities. However, the more variable effects of warming on evenness were better explained by the random effect of site identity, suggesting that effects on species' relative abundances were contingent on local species composition.

The relationship between biodiversity and ecosystem functioning (BEF) is a topic of considerable interest to scientists and managers because a better understanding of its underlying mechanisms may help us mitigate the consequences of biodiversity loss on ecosystems. Our current knowledge of BEF relies heavily on theoretical and experimental studies, typically conducted on a narrow range of spatio-temporal scales, environmental conditions, and trophic levels. Hence, whether a relationship holds in the natural environment is poorly understood, especially in exploited marine ecosystems. Using large-scale observations of marine fish communities, we applied a structural equation modelling framework to investigate the existence and significance of BEF relationships across northwestern European seas. We find that ecosystem functioning, here represented by spatial patterns in total fish biomass, is unrelated to species richness—the most commonly used diversity metric in BEF studies. Instead, community evenness, differences in species composition, and abiotic variables are significant drivers. In particular, we find that high fish biomass is associated with fish assemblages dominated by a few generalist species of a high trophic level, who are able to exploit both the benthic and pelagic energy pathway. Our study provides a better understanding of the mechanisms behind marine ecosystem functioning and allows for the integration of biodiversity into management considerations.

Reduced nitrogen species are key nutrients for biological productivity in the oceans. Ammonium is often present in low and growth-limiting concentrations, albeit peaks occur during collapse of algal blooms or via input from deep sea upwelling and riverine inflow. Autotrophic phytoplankton exploit ammonium peaks by storing nitrogen intracellularly. In contrast, the strategy of heterotrophic bacterioplankton to acquire ammonium is less well understood. This study revealed the marine bacterium Phaeobacter inhibens DSM 17395, a Roseobacter group member, to have already depleted the external ammonium when only ∼⅓ of the ultimately attained biomass is formed. This was paralleled by a three-fold increase in cellular nitrogen levels and rapid buildup of various nitrogen-containing intracellular metabolites (and enzymes for their biosynthesis) and biopolymers (DNA, RNA and proteins). Moreover, nitrogen-rich cells secreted potential RTX proteins and the antibiotic tropodithietic acid, perhaps to competitively secure pulses of external ammonium and to protect themselves from predation. This complex response may ensure growing cells and their descendants exclusive provision with internal nitrogen stocks. This nutritional strategy appears prevalent also in other roseobacters from distant geographical provenances and could provide a new perspective on the distribution of reduced nitrogen in marine environments, i.e. temporary accumulation in bacterioplankton cells., Phaeobacter inhibens DSM 17395 uncouples NH4+-utilization from growth to build up a broad array of intracellular nitrogen stocks, paralleled by the secretion of RTX-like proteins and the antibiotic tropodithietic acid.

Body size is related to an extensive number of species traits and ecological processes and has therefore been suggested as an effective metric to assess community changes and ecosystem's state. However, the applicability of body size as an ecological indicator in benthic environments has been hindered by the poor knowledge of the factors influencing the size spectra of organisms. By applying biological trait analysis (BTA) and generalized linear models to a species dataset collected in the German Wadden Sea (53A degrees 41'14'' N, 7A degrees 14'19'' E) between 1999 and 2012, we show that the size structure of the macrobenthic community changes predictably along environmental gradients. Specifically, body size increases with increasing current-induced shear stress and sediment organic matter content. In addition, the presence of oyster-mussel reefs in one of the sampling stations enhanced the survival of species belonging to the smallest size categories in habitats with high hydrodynamic energy. This was probably due to the local sheltering effects, which together with biodeposition also increased organic matter in the sediment, likely favoring large deposit feeders as well. Our results suggest that body size can be a useful trait for estimating effects of anthropogenic stressors, such as organic enrichment or alteration of hydrodynamic regime and could therefore be effectively included in current monitoring programs of intertidal macrobenthic communities.

Aquatic and terrestrial ecosystems are tightly linked through the fluxes of organisms and matter. However, aquatic and terrestrial ecologists have mainly studied these ecosystems separately, a "spl ...

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experimentsshow that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of communitycomposition and the potential interaction between diversity at different scales in maintaining ecosystem functioningremains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis ofeight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands?those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)?had higher levelsof multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities. Fil: Hautier, Yann. University of Utrecht; Países Bajos Fil: Isbell, Forest. University of Minnesota; Estados Unidos Fil: Borer, Elizabeth T.. University of Minnesota; Estados Unidos Fil: Seabloom, Eric W.. University of Minnesota; Estados Unidos Fil: Harpole, Stanley. Martin Luther University Halle-Wittenberg; Alemania Fil: Lind, Eric M.. University of Minnesota; Estados Unidos Fil: MacDougall, Andrew. University of Guelph; Canadá Fil: Stevens, Carly. Lancaster University, ; Reino Unido Fil: Adler, Peter. Utah State University; Estados Unidos Fil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Bakker, Jonathan D. University of Washington, ; Estados Unidos Fil: Brudvig, Lars A. Michigan State University; Estados Unidos Fil: Buckley, Yvonne M. Trinity College Dublin; Irlanda Fil: Cadotte, Marc. University of Toronto Scarborough,; Canadá Fil: Caldeira, María C. University of Lisbon; Portugal Fil: Chaneton, Enrique Jose. Universidad de Buenos Aires; Argentina Fil: Chu, Chengjin. Sun Yat-sen University; China Fil: Daleo, Pedro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Dickman, Christopher R. The University of Sydney; Australia Fil: Dwyer, John M. The University of Queensland; Australia Fil: Eskelinen, Anu. University of Oulu; Finlandia Fil: Fay, Philip. USDA-ARS, Grassland Soil and Water Lab; Estados Unidos Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: Hagenah, Nicole. University of KwaZulu-Natal; Sudáfrica Fil: Hillebrand, Helmut. University Oldenburg; Alemania Fil: Iribarne, Oscar Osvaldo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Kirkman, Kevin P.. University of KwaZulu-Natal; Sudáfrica Fil: Knops, Johannes M. H. University of Nebraska, Lincoln; Estados Unidos Fil: La Pierre, Kimberly J. Smithsonian Environmental Research Center; Estados Unidos Fil: McCulley, Rebecca L. University of Kentucky; Estados Unidos Fil: Morgan, John W.. La Trobe University; Australia Fil: Pärtel, Meelis. University of Tartu; Estonia Fil: Pascual, Jesus Maria. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Price, Jodi N. Charles Sturt University; Australia Fil: Prober, Suzanne M. CSIRO Land and Water; Australia Fil: Risch, Anita C. Swiss Federal Institute of Forest, Snow and Landscape Research; Suiza Fil: Sankaran, Mahesh. National Centre for Biological Sciences; India Fil: Schuetz, Martin. Swiss Federal Institute of Forest, Snow and Landscape Research; Suiza Fil: Standish, Rachel J. Murdoch University; Australia Fil: Virtanen, Risto. University of Oulu; Finlandia Fil: Wardle, Glenda M. The University of Sydney; Australia Fil: Yahdjian, María Laura. UBA; Argentina Fil: Hector, Andy. University of Oxford; Reino Unido

Shallow tidal coasts are characterised by shifting tidal flats and emerging or eroding islands above the high tide line. Salt marsh vegetation colonising new habitats distant from existing marshes are an ideal model to investigate metacommunity theory. We installed a set of 12 experimental salt marsh islands made from metal cages on a tidal flat in the German Wadden Sea to study the assembly of salt marsh communities in a metacommunity context. Experimental plots at the same elevation were established within the adjacent salt marsh on the island of Spiekeroog. For both, experimental islands and salt marsh enclosed plots, the same three elevational levels were realised while creating bare patches open for colonisation and vegetated patches with a defined transplanted community. One year into the experiment, the bare islands were colonised by plant species with high fecundity although with a lower frequency compared to the salt marsh enclosed bare plots. Initial plant community variations due to species sorting along the inundation gradient were evident in the transplanted vegetation. Competitive exclusion was not observed and is only expected to unfold in the coming years. Our study highlights that spatially and temporally explicit metacommunity dynamics should be considered in salt marsh plant community assembly and disassembly.

International audience; We examined the relationship between species richness (S) and evenness (J) within a novel, community assembly framework. We hypothesized that environmental stress leads to filtering (increasing the proportional abundance of tolerant species) and taxonomic dispersion (decreasing the number of species within genera and families). Environmental filtering would cause a decline in S by eliminating some stress-sensitive species and a reduction of J by allowing only tolerant species to maintain large populations. Taxonomic relatedness may influence both S and J by controlling the nature of interspecific interactions—positive under taxonomic dispersion versus negative under taxonomic clustering. Therefore, the S–J relationship may be a product of environmental filtering and taxonomic relatedness. We tested this framework with redundancy analyses and structural equation models using continental stream diatom and fish data. We confirmed that (i) environmental stress, defined by watershed forest cover, slope, and temperature, caused filtering (lower sensitive:tolerant species abundance ratios) and taxonomic dispersion (elevated genus:species richness and family:species richness ratios); (ii) S and J, which declined with filtering and taxonomic dispersion, exhibited a positive relationship; and (iii) the role of filtering on J was pronounced only under stressful conditions, while taxonomic dispersion remained an important predictor of J across stressful and favorable environments.

Pan et al . claim that our results actually support a strong linear positive relationship between productivity and richness, whereas Fridley et al . contend that the data support a strong humped relationship. These responses illustrate how preoccupation with bivariate patterns distracts from a deeper understanding of the multivariate mechanisms that control these important ecosystem properties.

The relationship between biodiversity and ecosystem functioning (BEF) and its consequence for ecosystem services has predominantly been studied by controlled, short-term and small-scale experiments under standardized environmental conditions and constant community compositions. However, changes in biodiversity occur in real-world ecosystems with varying environments and a dynamic community composition. In this theme issue, we present novel research on BEF in such dynamic communities. The contributions are organized in three sections on BEF relationships in (i) multi-trophic diversity, (ii) non-equilibrium biodiversity under disturbance and varying environmental conditions, and (iii) large spatial and long temporal scales. The first section shows that multi-trophic BEF relationships often appear idiosyncratic, while accounting for species traits enables a predictive understanding. Future BEF research on complex communities needs to include ecological theory that is based on first principles of species-averaged body masses, stoichiometry and effects of environmental conditions such as temperature. The second section illustrates that disturbance and varying environments have direct as well as indirect (via changes in species richness, community composition and species' traits) effects on BEF relationships. Fluctuations in biodiversity (species richness, community composition and also trait dominance within species) can severely modify BEF relationships. The third section demonstrates that BEF at larger spatial scales is driven by different variables. While species richness per se and community biomass are most important, species identity effects and community composition are less important than at small scales. Across long temporal scales, mass extinctions represent severe changes in biodiversity with mixed effects on ecosystem functions. Together, the contributions of this theme issue identify new research frontiers and answer some open questions on BEF relationships in dynamic communities of real-world landscapes.

Large-scale exploitation of higher trophic levels by humans, together with global-scale nutrient enrichment, highlights the need to explore interactions between predator loss and resource availability. The hypothesis of exploitation ecosystems suggests that top-down and bottom-up control alternate between trophic levels, resulting in a positive relationship between primary production and the abundance of every second trophic level. Specifically, in food webs with three effective trophic levels, primary producers and predators should increase with primary production, while in food webs with two trophic levels, only herbivores should increase. We provided short-term experimental support for these model predictions in a natural benthic community with three effective trophic levels, where the number of algal recruits, but not the biomass of gastropod grazers, increased with algal production. In contrast, when the food web was reduced to two trophic levels by removing larger predators, the number of algal recruits was unchanged while gastropod grazer biomass increased with algal production. Predator removal only affected the consumer-controlled early life-stages of algae, indicating that both the number of trophic levels and the life-stage development of the producer trophic level determine the propagation of trophic cascades in benthic systems. Our results support the hypothesis that predators interact with resource availability to determine food-web structure.

Fraser et al. (Reports, 17 July 2015, p. 302) report a unimodal relationship between productivity and species richness at regional and global scales, which they contrast with the results of Adler et al. (Reports, 23 September 2011, p. 1750). However, both data sets, when analyzed correctly, show clearly and consistently that productivity is a poor predictor of local species richness. EEA Santa Cruz Fil: Tredennick, Andrew T. Utah State University. Department of Wildland Resources and the Ecology Center; Estados Unidos Fil: Adler, Peter B. Utah State University. Department of Wildland Resources and the Ecology Center; Estados Unidos Fil: Grace, James B. U.S. Geological Survey, Wetland and Aquatic Research Center; Estados Unidos Fil: Harpole, W. Stanley. Helmholtz Center for Environmental Research – UFZ. Department of Physiological Diversity; Alemania Fil: Borer, Elizabeth T. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados Unidos Fil: Seabloom, Eric W. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados Unidos Fil: Anderson, T. Michael. Wake Forest University. Department of Biology; Estados Unidos Fil: Bakker, Jonathan D. University of Washington. School of Environmental and Forest Sciences; Estados Unidos Fil: Biederman, Lori A. lowa State University Ecology. Evolution and Organismal Biology; Estados Unidos Fil: Brown, Cynthia S. Colorado State University. Department of Bioagricultural Sciences and Pest Management; Estados Unidos Fil: Buckley, Yvonne M. University of Dublin. School of Natural Sciences. Trinity College Dublin. Zoology; Irlanda. Fil: Chu, Chengjin. Sun Yat-sen University. School of Life Sciences; China. Fil: Collins, Scott L. University of New Mexico. Department of Biology; Estados Unidos Fil: Crawley, Michael J. Imperial College London. Department of Biology; Reino Unido Fil: Fay, Philip A. USDA-ARS. Grassland, Soil, and Water Research Laboratory; Estados Unidos Fil: Firn, Jennifer. Queensland University of Technology. School of Earth, Environmental and Biological Sciences; Australia. Fil: Gruner, Daniel S. University of Maryland. Department of Entomology. Plant Sciences; Estados Unidos Fil: Hagenah, Nicole. University of KwaZulu-Natal. School of Life Sciences; Sudáfrica Fil: Hautier, Yann. Utrecht University. Department of Biology, Ecology and Biodiversity group; Países Bajos Fil: Hector, Andy. University of Oxford. Department of Plant Sciences; Reino Unido Fil: Hillebrand, Helmut. Carl-von-Ossietzky University. Institute for Chemistry and Biology of the Marine Environment; Alemania Fil: Kirkman, Kevin. University of KwaZulu-Natal. School of Life Sciences; Sudáfrica Fil: Knops, Johannes M. H. University of Nebraska. School of Biological Sciences; Estados Unidos Fil: Laungani, Ramesh. Doane College. Biology Department; Estados Unidos Fil: Lind, Eric M. University of Minnesota. Department of Ecology, Evolution and Behavior; Estados Unidos. Fil: MacDougall, Andrew S. University of Guelph. Department of Integrative Biology; Canadá. Fil: McCulley, Rebecca L. University of Kentucky. Department of Plant and Soil Science; Estados Unidos Fil: Mitchell, Charles E. University of North Carolina at Chapel Hill. Department of Biology; Estados Unidos Fil: Moore, Joslin L. Monash University. School of Biological Sciences; Australia. Fil: Morgan, John W. La Trobe University. Department of Ecology, Environment and Evolution; Australia. Fil: Orrock, John L. University of Wisconsin. Department of Zoology; Estados Unidos Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina. Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina. Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Prober, Suzanne M. Commonwealth Scientific and Industrial Research Organisation Land and Water; Australia. Fil: Risch, Anita C. Swiss Federal Institute for Forest, Snow and Landscape Research. Community Ecology; Suiza Fil: Schütz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research. Community Ecology; Suiza Fil: Speziale, Karina L. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones en Biodiversidad y Medioambiente (INIBIOMA). Universidad Nacional del Comahue; Argentina. Fil: Standish, Rachel J. Murdoch University. School of Veterinary and Life Sciences; Australia. Fil: Sullivan, Lauren L. University of Minnesota. Department of Ecology, Evolution and Behavior; Estados Unidos Fil: Wardle, Glenda M. University of Sydney. School of Biological Sciences; Australia. Fil: Williams, Ryan J. Iowa State University. Agricultural and Biosystems Engineering; Estados Unidos Fil: Yang, Louie H. University of California. Department of Entomology and Nematology; Estados Unidos

The North Sea region contains a vast number of lakes; from shallow, highly eutrophic water bodies in agricultural areas to deep, oligotrophic systems in pristine high-latitude or high-altitude areas. These freshwaters and the biota they contain are highly vulnerable to climate change. As largely closed systems, lakes are ideally suited to studying climate-induced effects via changes in ice cover, hydrology and temperature, as well as via biological communities (phenology, species and size distribution, food-web dynamics, life-history traits, growth and respiration, nutrient dynamics and ecosystem metabolism). This chapter focuses on change in natural lakes and on parameters for which their climate-driven responses have major impacts on ecosystem properties such as productivity, community composition, metabolism and biodiversity. It also points to the importance of addressing different temporal scales and variability in driving and response variables along with threshold-driven responses to environmental forces. Exceedance of critical thresholds may result in abrupt changes in particular elements of an ecosystem. Modelling climate-driven physical responses like ice-cover duration, stratification periods and thermal profiles in lakes have shown major advances, and the chapter provide recent achievements in this field for northern lakes. Finally, there is a tentative summary of the level of certainty for key climatic impacts on freshwater ecosystems. Wherever possible, data and examples are drawn from the North Sea region.

Effects of temperature changes on phytoplankton communities seem to be highly context-specific, but few studies have analyzed whether this context specificity depends on differences in the abiotic conditions or in species composition between studies. We present an experiment that allows disentangling the contribution of abiotic and biotic differences in shaping the response to two aspects of temperature change: permanent increase of mean temperature versus pulse disturbance in form of a heat wave. We used natural communities from six different sites of a floodplain system as well as artificially mixed communities from laboratory cultures and grew both, artificial and natural communities, in water from the six different floodplain lakes (sites). All 12 contexts (2 communities × 6 sites) were first exposed to three different temperature levels (12, 18, 24 °C, respectively) and afterward to temperature pulses (4 °C increase for 7 h day−1). Temperature-dependent changes in biomass and community composition depended on the initial composition of phytoplankton communities. Abiotic conditions had a major effect on biomass of phytoplankton communities exposed to different temperature conditions, however, the effect of biotic and abiotic conditions together was even more pronounced. Additionally, phytoplankton community responses to pulse temperature effects depended on the warming history. By disentangling abiotic and biotic effects, our study shows that temperature-dependent effects on phytoplankton communities depend on both, biotic and abiotic constraints. Electronic supplementary material The online version of this article (doi:10.1007/s00442-016-3693-3) contains supplementary material, which is available to authorized users.

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.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands., It remains unclear whether exotic and native species are functionally different. Using a global grassland experiment, Seabloom et al. show that native and exotic species respond differently to two globally pervasive environmental changes, addition of mineral nutrients and alteration of herbivore density.

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. peerReviewed

Die Artenvielfalt von Pflanzen strukturiert die Gesellschaften höherer trophischer Ebenen und verbessert die Fortpflanzung und Fitness von Pflanzen-assoziierten Insekten. Dieser Einfluss auf höhere trophische Ebenen kann von der Quantität (Verfügbarkeit) und Qualität (Zusammensetzung) der Ressourcen abhängen. Wir untersuchten die Stöchiometrie von Interaktionen zwischen Pflanzen und zwei funktionellen Insektengruppen (Herbivoren, Bestäuber) entlang des experimentellen Pflanzendiversitätsgradienten im „Jena-Experiment“. Wir analysierten den prozentuellen Anteil von Kohlenstoff (C), Stickstoff (N) und Phosphat (P), sowie die entsprechenden molaren Verhältnisse dieser Elemente (C:N, C:P, N:P) bei einer bestäubenden Bienenart (Chelostoma distinctum) und einer herbivoren Heuschreckenart (Chorthippus parallelus), welche wir auf Graslandflächen verschiedener Diversitätsstufen gezüchtet haben. Insekten zeigten höhere prozentuale Anteile in der Trockenmasse (C, N und P) und niedrigere C:N- und C:P-Verhältnisse als Pflanzen. Das C:N-Verhältnis war bei Heuschrecken signifikant höher als bei den Bienen, und in beiden Gruppen bei den Weibchen höher als bei den Männchen. Zunehmende Pflanzenvielfalt erhöhte das C:N-Verhältnis von männlichen Bienen und weiblichen Heuschrecken und die Insektenstöchiometrie beider Gruppen hängte positiv mit der Pflanzenstöchiometrie zusammen (männliche Bienen: C:P und N:P; Heuschrecken: C:N und N:P). Unsere Studie zeigt unterschiedliche Stöchiometrie von antagonistischen und mutualistischen Interaktionen. Daraus ergeben sich verschiedene Schlussfolgerungen für diversitätsgetriebene Veränderungen von der Elementzusammensetzung bei Pflanzen verschiedener trophischer und funktioneller Gruppen. Changes in plant diversity have consequences for higher trophic levels, e.g., higher plant diversity can enhance the reproduction and fitness of plant-associated insects. This response of higher trophic levels potentially depends on diversity-related changes in both resource quantity (abundance) and quality (nutritional content). The availability of elemental nutrients in plant resources is one aspect of nutritional quality, but has rarely been addressed as a pathway relating plant diversity to associated insects. Using the experimental plant diversity gradient of a large biodiversity grassland project, the Jena-Experiment, we analysed the %C, %N and %P and the molar ratios of those elements (C:N, C:P and N:P) in a pollinating bee, Chelostoma distinctum, and an herbivorous grasshopper, Chorthippus parallelus, reared on plots of different plant diversity. Insects showed higher content of C, N and P (% dry mass), and lower C:N and C:P ratios than plants. C:N ratios were significantly higher in grasshoppers than in bees and higher in females than in males of both species. Increasing plant species richness increased the C:N ratio of male bees and female grasshoppers. In both groups, stoichiometry was positively related to plant stoichiometry (male bees: C:P and N:P; grasshoppers: C:N and N:P). Path analysis revealed that diversity-driven changes in plant elemental composition can have consequences for abundance and chemical composition of higher trophic levels, with different responses of the two functional groups.

Ecological stoichiometry (ES) has become one of the most pervasive theoretical frameworks in environmental sciences and biology in the last two decades. ES allows predicting processes on all organizational levels from subcellular structures to ecosystems by relating the elemental composition and demand of organisms to the relative availability of resources. However, ES has been rarely used to understand and predict the relationship between biodiversity and ecosystem functioning (BEF), although ES would be ideally suited as it makes predictions on both population processes underlying biodiversity as well as on matter transformations underlying ecosystem processes. Here, we propose to link the two fields of research on ES and BEF relationships and highlight a number of potential avenues for further research. First, we cast a stoichiometric view on drivers of biodiversity change. Second, we address the stoichiometric underpinning of biodiversity–productivity relationships. Third, we discuss potential interactions between stoichiometry and diversity in a food web context.

Ecosystem functioning is affected by horizontal (within trophic groups) and vertical (across trophic levels) biodiversity. Theory predicts that the effects of vertical biodiversity depend on consumer specialization. In a microcosm experiment, we investigated ciliate consumer diversity and specialization effects on algal prey biovolume, evenness and composition, and on ciliate biovolume production. The experimental data was complemented by a process-based model further analyzing the ecological mechanisms behind the observed diversity effects. Overall, increasing consumer diversity had no significant effect on prey biovolume or evenness. However, consumer specialization affected the prey community. Specialist consumers showed a stronger negative impact on prey biovolume and evenness than generalists. The model confirmed that this pattern was mainly driven by a single specialist with a high per capita grazing rate, consuming the two most productive prey species. When these were suppressed, the prey assemblage became dominated by a less productive species, consequently decreasing prey biovolume and evenness. Consumer diversity increased consumer biovolume, which was stronger for generalists than for specialists and highest in mixed combinations, indicating that consumer functional diversity, i.e. more diverse feeding strategies, increased resource use efficiency. Overall, our results indicate that consumer diversity effects on prey and consumers strongly depend on species-specific growth and grazing rates, which may be at least equally important as consumer specialization in driving consumer diversity effects across trophic levels. Synthesis In a microcosm experiment, we investigated multitrophic consumer diversity and specialization effects using ciliate consumers and microalgal prey. Consumer diversity increased consumer biovolume, which was highest in combinations containing both generalists and specialists. Specialist consumers showed a stronger negative effect on prey biovolume and evenness than generalists. These experimental data were supported by a process-based model, indicating that the large effect of the specialists was based on high per capita grazing rate on the two most productive prey species. Species-specific traits such as growth and grazing rates were equally important for multitrophic diversity effects than consumer specialization.