Global change drives modern plankton communities away from the pre-industrial state.

The ocean-the Earth's largest ecosystem-is increasingly affected by anthropogenic climate change1,2. Large and globally consistent shifts have been detected in species phenology, range extension and community composition in marine ecosystems3-5. However, despite evidence for ongoing change, it remains unknown whether marine ecosystems have entered an Anthropocene6 state beyond the natural decadal to centennial variability. This is because most observational time series lack a long-term baseline, and the few time series that extend back into the pre-industrial era have limited spatial coverage7,8. Here we use the unique potential of the sedimentary record of planktonic foraminifera-ubiquitous marine zooplankton-to provide a global pre-industrial baseline for the composition of modern species communities. We use a global compilation of 3,774 seafloor-derived planktonic foraminifera communities of pre-industrial age9 and compare these with communities from sediment-trap time series that have sampled plankton flux since AD 1978 (33 sites, 87 observation years). We find that the Anthropocene assemblages differ from their pre-industrial counterparts in proportion to the historical change in temperature. We observe community changes towards warmer or cooler compositions that are consistent with historical changes in temperature in 85% of the cases. These observations not only confirm the existing evidence for changes in marine zooplankton communities in historical times, but also demonstrate that Anthropocene communities of a globally distributed zooplankton group systematically differ from their unperturbed pre-industrial state.

Estimation of functional diversity and species traits from ecological monitoring data.

The twin crises of climate change and biodiversity loss define a strong need for functional diversity monitoring. While the availability of high-quality ecological monitoring data is increasing, the quantification of functional diversity so far requires the identification of species traits, for which data are harder to obtain. However, the traits that are relevant for the ecological function of a species also shape its performance in the environment and hence, should be reflected indirectly in its spatiotemporal distribution. Thus, it may be possible to reconstruct these traits from a sufficiently extensive monitoring dataset. Here, we use diffusion maps, a deterministic and de facto parameter-free analysis method, to reconstruct a proxy representation of the species' traits directly from monitoring data and use it to estimate functional diversity. We demonstrate this approach with both simulated data and real-world phytoplankton monitoring data from the Baltic Sea. We anticipate that wider application of this approach to existing data could greatly advance the analysis of changes in functional biodiversity.

Failures to disagree are essential for environmental science to effectively influence policy development.

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 polarised 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 four 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.

Elemental and biochemical nutrient limitation of zooplankton: A meta-analysis.

Primary consumers in aquatic ecosystems are frequently limited by the quality of their food, often expressed as phytoplankton elemental and biochemical composition. However, the effects of these food quality indicators vary across studies, and we lack an integrated understanding of how elemental (e.g. nitrogen, phosphorus) and biochemical (e.g. fatty acid, sterol) limitations interactively influence aquatic food webs. Here, we present the results of a meta-analysis using >100 experimental studies, confirming that limitation by N, P, fatty acids, and sterols all have significant negative effects on zooplankton performance. However, effects varied by grazer response (growth vs. reproduction), specific manipulation, and across taxa. While P limitation had greater effects on zooplankton growth than fatty acids overall, P and fatty acid limitation had equal effects on reproduction. Furthermore, we show that: nutrient co-limitation in zooplankton is strong; effects of essential fatty acid limitation depend on P availability; indirect effects induced by P limitation exceed direct effects of mineral P limitation; and effects of nutrient amendments using laboratory phytoplankton isolates exceed those using natural field communities. Our meta-analysis reconciles contrasting views about the role of various food quality indicators, and their interactions, for zooplankton performance, and provides a mechanistic understanding of trophic transfer in aquatic environments.

Low statistical power and overestimated anthropogenic impacts, exacerbated by publication bias, dominate field studies in global change biology.

Field studies are essential to reliably quantify ecological responses to global change because they are exposed to realistic climate manipulations. Yet such studies are limited in replicates, resulting in less power and, therefore, potentially unreliable effect estimates. Furthermore, while manipulative field experiments are assumed to be more powerful than non-manipulative observations, it has rarely been scrutinized using extensive data. Here, using 3847 field experiments that were designed to estimate the effect of environmental stressors on ecosystems, we systematically quantified their statistical power and magnitude (Type M) and sign (Type S) errors. Our investigations focused upon the reliability of field experiments to assess the effect of stressors on both ecosystem's response magnitude and variability. When controlling for publication bias, single experiments were underpowered to detect response magnitude (median power: 18%-38% depending on effect sizes). Single experiments also had much lower power to detect response variability (6%-12% depending on effect sizes) than response magnitude. Such underpowered studies could exaggerate estimates of response magnitude by 2-3 times (Type M errors) and variability by 4-10 times. Type S errors were comparatively rare. These observations indicate that low power, coupled with publication bias, inflates the estimates of anthropogenic impacts. Importantly, we found that meta-analyses largely mitigated the issues of low power and exaggerated effect size estimates. Rather surprisingly, manipulative experiments and non-manipulative observations had very similar results in terms of their power, Type M and S errors. Therefore, the previous assumption about the superiority of manipulative experiments in terms of power is overstated. These results call for highly powered field studies to reliably inform theory building and policymaking, via more collaboration and team science, and large-scale ecosystem facilities. Future studies also require transparent reporting and open science practices to approach reproducible and reliable empirical work and evidence synthesis.

Integrative modelling reveals mechanisms linking productivity and plant species richness.

How ecosystem productivity and species richness are interrelated is one of the most debated subjects in the history of ecology. Decades of intensive study have yet to discern the actual mechanisms behind observed global patterns. Here, by integrating the predictions from multiple theories into a single model and using data from 1,126 grassland plots spanning five continents, we detect the clear signals of numerous underlying mechanisms linking productivity and richness. We find that an integrative model has substantially higher explanatory power than traditional bivariate analyses. In addition, the specific results unveil several surprising findings that conflict with classical models. These include the isolation of a strong and consistent enhancement of productivity by richness, an effect in striking contrast with superficial data patterns. Also revealed is a consistent importance of competition across the full range of productivity values, in direct conflict with some (but not all) proposed models. The promotion of local richness by macroecological gradients in climatic favourability, generally seen as a competing hypothesis, is also found to be important in our analysis. The results demonstrate that an integrative modelling approach leads to a major advance in our ability to discern the underlying processes operating in ecological systems.

Addition of multiple limiting resources reduces grassland diversity.

Niche dimensionality provides a general theoretical explanation for biodiversity-more niches, defined by more limiting factors, allow for more ways that species can coexist. Because plant species compete for the same set of limiting resources, theory predicts that addition of a limiting resource eliminates potential trade-offs, reducing the number of species that can coexist. Multiple nutrient limitation of plant production is common and therefore fertilization may reduce diversity by reducing the number or dimensionality of belowground limiting factors. At the same time, nutrient addition, by increasing biomass, should ultimately shift competition from belowground nutrients towards a one-dimensional competitive trade-off for light. Here we show that plant species diversity decreased when a greater number of limiting nutrients were added across 45 grassland sites from a multi-continent experimental network. The number of added nutrients predicted diversity loss, even after controlling for effects of plant biomass, and even where biomass production was not nutrient-limited. We found that elevated resource supply reduced niche dimensionality and diversity and increased both productivity and compositional turnover. Our results point to the importance of understanding dimensionality in ecological systems that are undergoing diversity loss in response to multiple global change factors.

The impact of climate warming on species diversity across scales: Lessons from experimental meta-ecosystems.

Aim: The aim was to evaluate the effects of climate warming on biodiversity across spatial scales (i.e., α-, ß- and γ-diversity) and the effects of patch openness and experimental context on diversity responses. Location: Global. Time period: 1995-2017. Major taxa studied: Fungi, invertebrates, phytoplankton, plants, seaweed, soil microbes and zooplankton. Methods: We compiled data from warming experiments and conducted a meta-analysis to evaluate the effects of warming on different components of diversity (such as species richness and equivalent numbers) at different spatial scales (α-, ß- and γ-diversity, partitioning ß-diversity into species turnover and nestedness components). We also investigated how these effects were modulated by system openness, defined as the possibility of replicates being colonized by new species, and experimental context (duration, mean temperature change and ecosystem type). Results: Experimental warming did not affect local species richness (α-diversity) but decreased effective numbers of species by affecting species dominance. Warming increased species spatial turnover (ß-diversity), although no significant changes were detected at the regional scale (γ-diversity). Site openness and experimental context did not significantly affect our results, despite significant heterogeneity in the effect sizes of α- and ß-diversity. Main conclusions: Our meta-analysis shows that the effects of warming on biodiversity are scale dependent. The local and regional inventory diversity remain unaltered, whereas species composition across temperature gradients and the patterns of species dominance change with temperature, creating novel communities that might be harder to predict.

Effects of site-selection bias on estimates of biodiversity change.

Estimates of biodiversity change are essential for the management and conservation of ecosystems. Accurate estimates rely on selecting representative sites, but monitoring often focuses on sites of special interest. How such site-selection biases influence estimates of biodiversity change is largely unknown. Site-selection bias potentially occurs across four major sources of biodiversity data, decreasing in likelihood from citizen science, museums, national park monitoring, and academic research. We defined site-selection bias as a preference for sites that are either densely populated (i.e., abundance bias) or species rich (i.e., richness bias). We simulated biodiversity change in a virtual landscape and tracked the observed biodiversity at a sampled site. The site was selected either randomly or with a site-selection bias. We used a simple spatially resolved, individual-based model to predict the movement or dispersal of individuals in and out of the chosen sampling site. Site-selection bias exaggerated estimates of biodiversity loss in sites selected with a bias by on average 300-400% compared with randomly selected sites. Based on our simulations, site-selection bias resulted in positive trends being estimated as negative trends: richness increase was estimated as 0.1 in randomly selected sites, whereas sites selected with a bias showed a richness change of -0.1 to -0.2 on average. Thus, site-selection bias may falsely indicate decreases in biodiversity. We varied sampling design and characteristics of the species and found that site-selection biases were strongest in short time series, for small grains, organisms with low dispersal ability, large regional species pools, and strong spatial aggregation. Based on these findings, to minimize site-selection bias, we recommend use of systematic site-selection schemes; maximizing sampling area; calculating biodiversity measures cumulatively across plots; and use of biodiversity measures that are less sensitive to rare species, such as the effective number of species. Awareness of the potential impact of site-selection bias is needed for biodiversity monitoring, the design of new studies on biodiversity change, and the interpretation of existing data.

Efectos del Sesgo en la Selección de Sitio sobre las Estimaciones del Cambio en la Biodiversidad Resumen Las estimaciones del cambio en la biodiversidad son esenciales para el manejo y la conservación de los ecosistemas. Las estimaciones precisas dependen de la selección de sitios representativos pero su monitoreo con frecuencia se enfoca en los sitios de interés especial. En su mayoría se desconoce cómo influyen tales sesgos en la selección de sitios sobre las estimaciones del cambio en la biodiversidad. El sesgo en la selección de sitios ocurre potencialmente en cuatro fuentes principales de datos sobre biodiversidad, disminuyendo en probabilidad cuando los datos vienen de la ciencia ciudadana, museos, el monitoreo de los parques nacionales y la investigación académica. Definimos al sesgo en la selección de sitios como la preferencia por sitios que están densamente poblados (es decir, sesgo por abundancia) o que son ricos en especies (es decir, sesgo por riqueza). Simulamos el cambio en la biodiversidad en un paisaje virtual y le dimos seguimiento a la biodiversidad observada en un sitio muestreado. El sitio fue seleccionado al azar o con un sesgo en la selección de sitio. Usamos un modelo simple basado en los individuos y resuelto espacialmente para predecir el movimiento o la dispersión de los individuos dentro y fuera del sitio de muestreo elegido. El sesgo en la selección de sitio exageró las estimaciones de la pérdida de la biodiversidad en los sitios seleccionados con un sesgo en promedio de 300-400% en comparación con sitios seleccionados al azar. Con base en nuestras simulaciones, el sesgo en la selección de sitio derivó en que las tendencias positivas se estimaran como tendencias negativas: se estimó que el incremento en la riqueza fue de 0.1 en sitios seleccionados al azar, mientras que en los sitios seleccionados con un sesgo mostraron un cambio en la riqueza de −0.1 a −0.2 en promedio. Así, el sesgo en la selección de sitio puede indicar erróneamente la existencia de disminuciones en la biodiversidad. Variamos el diseño del muestreo y las características de las especies y encontramos que los sesgos en la selección de sitio estaban más consolidados en las series de tiempo corto, para los granos pequeños, organismos con una baja habilidad de dispersión, grandes patrimonios genéticos de especies regionales y una agregación espacial fuerte. Con base en estos resultados, para lograr minimizar el sesgo en la selección de sitio, recomendamos usar esquemas sistemáticos de selección de sitio; maximizar el área de muestreo; calcular las medidas de biodiversidad acumulativamente en los lotes; y usar las medidas de biodiversidad que son menos sensibles a las especies raras, como el número efectivo de especies. Se necesita tener conciencia sobre el impacto potencial del sesgo en la selección de sitio para el monitoreo de la biodiversidad, el diseño de nuevos estudios sobre el cambio en la biodiversidad y la interpretación de los datos existentes.

Meta-analysis on pulse disturbances reveals differences in functional and compositional recovery across ecosystems.

Most ecosystems are affected by anthropogenic or natural pulse disturbances, which alter the community composition and functioning for a limited period of time. Whether and how quickly communities recover from such pulses is central to our understanding of biodiversity dynamics and ecosystem organisation, but also to nature conservation and management. Here, we present a meta-analysis of 508 (semi-)natural field experiments globally distributed across marine, terrestrial and freshwater ecosystems. We found recovery to be significant yet incomplete. At the end of the experiments, disturbed treatments resembled controls again when considering abundance (94%), biomass (82%), and univariate diversity measures (88%). Most disturbed treatments did not further depart from control after the pulse, indicating that few studies showed novel trajectories induced by the pulse. Only multivariate community composition on average showed little recovery: disturbed species composition remained dissimilar to the control throughout most experiments. Still, when experiments revealed a higher compositional stability, they tended to also show higher functional stability. Recovery was more complete when systems had high resistance, whereas resilience and resistance were negatively correlated. The overall results were highly consistent across studies, but significant differences between ecosystems and organism groups appeared. Future research on disturbances should aim to understand these differences, but also fill obvious gaps in the empirical assessments for regions (especially the tropics), ecosystems and organisms. In summary, we provide general evidence that (semi-)natural communities can recover from pulse disturbances, but compositional aspects are more vulnerable to long-lasting effects of pulse disturbance than the emergent functions associated to them.

Eutrophication weakens stabilizing effects of diversity in natural grasslands.

Studies of experimental grassland communities have demonstrated that plant diversity can stabilize productivity through species asynchrony, in which decreases in the biomass of some species are compensated for by increases in others. However, it remains unknown whether these findings are relevant to natural ecosystems, especially those for which species diversity is threatened by anthropogenic global change. Here we analyse diversity-stability relationships from 41 grasslands on five continents and examine how these relationships are affected by chronic fertilization, one of the strongest drivers of species loss globally. Unmanipulated communities with more species had greater species asynchrony, resulting in more stable biomass production, generalizing a result from biodiversity experiments to real-world grasslands. However, fertilization weakened the positive effect of diversity on stability. Contrary to expectations, this was not due to species loss after eutrophication but rather to an increase in the temporal variation of productivity in combination with a decrease in species asynchrony in diverse communities. Our results demonstrate separate and synergistic effects of diversity and eutrophication on stability, emphasizing the need to understand how drivers of global change interactively affect the reliable provisioning of ecosystem services in real-world systems.

Herbivores and nutrients control grassland plant diversity via light limitation.

Human alterations to nutrient cycles and herbivore communities are affecting global biodiversity dramatically. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.

Scale Both Confounds and Informs Characterization of Species Coexistence in Empirical Systems.

Identifying stable coexistence in empirical systems is notoriously difficult. Here, we show how spatiotemporal structure and complex system dynamics can confound two commonly used stability metrics in empirical contexts: response to perturbation and invasion rate when rare. We use these metrics to characterize stable coexistence across a range of spatial and temporal scales for five simulated models in which the ability of species to coexist in the long term is known a priori and for an empirical old field successional time series. We term the resulting multivariate distribution of metrics a "stability fingerprint." In accordance with a wide range of classic and recent studies, our results demonstrate that no combination of empirically tractable metrics or measurements is guaranteed to "correctly" characterize coexistence. However, we also find that heuristic information from the stability fingerprint can be used to broadly characterize dynamic behavior and identify circumstances under which particular combinations of species are likely to persist. Moreover, stability fingerprints appear to be particularly well suited for matching potential theoretical models to observed dynamics. These findings suggest that it may be prudent to shift the focus of empirical stability analysis away from quantifying single measures of stability and toward more heuristic, multivariate characterizations of community dynamics.

Metaecosystem Dynamics of Marine Phytoplankton Alters Resource Use Efficiency along Stoichiometric Gradients.

Metaecosystem theory addresses the link between local (within habitats) and regional (between habitats) dynamics by simultaneously analyzing spatial community ecology and abiotic matter flow. Here we experimentally address how spatial resource gradients and connectivity affect resource use efficiency (RUE) and stoichiometry in marine phytoplankton as well as the community composition at local and regional scales. We created gradostat metaecosystems consisting of five linearly interconnected patches, which were arranged either in countercurrent gradients of nitrogen (N) and phosphorus (P) supply or with a uniform spatial distribution of nutrients and which had either low or high connectivity. Gradient metaecosystems were characterized by higher remaining N and P concentrations (and N∶P ratios) than uniform ones, a difference reduced by higher connectivity. The position of the patch in the gradient strongly constrained elemental stoichiometry, local biovolume production, and RUE. As expected, algal carbon (C)∶N, biovolume, and N-specific RUE decreased toward the N-rich end of the gradient metaecosystem, whereas the opposite was observed for most of the gradient for C∶P, N∶P, and P-specific RUE. However, at highest N∶P supply, unexpectedly low C∶P, N∶P, and P-specific RUE values were found, indicating that the low availability of P inhibited efficient use of N and biovolume production. Consequently, gradient metaecosystems had lower overall biovolume at the regional scale. Whereas treatment effects on local richness were weak, gradients were characterized by higher dissimilarity in species composition. Thus, the stoichiometry of resource supply and spatial connectivity between patches appeared as decisive elements constraining phytoplankton composition and functioning in metaecosystems.

Biodiversity-ecosystem functioning relationships in fish communities: biomass is related to evenness and the environment, not to species richness.

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.

Decomposing multiple dimensions of stability in global change experiments.

Ecological stability is the central framework to understand an ecosystem's ability to absorb or recover from environmental change. Recent modelling and conceptual work suggests that stability is a multidimensional construct comprising different response aspects. Using two freshwater mesocosm experiments as case studies, we show how the response to single perturbations can be decomposed in different stability aspects (resistance, resilience, recovery, temporal stability) for both ecosystem functions and community composition. We find that extended community recovery is tightly connected to a nearly complete recovery of the function (biomass production), whereas systems with incomplete recovery of the species composition ranged widely in their biomass compared to controls. Moreover, recovery was most complete when either resistance or resilience was high, the latter associated with low temporal stability around the recovery trend. In summary, no single aspect of stability was sufficient to reflect the overall stability of the system.

Integrating community assembly and biodiversity to better understand ecosystem function: the Community Assembly and the Functioning of Ecosystems (CAFE) approach.

The research of a generation of ecologists was catalysed by the recognition that the number and identity of species in communities influences the functioning of ecosystems. The relationship between biodiversity and ecosystem functioning (BEF) is most often examined by controlling species richness and randomising community composition. In natural systems, biodiversity changes are often part of a bigger community assembly dynamic. Therefore, focusing on community assembly and the functioning of ecosystems (CAFE), by integrating both species richness and composition through species gains, losses and changes in abundance, will better reveal how community changes affect ecosystem function. We synthesise the BEF and CAFE perspectives using an ecological application of the Price equation, which partitions the contributions of richness and composition to function. Using empirical examples, we show how the CAFE approach reveals important contributions of composition to function. These examples show how changes in species richness and composition driven by environmental perturbations can work in concert or antagonistically to influence ecosystem function. Considering how communities change in an integrative fashion, rather than focusing on one axis of community structure at a time, will improve our ability to anticipate and predict changes in ecosystem function.

Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation.

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change - fertilisation and herbivore loss - are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics.

Warming and oligotrophication cause shifts in freshwater phytoplankton communities.

While there is a lot of data on interactive effects of eutrophication and warming, to date, we lack data to generate reliable predictions concerning possible effects of nutrient decrease and temperature increase on community composition and functional responses. In recent years, a wide-ranging trend of nutrient decrease (re-oligotrophication) was reported for freshwater systems. Small lakes and ponds, in particular, show rapid responses to anthropogenic pressures and became model systems to investigate single as well as synergistic effects of warming and fertilization in situ and in experiments. Therefore, we set up an experiment to investigate the single as well as the interactive effects of nutrient reduction and gradual temperature increase on a natural freshwater phytoplankton community, using an experimental indoor mesocosm setup. Biomass production initially increased with warming but decreased with nutrient depletion. If nutrient supply was constant, biomass increased further, especially under warming conditions. Under low nutrient supply, we found a sharp transition from initially positive effects of warming to negative effects when resources became scarce. Warming reduced phytoplankton richness and evenness, whereas nutrient reduction at ambient temperature had positive effects on diversity. Our results indicate that temperature effects on freshwater systems will be altered by nutrient availability. These interactive effects of energy increase and resource decrease have major impacts on biodiversity and ecosystem function and thus need to be considered in environmental management plans.

Genetic variation of a foundation rockweed species affects associated communities.

Genetic variation in a foundation species may affect the composition of associated communities as well as modify ecosystem function. While the ecological consequences of genetic diversity of foundation species have been widely reported, the ability of individual genotypes to support dissimilar communities has been documented only in forest ecosystems. Here, for the first time in a marine ecosystem, we test whether the different genotypes of the rockweed Fucus vesiculosus harbor distinct community phenotypes and whether the genetic similarity of individual genotypes or their defensive compound content can explain the variation of the associated communities. We reared replicated genotypes in a common garden in the sea and analyzed their associated communities of periphytic algae and invertebrates as well as determined their contents of defense compounds, phlorotannins, and genetic distance based on neutral molecular markers. The periphytic community was abundant in mid-summer and its biovolume, diversity and community composition varied among the rockweed genotypes. The diversity of the periphytic community decreased with its increasing biovolume. In autumn, when grazers were abundant, periphytic community biomass was lower and less variable among rockweed genotypes, indicating different relative importance of bottom-up regulation through heritable variation of the foundation species and top-down regulation through grazing intensity. Similarly, composition of the invertebrate community varied among the rockweed genotypes. Although the genotype explained about 10-18% of the variation in associated communities, the variation was explained neither by the genetic distance nor the phlorotannin content. Thus, neither neutral genetic markers nor a single phenotypic trait could provide a mechanistic understanding of the genetic basis of community specificity. Therefore, a more comprehensive mapping of quantitative trait variation is needed to understand the underlying mechanisms. The community specificity implies that genetic variation within a foundation species is crucial for the biodiversity and assembly of associated organisms and, thus, for the functioning of associated communities. The result highlights the importance of ensuring the genetic variation of foundation species as a conservation target.