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The high number of freshwater species at low salinity and the correspondingly high number of marine species at high salinity enveloping a conspicuous richness minimum at intermediate salinities has shaped our basic understanding of biodiversity along a coastal salinity gradient for almost 80 years. Visualized as the Remane curve, this iconic concept was originally based on sedentary macroinvertebrates in the Baltic Sea. To what extent the concept can be generalized, particularly to free-drifting organisms, is currently debated. Here we use approximately 16,000 phytoplankton samples from two large coastal ecosystems-the Baltic Sea and Chesapeake Bay-to analyze the relationship between salinity and phytoplankton species richness. Alpha diversity showed a consistent variation along the salinity gradient, with a minimum at mesohaline salinities of around 7-9. Rarefied species pools at narrow salinity intervals also showed reduced diversity at intermediate salinities, surrounded by high richness toward both ends of the gradient. The cumulative likelihood of species presence validated the minimum at intermediate salinities. Community composition changed abruptly at the α diversity minimum in the Baltic Sea, while it changed gradually along the salinity gradient in Chesapeake Bay. We conclude that the Remane concept is in every respect valid for phytoplankton.

Recent research has highlighted that positive biodiversity-ecosystem functioning relationships hold for all groups of organisms, including microbes. Yet, we still lack understanding regarding the drivers of microbial diversity, in particular, whether diversity of microbial communities is a matter of local factors, or whether metacommunities are of similar importance to what is known from higher organisms. Here, we explore the driving forces behind spatial variability in lake phytoplankton diversity in Fennoscandia. While phytoplankton biovolume is best predicted by local phosphorus concentrations, phytoplankton diversity (measured as genus richness, G) only showed weak correlations with local concentrations of total phosphorus. By estimating spatial averages of total phosphorus concentrations on various scales from an independent, spatially representative lake survey, we found that close to 70 per cent of the variability in local phytoplankton diversity can be explained by regionally averaged phosphorus concentrations on a scale between 100 and 400 km. Thus, the data strongly indicate the existence of metacommunities on this scale. Furthermore, we show a strong dependency between lake productivity and spatial community turnover. Thus, regional productivity affects beta-diversity by controlling spatial community turnover, resulting in scale-dependent productivity-diversity relationships. As an illustration of the interaction between local and regional processes in shaping microbial diversity, our results offer both empirical support and a plausible mechanism for the existence of common scaling rules in both the macrobial and the microbial worlds. We argue that awareness of regional species pools in phytoplankton and other unicellular organisms may critically improve our understanding of ecosystems and their susceptibility to anthropogenic stressors.

The relationship between species diversity and ecosystem functioning has been debated for decades, especially in relation to the "macroscopic" realm (higher plants and metazoans). Although there is emerging consensus that diversity enhances productivity and stability in communities of higher organisms; however, we still do not know whether these relationships apply also for communities of unicellular organisms, such as phytoplankton, which contribute approximately 50% to the global primary production. We show here that phytoplankton resource use, and thus carbon fixation, is directly linked to the diversity of phytoplankton communities. Datasets from freshwater and brackish habitats show that diversity is the best predictor for resource use efficiency of phytoplankton communities across considerable environmental gradients. Furthermore, we show that the diversity requirement for stable ecosystem functioning scales with the nutrient level (total phosphorus), as evidenced by the opposing effects of diversity (negative) and resource level (positive) on the variability of both resource use and community composition. Our analyses of large-scale observational data are consistent with experimental and model studies demonstrating causal effects of microbial diversity on functional properties at the system level. Our findings point at potential linkages between eutrophication and pollution-mediated loss of phytoplankton diversity. Factors reducing phytoplankton diversity may have direct detrimental effects on the amount and predictability of aquatic primary production.

Climate change‐related heatwaves are major threats to biodiversity and ecosystem functioning. However, our current understanding of the mechanisms governing community resistance to and recovery from extreme temperature events is still rudimentary. The spatial insurance hypothesis postulates that diverse regional species pools can buffer ecosystem functioning against local disturbances through the immigration of better‐adapted taxa. Yet, experimental evidence for such predictions from multi‐trophic communities and pulse‐type disturbances, like heatwaves, is largely missing. We performed an experimental mesocosm study to test whether species dispersal from natural lakes prior to a simulated heatwave could increase the resistance and recovery of plankton communities. As the buffering effect of dispersal may differ among trophic groups, we independently manipulated the dispersal of organisms from lower (phytoplankton) and higher (zooplankton) trophic levels. The experimental heatwave suppressed total community biomass by having a strong negative effect on zooplankton biomass, probably due to a heat‐induced increase in metabolic costs, resulting in weaker top‐down control on phytoplankton. While zooplankton dispersal did not alleviate the negative heatwave effects on zooplankton biomass, phytoplankton dispersal enhanced biomass recovery at the level of primary producers, providing partial evidence for spatial insurance. The differential responses to dispersal may be linked to the much larger regional species pool of phytoplankton than of zooplankton. Our results suggest high recovery capacity of community biomass independent of dispersal. However, community composition and trophic structure remained altered due to the heatwave, implying longer‐lasting changes in ecosystem functioning. [ABSTRACT FROM AUTHOR]

Salinity is a major environmental predictor of phytoplankton species richness and composition. We hypothesize that the variation in phytoplankton richness along coastal salinity gradients is reflected in essential ecosystem functions like resource use efficiency (RUE)—the proportion of limiting resource that is converted into biomass. To test the hypothesis, we analyzed time series of phytoplankton and environmental variables from the Chesapeake Bay and the Baltic Sea. We analyzed the relationship among salinity, diversity, and RUE in bivariate and interaction association, and in structural equation model (SEM)—a form of path analysis to resolve multivariate relationships among interrelated variables. We concluded that the intrusion of marine water will lead to rapid change in species diversity on the fresh side. Despite diversity drop, individual community functions, like RUE, remain relatively resilient, reflecting functional redundancy. We propose that the salinity gradient also reflects trophic complexity, allowing stable resource use at reduced diversity. [ABSTRACT FROM AUTHOR]

Aim: Waterbirds are important dispersal vectors of multicellular organisms; however, no study to date has focused on their potential role in dispersing aquatic microbial communities. We explicitly studied endozoochory of prokaryotes and unicellular microeukaryotes by waterbirds using DNA metabarcoding. By directly comparing the dispersed set of organisms to the source pool of a natural metacommunity, we aimed at a realistic estimate of the importance of waterbird zoochory for natural microbial communities. Location: Temporary saline soda pans in Austria and Hungary. Taxon: Prokaryotes and unicellular microeukaryotes. Methods: In 2017 and 2018, water samples were collected from a network of 25 temporary ponds along with fresh droppings of five waterbird species including the dominant greylag goose (Anser anser). Prokaryotic and microeukaryotic communities were identified via 16S and 18S rRNA gene amplicon sequencing. After quality filtering of sequence reads, pro‐ and microeukaryotic amplicon sequence variant (ASV) compositions were compared between the aquatic and dropping samples, across years and waterbird species. Results: 28% of the dominant aquatic prokaryotic and 19% of the microeukaryotic ASVs were transported by A. anser. ASV richness was lower, but compositional variation was higher in A. anser droppings than in aquatic communities, probably resulting from stochastic pick‐up from multiple aquatic habitats. The composition of prokaryotic ASVs in bird droppings differed among the 2 years and reflected the actual aquatic communities. The dispersed set of microbes were largely similar among the waterbird species except for the planktivore filter‐feeder northern shoveler (Spatula clypeata), which dispersed more microeukaryotes than the other waterbirds. Main conclusions: Using an amplicon sequencing approach to characterize aquatic microorganisms in waterbird droppings and in the associated environment, our study provides strong evidence for endozoochory of natural communities. These results imply that waterbirds may be crucial in maintaining ecological connectivity between aquatic habitats at the level of microbial communities. [ABSTRACT FROM AUTHOR]

Omnivory is widespread in food webs, with an important stabilising effect. The strength of omnivorous trophic interactions may change considerably with changes in the local environment.Shallow temporary waters are often characterised by high levels of inorganic turbidity that may directly limit the food uptake of filter‐feeding organisms, but there is little evidence on how it might affect omnivorous species. Anostracans are key species of temporary waters and recent evidence suggests that these organisms are omnivorous consumers of both phyto‐ and zooplankton.Using Branchinecta orientalis as a model species, our aim was to test how turbidity affects the feeding of an omnivorous anostracan. To do this, we used short‐term feeding experiments and stable isotope analyses, with animals collected from soda pans in eastern Austria. In the feeding experiments, algae and zooplankton were offered as food either separately or in combination. The prey type treatments were crossed with turbidity levels in a factorial design.There was a pronounced decrease in the ingested algal biomass with increasing turbidity. Conversely, ingestion rates on zooplankton were less affected by turbidity. Stable isotope analyses from field material supported our experimental results by showing a positive relationship of the trophic position of anostracans and the trophic niche of the communities with turbidity.Our results show that turbidity modulates the intraguild trophic relationship between anostracans and their prey by shifting the diet of anostracans from more herbivorous in transparent to more carnivorous in turbid waters. Thus, inorganic turbidity might also have a community‐shaping role in plankton communities of temporary waters through altering trophic relationships. [ABSTRACT FROM AUTHOR]

Abstract: Species diversity is affected by processes operating at multiple spatial scales, although the most relevant scales that contribute to compositional variation and the temporal shifts of the involved mechanisms remain poorly explored. We studied spatial patterns of phytoplankton, rotifers and microcrustacean diversity across scales in a river floodplain system of the Danube in Austria under contrasting hydrological conditions (post‐flood versus low water level). The species turnover between water sections (β2) and between wetlands (β3) was the major components of regional diversity for all studied groups, with species turnover between habitats (β1) as a minor contributor. β1 diversity and β2 diversity were lower than expected by chance in most cases, suggesting that communities are more homogeneous than expected at these scales. β3 diversity was higher than expected by chance in many cases, indicating more distinct communities at the wetland level. Patterns were highly similar under different hydrological conditions, indicating no major immediate effect of flood events. Local environmental and spatial factors were similarly important in structuring phytoplankton, rotifer and microcrustacean communities in both hydrological conditions. Relevant environmental factors were spatially structured in post‐flood conditions especially between sections, suggesting flood‐driven homogenisation within the wetlands. Under low water level, spatial structuring of environment decreased and pure environmental factors gained relevance for phytoplankton and rotifers. Our results suggest that although β2 diversity between water sections is a major component of regional diversity, long‐term spatial processes responding to connectivity across the wetland structure phytoplankton, rotifer and microcrustacean communities. Aquatic sections within the limited spatial extent of the remaining floodplain areas appear more homogeneous than expected probably due to flood recurrence over the years. These results highlight that adequate planning of restoration and conservation strategies of floodplain wetlands should consider environmental heterogeneity together with long‐term spatial processes. [ABSTRACT FROM AUTHOR]

We used natural phytoplankton communities from four coastal regions to test diversity-functioning relationships, relations of N2-fixing cyanobacteria to nutrient imbalance, and the importance of metacommunity dynamics. Resource availability was measured as total nitrogen and phosphorus. Resource imbalance was determined as (1) the ratio of dissolved inorganic nitrogen to total phosphorus and (2) an experimentally verified indicator quantified by modelling responses in 14C-based primary production to nutrient additions. Resource availability explained variance in biomass, productivity, and species richness, as expected by the Species Energy Theory, but not evenness. Linear mixed-effects models confirmed the overall relation between productivity and resource availability, whereas other resource availability relations showed also notable clustering by region. The Resource Ratio Theory predicting that diversity increases with the number of limiting resources was not supported. Nutrient imbalance had a weak effect on the biomass of N2-fixing cyanobacteria, but not their share of total phytoplankton biomass. Contrary to many previous studies on biodiversity-ecosystem functioning relationships, we found a highly significant inverse relationship between evenness and biomass. This indicates that species-rich natural phytoplankton communities form the basis for opportunistic species to temporarily monopolize resources and create blooms. The result was consistent across regions, although their community composition differed. Metacommunity dynamics were important, since distance between the regions explained higher percentage of the variability than local resources together. As the species able to monopolize resources vary widely in their role for aquatic food webs and environmental consequences (e.g., food quality, toxicity), species-level trait data is essential to understand better diversity-productivity relationships. [ABSTRACT FROM AUTHOR]

The diversity-stability debate is a long-standing issue in ecology, asking whether more diverse communities show higher stability over time and more rapid recovery from disturbances. Connection to undisturbed habitats is thought to affect compositional and functional stability after disturbances. Therefore, we established marine phytoplankton metacommunities consisting of three microcosms (local patches), which were connected by tubes opened for different time intervals to create 5 levels of connectivity. We performed two experiments differing by homogeneous (HOM) or heterogeneous (HET) supply of irradiance across patches. As disturbance we either removed 75% of the algal biomass locally from one randomly chosen patch, or 25 or 75% regionally from each local patch. By comparing these treatments to an undisturbed control, we analyzed resilience (rate of recovery) and final recovery (recovery ratio) with regard to biomass (functional stability) and species composition (structural stability). In both experiments (HET, HOM), functional and structural aspects of stability responded significantly to connectivity and disturbance treatments. Functional resilience was enhanced by increasing connectivity (HET and HOM), which partially also increased functional recovery (HET) and structural resilience (HOM). By contrast, the treatment-induced gradients in diversity (species richness and evenness) had no clear effect on functional resilience or recovery. Instead, structural and functional resilience were strongly correlated regardless of patch quality, indicating that only a full recovery in community composition ensured functional stability. Our findings suggest that connectivity plays a pivotal role in maintaining ecosystem stability under pulse disturbance such that a more complete understanding of stability requires spatially explicit approaches. [ABSTRACT FROM AUTHOR]

Analysing and interpreting long-term phytoplankton time series present a number of challenges, arising from potential historical inconsistencies in data collection and taxonomic identification of organisms. In a previous paper, Pomati et al. (2012) found a remarkable increase in phytoplankton diversity that coincided with oligotrophication and warming of Lake Zurich over a 32-year period. These findings were recently challenged on the basis of potential biases in detection limits and taxonomic classification over the time series (Straile, Jochimsen & Kümmerlin, 2013). We agree that being cautious with long-term phytoplankton data series is extremely important, but argue that the increase in richness detected in Lake Zurich cannot be due only to methodological bias., Following additional analysis of the Lake Zurich phytoplankton dataset, we found that the shift in taxon detection limits reported by Straile et al. (2013) is not supported by the data and stems from a rounding error in the calculation of density in the dataset available to those authors. We found a decline in the proportional abundance for common taxa, an increase in the annual prevalence of taxa and reduced community turnover over the time series. Taken together, the data clearly indicate a trend of decreasing dominance, while more taxa coexist simultaneously. We also argue that the taxonomic classification has been robust (at least at the family level) and propose a diagnostic plot that can help detect an unbiased signal of change in plankton richness through time., Straile et al. (2013) observed perfect synchrony in species occurrence between Lake Zurich and nearby Lake Walen. While we agree that such perfect synchrony in community composition can reflect a bias in the database compilation, it can also be an important ecological signal of changes in regional species pools that deserves further analysis., We conclude that the results of Pomati et al. (2012) are robust and not substantially undermined by the criticisms of Straile et al. (2013). More generally, it is indeed possible to extract meaningful signals of biodiversity change from long-term phytoplankton monitoring datasets, provided there is a clear understanding of how the data have been sampled, recorded and analysed over the history of the time series. [ABSTRACT FROM AUTHOR]

In their seminal paper, Goldman et al. suggested that phytoplankton close to maximum growth rate attains a restricted optimal N : P ratio close to the Redfield ratio of molar N : P = 16. Recently, the presence of such a global attractor for optimal phytoplankton stoichiometry has been questioned in models and empirical analyses. As the chemical composition of phytoplankton is of major importance for our understanding of global elemental cycles and biogeochemical transformations, we assembled 55 data sets of phytoplankton growth rate and biomass N : P ratios in a meta-analysis testing (1) whether phytoplankton N : P converges at high growth rates, (2) whether N : P ratios scale with growth rate, and (3) whether the optimal N : P ratios achieved at highest growth rates reflect organism traits or environmental conditions. Across systems and species, phytoplankton N : P decreased with increasing growth rate and at the same time showed decreasing variance, i.e., fast-growing phytoplankton is more P rich and has a more confined elemental composition. Optimal N : P increased with increasing N : P of available nutrients, i.e., with increasing P limitation. Other differences were rare, except cyanobacteria showed higher optimal N : P than diatoms. Understanding the role of phytoplankton in biogeochemical transformation requires modeling approaches that are stoichiometrically flexible to reflect the dynamics of growth and nutrient supply in primary producers. [ABSTRACT FROM AUTHOR]

Microbial parasites have only recently been included in planktonic food web studies, but their functional role in conveying dietary energy still remains to be elucidated. Parasitic fungi (chytrids) infecting phytoplankton may constitute an alternative trophic link and promote organic matter transfer through the production of dissemination zoospores. Particularly, during proliferation of inedible or toxic algal species, such as large Cyanobacteria fostered by global warming, parasites can constitute an alternative trophic link providing essential dietary nutrients that support somatic growth and reproduction of consumers. Using phytoplankton-parasites associations grown under laboratory controlled conditions we assessed the fatty acids and biochemical composition of species with different nutritional quality and followed the metabolic pathway from the algal host and their parasites zoospores using compound-specific stable isotope analysis. This study demonstrated that chytrids are trophic upgraders able to retain essential nutrients that can be transferred to upper trophic levels both in terms of organic matter quantity and nutritional quality. Through the production of zoospores, nutritionally important long-chain polyunsaturated fatty acids that can be consequently assimilated by consumers. We conclude that parasitism at the base of aquatic food webs may represent a crucial trophic link for dietary nutrients and essential biomolecules alternative to herbivory or bacterivory, which can be particularly crucial during the proliferation of inedible or nutritionally inadequate algal species fostered by climate change. [ABSTRACT FROM AUTHOR]

Cell size determination is a critical part of sampling phytoplankton communities, as size is both a fundamental trait determining species' ecological niches and a quantity necessary for generating unbiased estimates of community composition. The sizes of algal cells span orders of magnitude, driving variation in growth parameters, sinking rates, herbivore defense, and biogeochemical roles. Size variation occurs at both intra- and interspecific levels. Many central questions in community and ecosystem ecology require estimating and comparing the relative abundances of species. Cell counts provide insufficient estimates, due to interspecific variation in cell size, as they over-estimate the importance of abundant, small species. While cell size (or biovolume) estimation has become standard practice in limnology, many older data sets lack this information, limiting their interpretation and utility. To address these challenges, and advance the study of freshwater phytoplankton communities, we have compiled a database of >260000 algal biovolume estimates, spanning ~400 genera and ~1200 species. This represents the most extensive, standardized, and taxonomically diverse data set of its kind, drawing on both publicly and privately held sources. Estimates of the sizes of colonies or filaments are included, where applicable. With these data researchers will be able to account for intra- and interspecific variation in cell size, enabling new and improved analyses of classic data sets. [ABSTRACT FROM AUTHOR]

Biodiversity losses in marine, terrestrial, and freshwater ecosystems have raised concerns about the maintenance of sustainable ecosystem functions and services ("biodiversity crisis"). A positive diversity--productivity relationship has previously been supported by theoretical models, and by laboratory and field experiments in a variety of ecosystems including unicellular microbial communities. Here we show an increasing biomass yield of aquatic primary producers at the ecosystem scale, paralleled by a long- term positive biodiversity change, which contrasts with the trend of global biodiversity loss. The implied direct long-term biodiversity effect on ecosystem functioning was an increase of phytoplankton biomass per unit limiting nutrient by a factor of 1.2 to 1.4. Changes in diversity of microorganisms may have immediate implications for essential ecosystem processes like productivity and biomass yield. Diversity-driven enhancement of resource use in primary production can lead to increased food web yields, but they also can cause a stoichiometric mismatch between autotrophs and primary consumers. Unveiling the functional roles of planktonic biodiversity therefore has essential implications both for global change and for harvestable marine resources. [ABSTRACT FROM AUTHOR]