European river typologies fail to capture diatom, fish, and macrophyte community composition.

Typology systems are frequently used in applied and fundamental ecology and are relevant for environmental monitoring and conservation. They aggregate ecosystems into discrete types based on biotic and abiotic variables, assuming that ecosystems of the same type are more alike than ecosystems of different types with regard to a specific property of interest. We evaluated whether this assumption is met by the Broad River Types (BRT), a recently proposed European river typology system, that classifies river segments based on abiotic variables, when it is used to group biological communities. We compiled data on the community composition of diatoms, fishes, and aquatic macrophytes throughout Europe and evaluated whether the composition is more similar in site groups with the same river type than in site groups of different river types using analysis of similarities, classification strength, typical species analysis, and the area under zeta diversity decline curves. We compared the performance of the BRT with those of four region-based typology systems, namely, Illies Freshwater Ecoregions, the Biogeographic Regions, the Freshwater Ecoregions of the World, and the Environmental Zones, as well as spatial autocorrelation (SA) classifications. All typology systems received low scores from most evaluation methods, relative to predefined thresholds and the SA classifications. The BRT often scored lowest of all typology systems. Within each typology system, community composition overlapped considerably between site groups defined by the types of the systems. The overlap tended to be the lowest for fishes and between Illies Freshwater Ecoregions. In conclusion, we found that existing broad-scale river typology systems fail to delineate site groups with distinct and compositionally homogeneous communities of diatoms, fishes, and macrophytes. A way to improve the fit between typology systems and biological communities might be to combine segment-based and region-based typology systems to simultaneously account for local environmental variation and historical distribution patterns, thus potentially improving the utility of broad-scale typology systems for freshwater biota.

On the shape and origins of the freshwater species-area relationship.

The species-area relationship (SAR) has over a 150-year-long history in ecology, but how its shape and origins vary across scales and organisms remains incompletely understood. This is the first subcontinental freshwater study to examine both these properties of the SAR in a spatially explicit way across major organismal groups (diatoms, insects, and fish) that differ in body size and dispersal capacity. First, to describe the SAR shape, we evaluated the fit of three commonly used models, logarithmic, power, and Michaelis-Menten. Second, we proposed a hierarchical framework to explain the variability in the SAR shape, captured by the parameters of the SAR model. According to this framework, scale and species group were the top predictors of the SAR shape, climatic factors (heterogeneity and median conditions) represented the second predictor level, and metacommunity properties (intraspecific spatial aggregation, γ-diversity, and species abundance distribution) the third predictor level. We calculated the SAR as a sample-based rarefaction curve using 60 streams within landscape windows (scales) in the United States, ranging from 160,000 to 6,760,000 km2 . First, we found that all models provided good fits (R2 ≥ 0.93), but the frequency of the best-fitting model was strongly dependent on organism, scale, and metacommunity properties. The Michaelis-Menten model was most common in fish, at the largest scales, and at the highest levels of intraspecific spatial aggregation. The power model was most frequent in diatoms and insects, at smaller scales, and in metacommunities with the lowest evenness. The logarithmic model fit best exclusively at the smallest scales and in species-poor metacommunities, primarily fish. Second, we tested our framework with the parameters of the most broadly used SAR model, the log-log form of the power model, using a structural equation model. This model supported our framework and revealed that the SAR slope was best predicted by scale- and organism-dependent metacommunity properties, particularly spatial aggregation, whereas the intercept responded most strongly to species group and γ-diversity. Future research should investigate from the perspective of our framework how shifts in metacommunity properties due to climate change may alter the SAR.

Wind Exposure Regulates Water Oxygenation in Densely Vegetated Shallow Lakes.

The presence of dense macrophyte canopies in shallow lakes locally generates thermal stratification and the buildup of labile organic matter, which in turn stimulate the biological oxygen demand. The occurrence of hypoxic conditions may, however, be buffered by strong wind episodes, which favor water mixing and reoxygenation. The present study aims at explicitly linking the wind action and water oxygenation within dense hydrophytes stands in shallow lakes. For this purpose, seasonal 24 h-cycle campaigns were carried out for dissolved gases and inorganic compounds measurements in vegetated stands of an oligo-mesotrophic shallow lake. Further, seasonal campaigns were carried out in a eutrophic shallow lake, at wind-sheltered and -exposed sites. Overall results showed that dissolved oxygen (DO) daily and seasonal patterns were greatly affected by the degree of wind exposure. The occurrence of frequent wind episodes favored the near-bottom water mixing, and likely facilitated mechanical oxygen supply from the atmosphere or from the pelagic zone, even during the maximum standing crop of plants (i.e., summer and autumn). A simple model linking wind exposure (Keddy Index) and water oxygenation allowed us to produce an output management map, which geographically identified wind-sheltered sites as the most subjected to critical periods of hypoxia.

Corrigendum: Invasive Aquatic Plants as Ecosystem Engineers in an Oligo-Mesotrophic Shallow Lake.

[This corrects the article DOI: 10.3389/fpls.2018.01781.].

Metabolome response to anthropogenic contamination on microalgae: a review.

BACKGROUND: Microalgae play a key role in ecosystems and are widely used in ecological status assessment. Research focusing on such organisms is then well developed and essential. Anyway, approaches for a better comprehension of their metabolome's response towards anthropogenic stressors are only emerging. AIM OF REVIEW: This review presents the biochemical responses of various microalgae species towards several contaminants including metals and chemicals as pesticides or industrial compounds. We aim to provide a comprehensive and up-to-date overview of analytical approaches deciphering anthropogenic contaminants impact on microalgae metabolome dynamics, in order to bring out relevant biochemical markers that could be used for risk assessment. KEY SCIENTIFIC CONCEPTS OF REVIEW: Studies to date on ecotoxicological metabolomics on microalgae are highly heterogeneous in both analytical techniques and resulting metabolite identification. There is a real need for studies using complementary approaches to determine biomarkers usable for ecological risk assessment.

Invasive Aquatic Plants as Ecosystem Engineers in an Oligo-Mesotrophic Shallow Lake.

Exotic hydrophytes are often considered as aquatic weeds, especially when forming dense mats on an originally poorly colonized environment. While management efforts and research are focused on the control and on the impacts of aquatic weeds on biodiversity, their influence on shallow lakes' biogeochemical cycles is still unwell explored. The aim of the present study is to understand whether invasive aquatic plants may affect the biogeochemistry of shallow lakes and act as ecosystem engineers. We performed a multi-year investigation (2013-2015) of dissolved biogeochemical parameters in an oligo-mesotrophic shallow lake of south-west of France (Lacanau Lake), where wind-sheltered bays are colonized by dense mats of exotic Egeria densa Planch. and Lagarosiphon major (Ridl.) Moss. We collected seasonal samples at densely vegetated and plant-free areas, in order to extrapolate and quantify the role of the presence of invasive plants on the biogeochemistry, at the macrophyte stand scale and at the lake scale. Results revealed that elevated plant biomass triggers oxygen (O2), dissolved inorganic carbon (DIC) and nitrogen (DIN) stratification, with hypoxia events frequently occurring at the bottom of the water column. Within plants bed, elevated respiration rates generated important amounts of carbon dioxide (CO2), methane (CH4) and ammonium (NH4 +). The balance between benthic nutrients regeneration and fixation into biomass results strictly connected to the seasonal lifecycle of the plants. Indeed, during summer, DIC and DIN regenerated from the sediment are quickly fixed into plant biomass and sustain elevated growth rates. On the opposite, in spring and autumn, bacterial and plant respiration overcome nutrients fixation, resulting in an excess of nutrients in the water and in the increase of carbon emission toward the atmosphere. Our study suggests that aquatic weeds may perform as ecosystem engineers, by negatively affecting local oxygenation and by stimulating nutrients regeneration.

Biogeographical Patterns of Species Richness and Abundance Distribution in Stream Diatoms Are Driven by Climate and Water Chemistry.

In this intercontinental study of stream diatoms, we asked three important but still unresolved ecological questions: (1) What factors drive the biogeography of species richness and species abundance distribution (SAD)? (2) Are climate-related hypotheses, which have dominated the research on the latitudinal and altitudinal diversity gradients, adequate in explaining spatial biotic variability? and (3) Is the SAD response to the environment independent of richness? We tested a number of climatic theories and hypotheses (i.e., the species-energy theory, the metabolic theory, the energy variability hypothesis, and the climatic tolerance hypothesis) but found no support for any of these concepts, as the relationships of richness with explanatory variables were nonexistent, weak, or unexpected. Instead, we demonstrated that diatom richness and SAD evenness generally increased with temperature seasonality and at mid- to high total phosphorus concentrations. The spatial patterns of diatom richness and the SAD-mainly longitudinal in the United States but latitudinal in Finland-were defined primarily by the covariance of climate and water chemistry with space. The SAD was not entirely controlled by richness, emphasizing its utility for ecological research. Thus, we found support for the operation of both climate and water chemistry mechanisms in structuring diatom communities, which underscores their complex response to the environment and the necessity for novel predictive frameworks.

Assessing anthropogenic pressures on streams: A random forest approach based on benthic diatom communities.

Benthic diatoms have been widely used to assess the ecological status of freshwater ecosystems, especially in the context of recent international water framework directive policies (e.g. the WFD). Despite diatom-based indices are known to respond fastly to water quality degradation, they are not designed to precisely identify the nature of pressures co-occurring in the environment. Based on large scale monitoring data, we aimed at building models able to estimate the risk of stream impairment by many types of anthropogenic pressures from taxonomy-based and trait-based characteristics of diatom assemblages. Random forest models were built to individually evaluate the impairment risk of diatom assemblages for six chemical and five hydromorphological or land-use related pressure categories. Eight models provided good impairment risk assessment (Area Under the Curve≥0.70). Under multi-pressure scenarios, models built for chemical pressures exhibited a better accuracy than hydromorphological or land-use related ones. Models were able to detect both ecological restoration and degradation, based on long-term surveys. These models have been implemented in a R user-friendly routine, to help stream managers to early identify degrading processes and prioritize management actions.

Diatom Cooccurrence Shows Less Segregation than Predicted from Niche Modeling.

Species cooccurrence patterns give significant insights into the processes shaping communities. While biotic interactions have been widely studied using cooccurrence analyses in animals and larger plants, studies about cooccurrences among micro-organisms are still relatively rare. We examined stream diatom cooccurrences in France through a national database of samples. In order to test the relative influence of environmental, biotic and spatial constraints on species' incidence distribution, cooccurrence and nestedness patterns of real communities were compared with the patterns generated from a set of standard and environmentally constrained null models. Real communities showed a higher level of segregation than the most conservative standard null models, but a general aggregation of cooccurrences when compared to environmentally constrained null models. We did not find any evidence of limiting similarity between cooccurring species. Aggregations of species cooccurrences were associated with the high levels of nestedness. Altogether, these results suggested that biotic interactions were not structuring cooccurrences of diatom species at our study scale. Instead, the patterns were more likely to be related with colonization patterns, mass effect, and local temporal dynamics of diatom biofilms. We further highlight that the association of standard and environmentally constrained null models may give realistic insight into the cooccurrence patterns of microbial communities.

A multicompartment approach--diatoms, macrophytes, benthic macroinvertebrates and fish--to assess the impact of toxic industrial releases on a small French river.

The River Luzou flows through a sandy substrate in the South West of France. According to the results of two assessment surveys, the Water Agency appraised that this river may not achieve the good ecological status by 2015 as required by the Water Framework Directive (2000/60/EC). This ecosystem is impacted by industrial effluents (organic matter, metals and aromatic compounds). In order to assess and characterize the impact, this study aimed to combine a set of taxonomic and non-taxonomic metrics for diatoms, macrophytes, macroinvertebrates and fish along the up- to downstream gradient of the river. Diversity metrics, biological indices, biological and ecological traits were determined for the four biological quality elements (BQE). Various quantitative metrics (biomass estimates) were also calculated for diatom communities. The results were compared to physicochemical analysis. Biological measurements were more informative than physicochemical analysis, in the context of the study. Biological responses indicated both the contamination of water and its intensity. Diversity metrics and biological indices strongly decreased with pollution for all BQE but diatoms. Convergent trait selection with pollution was observed among BQE: reproduction, colonization strategies, or trophic regime were clearly modified at impaired sites. Taxon size and relation to the substrate diverged among biological compartments. Multiple anthropogenic pollution calls for alternate assessment methods of rivers' health. Our study exemplifies the fact that, in the case of complex contaminations, biological indicators can be more informative for environmental risk, than a wide screening of contaminants by chemical analysis alone. The combination of diverse biological compartments provided a refined diagnostic about the nature (general mode of action) and intensity of the contamination.

Water quality assessment of rivers using diatom metrics across Mediterranean Europe: a methods intercalibration exercise.

The European Water Framework Directive establishes a framework for the protection of water resources. However, common water management tools demand common understanding of assessment methods, so quality goals are equally met. Intercalibration of methods ensures the comparability of biological elements across similar geographical areas. Many aspects can influence the outcome of intercalibration: data sampling, treatment methods, taxonomic reliability of databases, choice of metrics for ecological quality status classification, and criteria for selecting reference sites. This study describes the potentials and constraints of the intercalibration of indices using diatoms for assessment of Mediterranean rivers. Harmonisation of diatom taxonomy and nomenclature was based on a previous ring test which took place at the European level. Four diatom indices (Indice de Polluosensibilité Spécifique-IPS, Indice Biologique Diatomées-IBD 2007, Intercalibration Common Metric Italy-ICMi and Slovenian Ecological Status assessment system) were intercalibrated using data from six European Mediterranean countries (Cyprus, France, Italy, Portugal, Slovenia and Spain). Boundaries between High/Good and Good/Moderate quality classes were harmonised by means of the Intercalibration Common Metric (ICM). Comparability between countries was assured through boundary bias and class agreement. The national boundaries were adjusted when they deviated more than a quarter of a class equivalent (0.25) from the global mean. All national methods correlated well with the ICM, which was sensitive to water quality (negatively correlated to nutrients). Achnanthidium minutissimum sensu lato was the most discriminative species of Good ecological status class. Planothidium frequentissimum, Gomphonema parvulum and Nitzschia palea were the most contributive to Moderate ecological status class. Some taxa were discriminative for both Good and Moderate ecological status classes due to low indication and ecological discriminative power but also due to differences in taxonomy between countries. This intercalibration exercise allowed establishment of common water quality goals across Mediterranean Europe, which is substantiated with the ICM.

Assessing the impact of chloroacetanilide herbicides and their metabolites on periphyton in the Leyre River (SW France) via short term growth inhibition tests on autochthonous diatoms.

The Leyre River is the main tributary to the Bassin d'Arcachon lagoon. Herbicides belonging to the chloroacetanilide class have been found in the river (S-metolachlor and acetochlor) as well as some of their metabolites at higher concentrations. As the environmental toxicity of these molecules is not well known, ecotoxicological tests have been carried out on river periphyton at different levels of biological diversity: from the clone of one diatom species (Nitzschia nana) to the population of the same species (several clones) up to the multi-specific species community dominated by diatoms. Moreover, tests were performed on diatoms coming from an unpolluted upstream site and from a contaminated downstream site, in order to investigate possible tolerance acquisition to pollutants. The method consisted in measuring diatom growth inhibition at different doses of each substance from the increase of chlorophyll-a concentration after 4 days. It resulted that acetochlor was clearly more toxic than S-metolachlor at all levels of biological diversity. EC(50) values estimated from the tests suggest no effect of contaminants on diatom growth or biomass in the river. The toxicity of the metabolites appeared very low compared to that of their parent compounds. No difference in tolerance to the herbicides was demonstrated between summer diatom communities from the two sites in spite of different specific compositions. However, concerning the populations of N. nana isolated in winter following the highest herbicide concentrations in the river (about 0.5 µg L(-1)), the downstream population showed a higher tolerance to acetochlor but there was no co-tolerance to S-metolachlor. Thus, it appeared that acetochlor represents the highest toxic pressure on periphyton among the other contaminants in the Leyre River.