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Climate change threatens surface waters worldwide, especially shallow lakes where one of the expected consequences is a sharp increase in their water temperatures. Phytobenthos is an essential, but still less studied component of aquatic ecosystems, and it would be important to learn more about how global warming will affect this community in shallow lakes. In this research, the effects of different climate change scenarios (SSP2-4.5 and SSP5-8.5, as intermediate and high emission scenarios) on the structure and function of the entire phytobenthos community using species- and trait-based approaches were experimentally investigated in an outdoor mesocosm system. Our results show that the forecasted 3 °C increase in temperature will already exert significant impacts on the benthic algal community by (1) altering its species and (2) trait composition (smaller cell size, lower abundance of colonial and higher of filamentous forms); (3) decreasing Shannon diversity; and (4) enhancing the variability of the community. Higher increase in the temperature (+5 °C) will imply more drastic alterations in freshwater phytobenthos by (1) inducing very high variability in species composition and compositional changes even in phylum level (towards higher abundance of Cyanobacteria and Chlorophyta at the expense of Bacillariophyta); (2) continuing shift in trait composition (benefits for smaller cell volume, filamentous life-forms, non-motile and weakly attached taxa); (3) further reducing the functional diversity; (4) increasing biofilm thickness (1.4 μm/°C) and (5) decreasing maximum quantum yield of photosystem II. In conclusion, already the intermediate emission scenario will predictably induce high risk in biodiversity issues, the high emission scenario will imply drastic impacts on the benthic algae endangering even the function of the ecosystem., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier Inc. All rights reserved.)

Since 2000, after the Water Framework Directive came into force, aquatic ecosystems' bioassessment has acquired immense practical importance for water management. Currently, due to extensive scientific research and monitoring, we have gathered comprehensive hydrobiological databases. The amount of available data increases with each subsequent year of monitoring, and the efficient analysis of these data requires the use of proper mathematical tools. Our study challenges the comparison of the modelling potential between four indices for the ecological status assessment of lakes based on three groups of aquatic organisms, i.e. phytoplankton, phytobenthos and macrophytes. One of the deep learning techniques, artificial neural networks, has been used to predict values of four biological indices based on the limited set of the physicochemical parameters of water. All analyses were conducted separately for lakes with various stratification regimes as they function differently. The best modelling quality in terms of high values of coefficients of determination and low values of the normalised root mean square error was obtained for chlorophyll a followed by phytoplankton multimetric. A lower degree of fit was obtained in the networks for macrophyte index, and the poorest model quality was obtained for phytobenthos index. For all indices, modelling quality for non-stratified lakes was higher than this for stratified lakes, giving a higher percentage of variance explained by the networks and lower values of errors. Sensitivity analysis showed that among physicochemical parameters, water transparency (Secchi disk reading) exhibits the strongest relationship with the ecological status of lakes derived by phytoplankton and macrophytes. At the same time, all input variables indicated a negligible impact on phytobenthos index. In this way, different explanations of the relationship between biological and trophic variables were revealed.

Although the Water Framework Directive specifies that macrophytes and phytobenthos should be used for the ecological assessment of lakes and rivers, practice varies widely throughout the EU. Most countries have separate methods for macrophytes and phytobenthos in rivers; however, the situation is very different for lakes. Here, 16 countries do not have dedicated phytobenthos methods, some include filamentous algae within macrophyte survey methods whilst others use diatoms as proxies for phytobenthos. The most widely-cited justification for not having a dedicated phytobenthos method is redundancy, i.e. that macrophyte and phytoplankton assessments alone are sufficient to detect nutrient impacts. Evidence from those European Union Member States that have dedicated phytobenthos methods supports this for high level overviews of lake condition and classification; however, there are a number of situations where phytobenthos may contribute valuable information for the management of lakes., (Copyright © 2016 The Authors. Published by Elsevier B.V. All rights reserved.)

Eutrophication impairs lake ecosystems at a global scale. In this context, as benthic microalgae are well-established warnings for a large range of stressors, particularly nutrient enrichment, the Water Framework Directive required the development of diatom-based methods to monitor lake eutrophication. Here, we present the diatom-based index we developed for French lakes, named IBDL (Indice Biologique Diatomées en Lacs). Data were collected in 93 lakes from 2015 to 2020. A challenge arose from the discontinuous pressure gradient of our dataset, especially the low number of nutrient-impacted lakes. To analyze the data we opted for the so-called "Threshold Indicator Taxa ANalysis" method, which makes it possible to determine a list of "alert taxa." We obtained a multimetric index based on specific pressure gradients (Kjeldahl nitrogen, suspended matter, biological oxygen demand, and total phosphorous). Considering the European intercalibration process, the very good correlation between IBDL and the common metric (R2 from 0.52 to 0.87 according to the lake alkalinity type) makes us very confident in our ability to match future IBDL quality thresholds with European standards. The IBDL proved at last to be particularly relevant as it has a twofold interest: an excellent relationship with total phosphorus (R2 from 0.63 to 0.83 according to the lake alkalinity type) and a possible application to any lake metatype. Its complementarity with macrophyte-based indices moreover justifies the use of at least two primary producer components for lake ecological status classification. [ABSTRACT FROM AUTHOR]

Mediterranean lakes present particular challenges when developing ecological assessment techniques, due partly to long histories of human activity in their catchments but also to strong interactions with climate, intensifying in recent decades due to global warming. A survey of diatoms in lakes in Greece revealed both a strong nutrient gradient and a salinity gradient. It is necessary to disentangle these when assessing ecological status in order that appropriate measures for restoring good ecological status are applied. Two approaches were used to develop a Water Framework Directive-compatible metric for assessing ecological status using diatoms. The strongest relationship was obtained using weighted averaging, but an alternative metric, generated by rescaling a well-established and widely used diatom metric performed only slightly less well. The limited number of lakes in Greece resulted in a small dataset, creating a risk of overfitting models and the rescaled metric has the advantage of drawing on a broader autecological knowledge base. Supplementary metrics to evaluate salinity are also proposed in order to ensure that lakes subject to this pressure are clearly differentiated from those where nutrient impacts predominate. Approaches to defining reference conditions and status class boundaries draw on established criteria developed for Water Framework Assessments in Europe to ensure that status assessments in Greece reflect similar levels of ambition to those established elsewhere in the European Union.

Diatom communities are composed of three distinct water-depth-associated assemblages: a nearshore benthic assemblage, a deep-water benthic assemblage, and a primarily planktonic assemblage. Light attenuation with depth has been hypothesized to be the factor that controls benthic-to-planktonic (B:P) assemblage transitions. Previous studies, however, could not distinguish between the individual effects of light and temperature on diatom assemblage transition because the depth of the B:P transition occurred at or near the depth of the thermocline. This has caused confusion regarding which factor is the more influential variable on benthic-planktonic diatom assemblage transition in surface sediments along water-depth transects in lakes. Two boreal lakes in northeast Ontario were selected to examine this problem. The study lakes are sufficiently deep and clear (Secchi disk depths ~ 9–10 m; dissolved organic carbon ~ 3 mg/L), making it possible to distinguish between the depth of 1% light penetration and the depth of the thermocline. Surface sediment samples were collected along water-depth transects from these lakes to examine intra-lake diatom assemblage structure. In both lakes, the water depth of the diatom assemblage B:P transition was closely associated with the depth of the ~ 1% photosynthetically active radiation value, and was well below the depth of the thermocline, suggesting that light penetration has more influence on this transition than does the thermocline. Because temporal fluxes in nutrients and DOC concentrations can attenuate light, understanding how the light regime influences benthic diatoms in lakes can help inform interpretations of diatom assemblage changes in paleolimnological studies. [ABSTRACT FROM AUTHOR]

To test if phytobenthic algae provide additional important information to macrophytes and phytoplankton for lake monitoring, we sampled two large lakes in Norway. In each lake, we analyzed water chemistry and phytoplankton above the deepest site, recorded macrophytes and non-diatom phytobenthic algae at 20 sites around the shoreline and estimated site-specific nutrient input from land cover. Since no ready-to-use phytobenthos index exists for lakes in Norway, we tested the PIT index developed for rivers, commonly perceived signs of disturbance such as high algal cover, and taxon richness as well as similarity patterns. Both lakes were nutrient poor, but had potential local nutrient inputs (villages, agriculture). In neither of the lakes did phytobenthos indicate a worse overall ecological status than macrophytes and phytoplankton. Our data therefore, did not suggest that it would be useful to add phytobenthos into surveillance monitoring of lakes in Norway. There was a loose correlation between macrophyte and phytobenthic site-specific taxon richness and similarities. This means that macrophytes and phytobenthos do indeed give partly redundant information. High algal cover was found at sites with both high and low phosphorus input. Using algal cover as indicator of site-specific nutrient input is therefore overly simplistic. Urban and cultivated areas were associated with a more eutrophic PIT. This indicates that the PIT, despite being developed for lotic waters, may be used to detect site specific nutrient input in lakes. [ABSTRACT FROM AUTHOR]

• A new diatom-based metric for ecological assessment in Mediterranean lakes. • Rescaling algorithm to optimise existing metric for local conditions. • Need to assess nutrients and salinity simultaneously in Mediterranean lakes. • Draw on Water Framework Directive knowledge base to set reference conditions and class boundaries. Mediterranean lakes present particular challenges when developing ecological assessment techniques, due partly to long histories of human activity in their catchments but also to strong interactions with climate, intensifying in recent decades due to global warming. A survey of diatoms in lakes in Greece revealed both a strong nutrient gradient and a salinity gradient. It is necessary to disentangle these when assessing ecological status in order that appropriate measures for restoring good ecological status are applied. Two approaches were used to develop a Water Framework Directive-compatible metric for assessing ecological status using diatoms. The strongest relationship was obtained using weighted averaging, but an alternative metric, generated by rescaling a well-established and widely used diatom metric performed only slightly less well. The limited number of lakes in Greece resulted in a small dataset, creating a risk of overfitting models and the rescaled metric has the advantage of drawing on a broader autecological knowledge base. Supplementary metrics to evaluate salinity are also proposed in order to ensure that lakes subject to this pressure are clearly differentiated from those where nutrient impacts predominate. Approaches to defining reference conditions and status class boundaries draw on established criteria developed for Water Framework Assessments in Europe to ensure that status assessments in Greece reflect similar levels of ambition to those established elsewhere in the European Union. [ABSTRACT FROM AUTHOR]

Eleven European countries participated in an exercise to harmonise diatom-based methods used for status assessment in lakes. Lakes were divided into low, medium and high alkalinity types for this exercise. However, it was not possible to perform a full intercalibration on low alkalinity lakes due to the short gradient and confounding factors. Values of the Trophie Index were computed for all samples in order that national datasets could all be expressed on a common scale. Not all participants had reference sites against which national methods could be standardised and, therefore, a Generalised Linear Modelling approach was used to control the effect of national differences in datasets. This enabled the high/good and good/moderate status boundaries to be expressed on a common scale and for deviations beyond ±0.25 class widths to be identified. Those countries which had relaxed boundaries were required to adjust these to within ±0.25 class widths whilst the intercalibration rules allowed those countries with more stringent boundaries to retain these. Despite biogeographical and typological differences between countries, there was broad agreement on the characteristics of high, good and moderate status diatom assemblages, and the exercise has ensured consistent application of Water Framework Directive assessments around Europe. [ABSTRACT FROM AUTHOR]

The European Union's Water Framework Directive (WFD) requires that all water bodies in Europe achieve good ecological status (GES) by 2015. We developed an ecological classification tool for UK lakes based on benthic diatoms, a key component of the biological-quality element macrophytes and phytobenthos. A database of 1079 epilithic and epiphytic diatom samples and matching environmental data was assembled from 228 UK lakes. The data set was divided into 3 lake types: low, medium, and high alkalinity. A lake trophic diatom index (LTDI) was developed based on modification of the trophic diatom index (TDI) for rivers, and ecological quality ratios (EQRs) were generated for each lake type. The high/good status boundary was defined as the 25th percentile of EQRs of all reference sites (identified based on independent sedimentary-diatomassemblage data or catchment point-source and landuse data), whereas the good/moderate boundary was set at the point at which nutrient-sensitive and nutrient-tolerant taxa were present in equal relative abundance. The moderate/poor and poor/bad boundaries were defined by equal division of the remaining EQR gradient. Samples from reference sites were used to predict the expected LTDI value for each sample, and these values were compared with the classifications derived from the LTDI. For lakes identified as reference sites, 68% were classified as having high status and 32% as having good. The model predicted 81% of nonreference lakes to have good or worse status. The model was applied to 17 English lakes (10 low- and 7 medium-alkalinity) for which classification based on other WFD tools was available. The classifications based on LTDI gave the same status (within 1 class) as other biological elements for 11 of the 17 lakes (65%). Thus, the LTDI gives a reliable assessment of the condition of the littoral biofilm and is a key component of a WFD-compliant tool kit for classifying UK standing waters. [ABSTRACT FROM AUTHOR]

The Water Framework Directive (WFD) provides the legal basis for water management in the European Union (EU). Twelve years after it was passed, all but five EU Member States had phytobenthos assessment methods for rivers, whilst nine had methods for lakes. Most are based on diatoms, although a few are supplemented by evaluations of non-diatoms and some include macroalgae as part of parallel macrophyte assessments. Norway is the exception, with assessment based on non-diatom algae alone. Over half of all states have methods based wholly or partly on weighted average metrics developed before the onset of the WFD, with nine choosing the Indice de Polluosensibilité Specifique. Such metrics generally have high correlations with the predominant nutrient or organic pollution gradient and, as such, represent pragmatic solutions to ecological status assessment. However, their widespread use raises questions about what, exactly, ‘ecological status’ means. Strong relationships with chemical pressure gradients may be a mixed blessing as pressure gradients are often composed of several intercorrelated variables, making it difficult to disentangle correlation and causation in the absence of ecophysiological studies of individual species. Moreover, the focus on strong relationships with chemical gradients means that most phytobenthos metrics describe the scale of hazard at a site rather than the risk posed to other trophic levels and to ecosystem services. This first generation of phytobenthos assessment tools may be inadequate when catchment managers need guidance on remediation strategies for particular water bodies. A second generation of assessment tools, focused on the fitness of the phytobenthos as part of aquatic ecosystems, rather than just as indicators of chemical conditions, is needed if the goal of good ecological status around Europe is to be achieved. [ABSTRACT FROM PUBLISHER]

Biological elements, including phytoplankton, phytobenthos, macrophytes, benthic invertebrates and fish, are employed by the EU Water Framework Directive (WFD) 2000/60/EC as ecological indicators for the assessment of surface waters. The use of primary producers (phytoplankton, phytobenthos and macrophytes) for water quality assessment has a long history, and several methods have been developed worldwide. In this study, we used these three communities to assess the ecological status of five natural lakes located in the Aquitaine region (southwest France). Several biological indices used in lakes from other European countries or in French rivers were employed and compared among the three communities. Each primary producer provided complementary information about the ecological status of the lakes, including the invasiveness of exotic taxa. Regardless of the producer community used, the response to the environment, as reflected by the indices (adequate for each community), was similar: Lakes Cazaux, Lacanau and Hourtin showed the best ecological status and Parentis and Soustons the worst. Phytoplankton diagnosis reflected and integrated unambiguously the water quality of the lakes, as demonstrated by the strong relationships between the phytoplankton index and the trophic status criteria. This community appeared as the best indicator, especially when macrophytes were absent. The methods applied here represent a potential tool for the assessment of the ecological status in the context of WFD, but they need to be refined. We propose modifications for phytobenthos index initially tailored for running waters for adequate use in lentic ecosystems. Indices for the three primary producers should be modified to incorporate exotic species which may provide information on potential biodiversity losses. [ABSTRACT FROM AUTHOR]

Abstract: The St. Lucia estuarine lake on the north coast of KwaZulu-Natal, South Africa, is one of the largest estuarine systems in Africa and of unique importance for the adjacent marine and terrestrial ecosystems. The area regularly experiences periods of drought, resulting in hypersaline conditions in its shallow lakes and the closure of the estuarine mouth. This study aimed to assess the primary production rates of phytoplankton and microphytobenthos throughout an annual cycle of this drought phase. Primary production rates were assessed at representative sites, namely the Mouth, Narrows, South and North Lakes from June 2006 to May 2007. Because of the drought, the salinity gradient from the mouth to the head of the estuary was reversed by comparison to estuarine systems with a steady freshwater inflow and regular marine exchange. In March 2007, during the study, the mouth opened as a result of rough seas, and the marine influence broke the existing reversed gradient, producing a marine salinity throughout the system. Microphytobenthic primary productivity varied between 0 and 34 mg C m−2 h−1 and showed strong correlations with salinity, DIN:DIP ratios and irradiance. Benthic productivity was high across the system after breaching of the mouth. Pelagic primary productivity (between 0 and 180 mg C m−2 h−1), showed a correlation with temperature and irradiance and was highest across the system in February 2007 when the mouth was still closed. There was no significant correlation between production rates and biomass (chl-a) in either the benthic or pelagic habitats. The negative correlation between DIN:DIP ratio and benthic primary productivity indicated that phosphorus was the limiting nutrient. This study shows that salinity, along with seasonally dependent parameters such as temperature and irradiance, correlates with the rate of microalgal production. Hence, in these shallow lakes, the largest primary productivity can occur in either the pelagic or benthic subsystems, depending on prevailing conditions at the time. [Copyright &y& Elsevier]

Light environment, community structure, pigments, and photophysiological properties of mat-forming phytobenthos were studied in four shallow Antarctic lakes in 2007 at maximum water depths of 1.7–2.5 m. All lakes were oligotrophic, and water transparencies were high, enabling 45–60% of photosynthetically active radiation (PAR, 400–700 nm) and 20–40% of ultraviolet radiation (300–400 nm) to reach the lake beds. Phytobenthic mats were dominated by cyanobacteria and green algae. Little PARL (500–700 nm) penetrated through the firm mat in the shallowest lake, while in the other lakes more (>20%) PARL got through the mats to the subsurface mat layers. Photochemical activities indicated almost no photoinhibition but low photosynthetic efficiency in all mat surface layers. Non-photochemical quenching was rarely detected, suggesting excess energy dissipation may not be efficient in the UV-rich environment. There was a positive correlation between photo-protective substances and incident radiation in the mats, and an inverse correlation between such substances and photochemical efficiency, suggesting that the phytobenthos survive by changing a light-protection/utilization balance. The communities under strong UV-B and PAR had firm mat textures and were characterized by high UV/photo-protective substance ratios that make them less transparent. Maximum relative electron transportation rates (rETRmax) and photochemical efficiencies, however, were low, possibly because the protective substances prevent efficient light usage. In contrast, communities under mild light were characterized by lower substance ratios and softer textures, while rETRmax values and photochemical efficiencies were greater. The phytobenthic mat surface seems to act as a filter for strong and harmful light, typically penetrating through the clear water of Antarctic lakes, and produces a milder light environment for the subsurface mat organisms. [ABSTRACT FROM AUTHOR]

Research to develop tools to assess the ecological status of phytobenthos, as required in Annex V of the European Union Water Framework Directive, has focussed largely on diatoms. Diatoms are often the most abundant and diverse group of algae within the phytobenthos and have been used widely for other monitoring purposes. However, there is little empirical justification for the use of diatoms as proxies for the wider phytobenthos. In this paper, we re-examine an existing dataset compiled largely from littoral samples from standing waters in the English Lake District and compared transfer functions for total phosphorus, dissolved inorganic carbon, conductivity and calcium concentration generated from diatoms and non-diatoms separately and together. The results show that transfer functions generated from diatoms alone are as powerful as transfer functions generated from diatoms and non-diatoms combined, while transfer functions generated from non-diatoms alone are less effective. These results provide support for the use of diatoms as proxies for phytobenthos when ecological status is being assessed. [ABSTRACT FROM AUTHOR]

1. Spatial and temporal variations in algal concentrations are controlled in many aquatic ecosystems by the availability of solar irradiance, rather that nutrients or grazing. In such light limiting conditions, changes in the optical or hydrological characteristics of the water column will directly impact biomass concentrations. Here we develop and test an approach based on the relationship between available solar irradiance within the mixed layer and algal biomass concentrations. 2. As with most nutrient/biomass relationships, an increase in available solar energy favours an increase in biomass when light limitation prevails. The ratio between light/biomass is then used to determine a critical light requirement that can be used to estimate critical depth and compensation irradiance and open the way to exploring how changes in mixing depth and vertical attenuation may influence algal biomass concentrations. 3. This approach is used to describe real conditions in two disparate algal communities; the phytoplankton community in Lake Victoria, East Africa and the microphytobenthos community in the lacustrine system of Esteros del Iberá (South America). 4. Differences in the critical light requirement were used to examine the relative efficiency of the algal communities in their use of available solar energy. The tropical phytoplankton community showed similar energetic requirements to theoretical estimates and were found to be less efficient when compared with the phytobenthos community. [ABSTRACT FROM AUTHOR]

This review summarises the existing literature and outlines the theoretical basis for using standard methods for sampling diatoms from rivers to sample littoral diatoms and other phytobenthos from standing waters. The European Union's Water Framework Directive has created a statutory obligation for EU Member States to monitor macrophytes and phytobenthos in lakes. Although there has been a considerable amount of work using phytobenthos (especially diatoms) to monitor river water quality in Europe, there are fewer studies on the use of phytobenthos for monitoring in lakes. European standard methods for sampling diatoms from rivers should be suitable, with only minor modifications, for sampling littoral diatoms from lakes and other standing water bodies. These recommendations should be applicable to all temperate regions. [ABSTRACT FROM AUTHOR]

Abstract: A new assessment system for macrophytes and phytobenthos in German lakes according to the Water Framework Directive of the European Community is described. Based on biological, chemical and hydromorphological data from about 100 lake sites covering the main ecoregions, hydromorphological lake types and degradation forms, biocoenotic types could be defined. For developing a classification system the quality element macrophytes and phytobenthos was divided into two components: macrophytes and benthic diatoms. For macrophytes 4 and for benthic diatoms 4 lake types were identified. The benthic vegetation at reference conditions is described and degradation is characterised as deviation in benthic vegetation species composition and abundance from the reference biocoenosis. For classification in five ecological status classes, several metrics were developed and used in combination with existing indices. For a few of the described lake types further investigations are necessary before a classification can be developed. [Copyright &y& Elsevier]

• Phytobenthos in lowland lakes in Romania does not demonstrate a strong response to phosphorus. • Assemblage composition is, instead, determined by biochemical oxygen demand and conductivity. • A new metric to evaluate ecological status using phytobenthos in these lakes is described. • Expected values of this metric are derived from a quantile regression of the index against conductivity. • As conductivity is susceptible to climate change, 2010 values are used as benchmarks when calculating expected index values. In contrast to diatom assemblages in lakes in most other parts of Europe, those in lowland lakes in Romania appear to be determined primarily by biochemical oxygen demand (BOD) and conductivity rather than by nutrients. This has confounded the development of a Water Framework Directive-compatible phytobenthos assessment system for Romanian lakes and led, instead, to the development of a new metric, the Romanian Diatom Index (RDI), which captures these gradients. The primary anthropogenic stressor is assumed to be BOD whilst conductivity is a product of background geology and climate. The RDI is strongly correlated with BOD, and ecological quality ratios (EQRs) are computed by dividing the observed RDI by an expected value, determined from the 90th percentile of the relationship between RDI and conductivity. This equates to the "best available" RDI at any point on the conductivity gradient and, in the absence of true "reference sites", has been assumed to represent the boundary between high and good status. The boundary between good and moderate status has been set at the point where the sum of valves of taxa tolerant to elevated BOD exceeds the sum of valves of taxa that are sensitive to elevated BOD. The position of this boundary has been further validated by the use of a Threshold Indicator Species Analysis (TITAN). In view of the susceptibility of conductivity to climate warming, 2010 values of conductivity have been used to ensure a stable benchmark against which future changes can be measured. This study highlights the challenges involved in performing reliable ecological assessments in situations that are, by the standards of much of Europe, atypical. [ABSTRACT FROM AUTHOR]