Recommendations for sampling littoral diatoms in lakes for ecological status assessments

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.     

Coherence of phytoplankton and attached diatom-based ecological status assessment in Lake Balaton

Coherence between ecological status assessment by phytoplankton and attached diatoms was analyzed in the littoral zone of Lake Balaton. Sampling of periphytic diatoms, phytoplankton, and water were carried out at ten different littoral sites in the northern and southern shores of the lake for a year. Phytoplankton species were sorted into functional groups and ecological status was assessed by means of the phytoplankton assemblage Q index. The index TDIL was calculated using quantitative attached diatom data. Significant differences were found between the ecological assessments based on phytoplankton and phytobenthos metrics, both seasonally and spatially. The Q index indicated ecological states varying from bad to good, while the average of diatom indices varied from moderate to high conditions. The Q index provided more realistic ecological status of Lake Balaton, compared with trophic status based on TP values, especially in the summer period. Differences in the response-time indication of phytoplankton and attached diatoms suggest that lack of coherence should also be expected between the responses of other BQEs.     

Nitrogen limitation of the phytobenthos in Alpine lakes: results from nutrient‐diffusing substrata

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Assessment of ecological status in U.K. rivers using diatoms

1. The European Union's Water Framework Directive requires all water bodies to achieve ‘good ecological status’ by 2015 and this paper describes a rationale for defining ‘good ecological status’ based on diatoms, a significant component of the biological quality element ‘macrophyte and phytobenthos’. 2. A database of benthic diatom samples collected over the past 20 years was assembled. New sampling, specifically for this project, was undertaken during 2004 to supplement these data. In total 1051 samples were included in the database with matching environmental data. 3. ‘Reference sites’, relatively unimpacted by human activity, were selected from this database by a series of screening steps and these sites were used to develop a site‐specific reference typology. 4. Environmental variables not related to the pressure gradient were used to predict the ‘expected’ Trophic Diatom Index (TDI) values at each site. Site‐specific TDI predictions were used to generate ecological quality ratios (EQRs) ranging from ≥1, where the diatom assemblage showed no impact, to (theoretically) 0, when the diatom assemblage was indicative of major anthropogenic activities. 5. The boundary between ‘high’ and ‘good’ status was defined as the 25th percentile of EQRs of all reference sites. The boundary between ‘good’ and ‘moderate’ status was set at the point at which nutrient‐sensitive and nutrient‐tolerant taxa were present in equal relative abundance. An ecological rationale for this threshold is outlined in the paper.     

Diatoms in a peatbog polluted with oil-field brines in the Kolva River basin,Komi Republic

The first data about the diatoms of phytobenthos and epiphyton in water biotopes of a peatbog in the Kolva River basin (a tributary of the Pechora River) polluted by oil-field brines are reported. As a total, 49 species with varieties and forms from 14 genera and 9 families have been found. Dominating complexes of communities with halophilic diatoms have been revealed. The ecology of the species resistant to mineral pollution is discussed.     

龙感湖沉积物泥芯硅藻群落和营养状态的反演

The importance of aquatic vegetation to the ecologi-cal restoration has been recognized commonly bylimnolo-gists and lake managements[1—4].As to the ecologicalrestoration in eutrophicated lakes,it is of great signifi-cance to knowthe dynamic process of the ecosystemevo-lution in a macrophyte-dominated lake under the humanimpacts in historical period,to make it clear whether thecommunitystructure and ecological function would be af-fected bythe extension of the vegetation growth,tofind asolutionto remain ste...     

An assessment of the reproductive biology of the Marico barb Barbus motebensis from the upper Groot Marico catchment,South Africa

Temporal changes in fatty acid composition and δ¹⁵N, δ¹³C stable isotope values of the phytobenthos growing on artificial clay substrates under natural conditions over a 28-day period at an upstream and a downstream site in the Kowie River near Grahamstown were investigated in 2012. High concentrations of diatom markers 16:1ω7 and 20:5ω3 fatty acids were recorded, especially at the downstream site, reflecting the importance of diatoms in contributing to the phytobenthos communities at that station. After day 7 at the downstream site the average δ¹⁵N value of the phytobenthos was lighter, gradually increasing by ～2‰ and ～5‰ overall to heavier values on day 28. At the upstream site there were no significant changes (<1‰ increase) in δ¹⁵N values of the phytobenthos over time. Stable nitrogen (δ¹⁵N) and carbon (δ¹³C) signatures in the phytobenthos communities were significantly different between sites (one-way ANOVA; p < 0.001). The stable isotope values and fatty acid concentrations of phytobenthos at the downstream site were different to those of the phytobenthos at the upstream site, and they changed concurrently with changes in the phytobenthos community structure. At the downstream site there was a strong correlation of the δ¹⁵N of phytobenthos with nitrates (R = 0.56) and time (weeks; R = 0.81). However, the fatty acids were not specific enough to characterise the composition of phytobenthos communities. Other biomarker methods, such as stable isotopes and microscopic examination of the communities, were found to be useful. The results from this relatively small-scale tile experiment indicate the complexity of changes in fatty acid composition and δ¹⁵N, δ¹³C stable isotope values of a phytobenthos community. Stable isotope and fatty acid composition can be successfully used to map changes in phytobenthos composition and carbon and nitrogen flow patterns along a river continuum.     

Multi-proxy studies in palaeolimnology

Multi-proxy studies are becoming increasingly common in palaeolimnology. Eight basic requirements and challenges for a multi-proxy study are outlined in this essay – definition of research questions, leadership, site selection and coring, data storage, chronology, presentation of results, numerical tools and data interpretation. The nature of proxy data is discussed in terms of physical proxies and biotic proxies. Loss-on-ignition changes and the use of transfer functions are reviewed as examples of problems in the interpretation of data from multi-proxy studies. The importance of pollen analysis and plant macrofossil analysis in multi-proxy studies is emphasised as lake history cannot be interpreted without knowledge of catchment history. Future directions are outlined about how multi-proxy studies can contribute to understanding biotic responses to environmental change.     

Effect of streamlining taxa lists on diatom-based indices: implications for intercalibrating ecological status

Intercalibration of ecological status class boundaries between member states is a requirement of the European Union’s Water Framework Directive. Although a preliminary intercalibration of boundaries established for phytobenthos has been performed, a number of questions remain, including the extent to which variations in taxonomic concepts used in different member states influences the position of these boundaries. In this paper, the robustness of the diatom-based metrics used for intercalibration is assessed. Whilst use of genus-level identification led to a loss of ecological information, merging representatives of closely-related taxa has little effect on these metrics. Similarly, taxa that occur only rarely or never have high relative abundances in a dataset can also be ignored without the loss of ecological information. Similar results were obtained when modern taxonomic concepts were compared with concepts in use 80 years ago. Fine scale taxonomy may play a valuable role within member states; however, our results suggest that, at a continental scale, a simplified approach to diatom taxonomy should not affect intercalibration results.     

Taxonomy‐free molecular diatom index for high‐throughput eDNA biomonitoring

Current biodiversity assessment and biomonitoring are largely based on the morphological identification of selected bioindicator taxa. Recently, several attempts have been made to use eDNA metabarcoding as an alternative tool. However, until now, most applied metabarcoding studies have been based on the taxonomic assignment of sequences that provides reference to morphospecies ecology. Usually, only a small portion of metabarcoding data can be used due to a limited reference database and a lack of phylogenetic resolution. Here, we investigate the possibility to overcome these limitations using a taxonomy‐free approach that allows the computing of a molecular index directly from eDNA data without any reference to morphotaxonomy. As a case study, we use the benthic diatoms index, commonly used for monitoring the biological quality of rivers and streams. We analysed 87 epilithic samples from Swiss rivers, the ecological status of which was established based on the microscopic identification of diatom species. We compared the diatom index derived from eDNA data obtained with or without taxonomic assignment. Our taxonomy‐free approach yields promising results by providing a correct assessment for 77% of examined sites. The main advantage of this method is that almost 95% of OTUs could be used for index calculation, compared to 35% in the case of the taxonomic assignment approach. Its main limitations are under‐sampling and the need to calibrate the index based on the microscopic assessment of diatoms communities. However, once calibrated, the taxonomy‐free molecular index can be easily standardized and applied in routine biomonitoring, as a complementary tool allowing fast and cost‐effective assessment of the biological quality of watercourses.     

Using diatom life-forms and ecological guilds to assess organic pollution and trophic level in rivers: a case study of rivers in south-eastern France

The European Union’s Water Framework Directive has set a target of achieving good ecological status for all aquatic environments in Europe by 2015. In order to determine the quality of aquatic environments, biological indicators such as diatoms are often used. However, biotic diatom indices can be difficult and time consuming to use because of complexity of species determination. We investigated whether the biological traits of diatoms in rivers (life-forms, size classes and ecological guilds) could be used to assess organic pollution and trophic level. We worked on a data set comprising 315 diatom species, determined at 328 river stations of south-east France and a variety of parameters. The abundances of some biological traits differed significantly between the different organic pollution and trophic levels, particularly stalked diatoms, and the motile and low-profile guilds.     

The use of periphytic diatoms as a means of assessing impacts of point source inorganic nutrient pollution in south-eastern Australia

1. Periphytic diatoms are used as indicators of water quality because their ecological tolerances or preferences to environmental variables are thought to be predictable. However, much of the present autecological information for periphytic diatoms has been derived from studies conducted in the northern hemisphere. In this present study we used periphytic diatoms to determine the impacts of inorganic nutrient pollution in a tidal river system in the temperate latitudes of south‐east Australia. In so doing, we assess the suitability of the use of the ‘northern hemisphere’ ecological tolerance/preference data for periphytic diatoms. 2. Artificial substrates were used to collect periphytic diatoms at 35 sites, which were positioned along the riverbanks and the middle of the river at various distances upstream and downstream of the sewage outfall. The sampling design took into account tidal excursions and the observed sewage plume dynamics. Periphytic diatoms were collected during the austral winter month of August and the austral spring months of September and October. We deployed the artificial substrates for 4 weeks to allow the periphytic diatoms to recruit and colonise, before identifying and enumerating the assemblages. 3. Data analysis included two approaches: multivariate visualisations of combinations of environmental and biological data to investigate shifts in species structure of the periphytic diatom assemblage and multimetric indices based on ecological tolerance/preference data. 4. We found that the spatial patterns inferred from multivariate and multimetric analyses were consistent. Temporal variation in the composition of the periphytic diatom assemblage was greater than the spatial variation along horizontal sections of the river (in any one deployment) due mainly to shifts between winter and spring species. 5. Outfall effects were most apparent in winter, possibly because subsequent deployments were swamped by growth of spring periphytic diatoms. The outfall effects included a shift towards pollutant tolerant species and a reduction in the variability of the periphytic diatom assemblage across the river. 6. We conclude that the use of periphytic diatoms and associated ecological tolerance/preference data as a means of assessing impacts of point source inorganic nutrient pollution is effective. An understanding of river and sewage flow patterns is essential to the design of appropriate monitoring programmes and to the interpretation of results, especially as periphytic diatoms are sensitive to many environmental variables.     

大理西湖流域开发历史与硅藻群落变化的模式识别

有关云南湖泊的研究长期集中于高原九大湖泊和水体富营养化评价,缺少对中小型水体及多重环境压力胁迫的综合研究.本文以大理西湖为例,结合沉积物记录与现代监测资料,甄别了气候变化和人类活动干扰下硅藻群落结构的长期响应模式及其驱动强度.结果表明: 20世纪50年代以前,大理西湖总体处于自然演化阶段;1950年代开始,围湖造田和流域改造的增强导致了水体营养水平增加、水动力条件改变,硅藻优势种由扁圆卵型藻替代为脆杆藻属;而1997年以来营养水平的快速增加和湖泊水动力的改变,促进了浮游藻类大量生长、底栖硅藻持续减少,同时水生植物快速退化、生态系统稳定性明显降低.因此,在长期流域开发的背景下,对云南中小型高山湖泊的有效保护需要评价流域开发类型、强度及全球变暖的长期影响.     

Factors controlling the distribution of planktonic foraminifera in the Red Sea and implications for the development of transfer functions

The Red Sea is an extreme marine environment, with conditions limiting the application of standard geochemical proxies for the reconstruction of paleoclimate. In order to develop paleoenvironmental reconstruction methods which are not dependent on chemical signals, we investigated the distribution of planktonic foraminifera in the surface sediments and assessed the viability of constructing foraminiferal transfer functions in this basin. We find a distinct gradient in the faunal assemblage along the basin's axis, which is reflected in a high correlation between faunal composition and all considered environmental parameters (temperature, salinity, chlorophyll a concentration, stratification, and oxycline depth). As a result, transfer functions constructed by different methods (ANN, MAT, IKM, WA-PLS) appear to be able to estimate all of these parameters with a high average accuracy (15% of the parameter's range in the Red Sea). However, redundancy analysis of the distribution of foraminiferal assemblages in surface sediments alone did not yield unambiguous results in terms of which of the considered factors exerts a primary control on the foraminifera distribution and which of the observed relationships are the result of the mutual correlation among the environmental factors. To disentangle the effect of individual environmental parameters, we applied the obtained transfer functions on a newly generated Holocene record from the central Red Sea. The integration of published paleoclimate reconstructions with our data allowed us to identify productivity as the most likely primary control of the planktonic foraminifera distribution in the Red Sea. The generated transfer functions can estimate paleoproductivity with acceptable accuracy (RMSEP chlorophyll a = 0.1 mg/m³; ~ 8% of recent range), but only under such conditions in the past when circulation patterns and salinity levels in the basin were fundamentally comparable to the present day. Since productivity in the central and southern Red Sea is closely linked with the Monsoon-driven water exchange across the Strait of Bab al Mandab, the resulting reconstructions can provide indirect information on the mode and intensity of the monsoonal system in the past.     

Ecological functions of volatile organic compounds in aquatic systems

In terrestrial ecosystems, volatile organic compounds (VOCs) are widely acknowledged as an important group of infochemicals. They play a major role in pollinator attraction by terrestrial plants and as insect pheromones. Furthermore, they are the mediating agent of so-called 'tritrophic interactions'. When plants are attacked by herbivorous insects, volatile signal substances are emitted, which act as attractants for parasitoids that kill the herbivores, thereby protecting the plant from herbivory. Despite the generally acknowledged importance of VOCs in terrestrial chemical ecology, their functions in aquatic food webs are largely unknown. VOCs produced by algae and cyanobacteria are a major concern in water processing, since aquatic primary producers are the reason for regularly encountered taste and odour problems in drinking water. Only very recently, research in aquatic chemical ecology has started to investigate possible ecological functions for the production of VOCs by algae and cyanobacteria. Volatile aldehydes released by wounded cells of marine planktonic diatoms seem to act as defensive compounds against herbivorous copepods on the population level. Just recently, it was found that VOCs released from benthic algae and cyanobacteria can be utilised as food and/or habitat finding cues by aquatic invertebrates such as freshwater gastropods and nematodes. Here, I review concepts and recent experimental studies on the ecological functions of such VOCs in aquatic ecosystems. Understanding the factors that lead to the liberation of volatile compounds is an essential prerequisite to properly assessing their ecological functions. It appears that (similar to terrestrial plant-herbivore interactions) VOCs can also play a steering role for both attraction and defence in aquatic ecosystems.     

Evolutionary origins and functions of the carotenoid biosynthetic pathway in marine diatoms

Carotenoids are produced by all photosynthetic organisms, where they play essential roles in light harvesting and photoprotection. The carotenoid biosynthetic pathway of diatoms is largely unstudied, but is of particular interest because these organisms have a very different evolutionary history with respect to the Plantae and are thought to be derived from an ancient secondary endosymbiosis between heterotrophic and autotrophic eukaryotes. Furthermore, diatoms have an additional xanthophyll-based cycle for dissipating excess light energy with respect to green algae and higher plants. To explore the origins and functions of the carotenoid pathway in diatoms we searched for genes encoding pathway components in the recently completed genome sequences of two marine diatoms. Consistent with the supplemental xanthophyll cycle in diatoms, we found more copies of the genes encoding violaxanthin de-epoxidase (VDE) and zeaxanthin epoxidase (ZEP) enzymes compared with other photosynthetic eukaryotes. However, the similarity of these enzymes with those of higher plants indicates that they had very probably diversified before the secondary endosymbiosis had occurred, implying that VDE and ZEP represent early eukaryotic innovations in the Plantae. Consequently, the diatom chromist lineage likely obtained all paralogues of ZEP and VDE genes during the process of secondary endosymbiosis by gene transfer from the nucleus of the algal endosymbiont to the host nucleus. Furthermore, the presence of a ZEP gene in Tetrahymena thermophila provides the first evidence for a secondary plastid gene encoded in a heterotrophic ciliate, providing support for the chromalveolate hypothesis. Protein domain structures and expression analyses in the pennate diatom Phaeodactylum tricornutum indicate diverse roles for the different ZEP and VDE isoforms and demonstrate that they are differentially regulated by light. These studies therefore reveal the ancient origins of several components of the carotenoid biosynthesis pathway in photosynthetic eukaryotes and provide information about how they have diversified and acquired new functions in the diatoms.     

Microphytobenthos as a biogeomorphological force in intertidal sediment stabilization

The properties and behavior of intertidal marine sediments cannot be understood without taking their biology into account. Biological factors are important for the stability and erosion threshold of intertidal sediments as well as for sediment transport. In this paper I focus on intertidal sediments that are colonized and dominated by phototrophic microorganisms and their impact on the morphodynamics and sediment stabilization. The emphasis is on epipelic diatoms. These organisms exude copious amounts of extracellular polymeric substances (EPS) that may contribute to the stability of the sediment by gluing and binding. I review the factors that lead to the development of such microphytobenthic communities and the processes that lead to the exudation of EPS and its fate in intertidal mudflats. Epipelic diatoms exude EPS partly as the result of unbalanced growth. Extraction of EPS from cultures of epipelic diatoms yields two operational fractions. While one fraction contains largely neutral EPS, which may serve as a carbon- and energy reserve for the organism, the other is acidic and more recalcitrant to degradation. The latter EPS fraction is therefore predominant in the muddy sediment and may be responsible for increasing the erosion threshold. However, since extracted EPS alone is incapable of increasing the erosion threshold, diatoms are apparently actively involved in the structuring of the biofilm matrix. Therefore, sediment stabilization cannot be attributed simply to EPS alone.