Should only live diatoms be used in the bioassessment of small mountain streams?

It is unclear whether differentiating live and dead diatoms would enhance the accuracy and precision of diatom-based stream bioassessment. We collected benthic diatom samples from 25 stream sites in the Northern Oregon Coast ecoregion. We counted live diatoms (cells with visible chloroplasts) and then compared the counts with those generated using the conventional method (clean counts). Non-metric multidimensional scaling (NMDS) showed that the diatom assemblages generated from the two counts were overall similar. The relationships between the two diatom assemblages (summarized as NMDS ordination axes) and the environmental variables were also similar. Both assemblages correlated well with in-stream physical habitat conditions (e.g., channel dimensions, substrate types, and canopy cover). The conventional diatom method provides taxonomic confidence while the live diatom count offers ecological reliability. Both methods can be used in bioassessment based on specific assessment objectives. Handling editor: J. Saros     

Using Diatom Assemblages to Assess Urban Stream Conditions

We characterized changes in diatom assemblages along an urban-to-rural gradient to assess impacts of urbanization on stream conditions. Diatoms, water chemistry, and physical variables of riffles at 19 urban and 28 rural stream sites were sampled and assessed during the summer base flow period. Near stream land use was characterized using GIS. In addition, one urban and one rural site were sampled monthly throughout a year to assess temporal variation of diatom assemblages between the urban and rural stream sites. Canonical correspondence analysis (CCA) showed that the 1st ordination axis distinctly separated rural and urban sites. This axis was correlated with conductivity (r = 0.75) and % near-stream commercial/industrial land use (r = 0.55). TWINSPAN classified all sites into four groups based on diatom assemblages. These diatom-based site groups were significantly different in water chemistry (e.g., conductivity, dissolved nutrients), physical habitat (e.g., % stream substrate as fines), and near-stream land use. CCA on the temporal diatom data set showed that diatom assemblages had high seasonal variation along the 2nd axis in both urban and rural sites, however, rural and urban sites were well separated along the 1st ordination axis. Our results suggest that changes in diatom assemblages respond to urban impacts on stream conditions.     

Comparative Environmental Assessment in the Studies of Benthic Diatom,Macroinvertebrate, and Fish Communities

Community response to environmental gradients operating at hierarchical scales was assessed in studies of benthic diatoms, macroinvertebrates and fish from 44 stream sites in the New York City watershed. Hierarchical cluster analysis (TWINSPAN) of diatoms and fish partitioned the study sites into four groups, i.e., acid streams, reservoir outlets and wetland streams, large eutrophic streams, and small eutrophic streams; macroinvertebrate TWINSPAN distinguished an additional group of silty eutrophic streams. The correspondence among the three assemblage TWINSPAN groupings was moderate, ranging from 51 to 57%. The similarity across the four major group types was the highest among large eutrophic stream and acid stream assemblages, and the lowest among small eutrophic stream assemblages. Stepwise discriminant function analysis revealed that environmental factors discriminated most effectively the diatom grouping and least effectively the fish grouping. The best environmental predictors for diatom and macroinvertebrate grouping were conductance and percent surface water, while population density was most powerful in separating the fish groups. Carbaryl was the only pesticide that correlated with macroinvertebrate grouping. Partial redundancy analyses suggested a differential dependence of freshwater communities on the scale of the environmental factors to which they respond. The role of small‐scale habitat and habitatland cover/land use interaction steadily increased across the diatom, macroinvertebrate, and fish assemblages, whereas the effect of large‐scale land cover/land use declined.     

Regional distribution of diatom assemblages in the headwater streams of Luxembourg

The aim of this study was to determine the influence of environmental variables on the structure of benthic diatom assemblages, and to propose type assemblages of diatoms characterizing unpolluted headwater streams of Luxembourg. A total of 289 diatom samples were collected in the headwater streams of Luxembourg. At each sampling site, physical and chemical variables were also measured. The relationships between environmental variables and the distribution of the taxa were estimated using univariate and multivariate statistical analyses. On the basis of geology and stream water chemistry, Luxembourg could be separated in two regions. The streams in the schistose northern part of Luxembourg presented a low carbonate hardness (median 3.5° F); the streams in the southern part of Luxembourg presented a high carbonate hardness (median 21.8° F) notably connected to the presence of sandstone and limestone substrata. A Twinspan classification carried out on diatom assemblages defined two groups of samples corresponding to these two regions. The carbonate hardness that is related to the nature of the geological substratum appeared to be the major structuring variable for the assemblage composition. Anthropogenic pollution was a secondary structuring variable for diatom assemblages since each group could be subdivided in subgroups presenting statistically different nutrient and organic matter concentrations. The diatom assemblages of these subgroups were characterised by differences of saprobic and trophic preferences. On the basis of these results, two type assemblages of diatoms are proposed for the unpolluted headwater streams in the two regions of Luxembourg. The presence of these different type assemblages in Luxembourg shows the necessity to adapt diatom bioindication to the different regions of the country.     

Relationships Between Environmental Variables and Benthic Diatom Assemblages in California Central Valley Streams (USA)

This study examines distributional patterns of benthic diatom assemblages in relation to environmental characteristics in streams and rivers in the California Central Valley ecoregion. Benthic diatoms, water quality, and physical habitat conditions were characterized from 53 randomly selected sites. The stream sites were characterized by low mid-channel canopy cover and high channel substrate embeddedness. The waters at these sites were enriched with minerals and turbidity varied from 1.3 to 185.0 NTU with an average of 13.5 NTU. A total of 249 diatom taxa were identified. Average taxa richness was 41 with a range of 7–76. The assemblages were dominated by Staurosira construens (11%), Epithemia sorex (8%), Cocconeis placentula (7%), and Nitzschia amphibia (6%). Multivariate analyses (cluster analysis, classification tree analysis, and canonical correspondence analysis) all showed that benthic diatom assemblages were mainly affected by channel morphology, in-stream habitat, and riparian conditions. The 1st CCA axis negatively correlated with mean wetted channel width (r = −0.66) and thalweg depth (r = −0.65) (Table 4). The 2nd axis correlated with % coarse substrates (r=0.60). Our results suggest that benthic diatoms can be used for assessing physical habitat alterations in streams.     

Distribution of Epilithic Diatoms in Response to Environmental Conditions in an Urban Tropical Stream, Central Kenya

Use of diatoms in monitoring water quality is well acknowledged in developed countries, but only recently has the assessment started gaining importance in developing countries. Diatoms can be obtained from natural and artificial substrates. Appreciating the differences and similarities of diatom assemblages on both substrates may contribute to a better understanding and standardization particularly during monitoring of water quality. During this study we assessed diatom assemblages, biodiversity and trophic indices in relation to water quality along the Nairobi River. Fifteen sites were sampled in September 2000 during the dry season. Diatoms were collected from natural substrates (stones, pebbles) and artificial substrates (100% acrylic wool). On artificial and natural substrates, a total of 190 and 151 taxa were found, respectively, the majority of these taxa (80%) have cosmopolitan distribution and are also widespread throughout tropical African. Species composition changed downstream, five taxa dominated upper and mid stream sites whereas lower stream sites were dominated by one or two taxa. Species richness, diversity, dominance and evenness were positively correlated with NO₃, O₂ and altitude but decreased markedly downstream with a simultaneous increase in total dissolved solids, alkalinity, chemical oxygen demand and PO₄. Ordination and classification (CANOCO and TWINSPAN) showed that diatom assemblages in the Nairobi River responded strongly to water quality changes with respect to concentrations of NO₃, NO₂, total dissolved solids and temperature. Taxa common at less impacted upstream sites included Gomphonema gracilis, Anomoeoneis brachysira and Fragilaria biceps; while common taxa at midstream sites with agricultural catchments were Gomphonema parvulum, Navicula cryptocephala, N. schroeteri, N. bryophila, N. halophila, Nitzschia linearis var. linearis and Cymbella silesica. Achnanthes minutissima var. saprophila, Gomphonema angustum, Navicula subminuscula, N. arvensis, Nitzschia palea and N. umbonata were most common at urban sites, which were polluted by residential and industrial effluents. Trophic diatom indices suggested that water quality was poor at most sites in the Nairobi River. Most sites along the river had low Generic Diatom Index values, GDI (<12) and high Trophic Diatom Index values, TDI 73–78 (median = 76) and 75–84 (median = 77) for artificial and natural substrates, respectively. This study showed that diatoms' response on natural and artificial substrates were similar and reflected environmental conditions correctly.     

Macroinvertebrate community response to natural and forest harvest gradients in western Oregon headwater streams

1. To examine the effects of forest harvest practices on headwater stream macroinvertebrates, we compiled a 167 site database with macroinvertebrate, fish, physical habitat and catchment land cover data from the three forested ecoregions in western Oregon. For our analysis, headwater streams were defined by catchment areas <10 km² and perennial water during summer low flows. Almost all sites in the database were selected using a randomised survey design, constituting a representative sample of headwater streams in these ecoregions. 2. Macroinvertebrate taxonomic and functional feeding group composition were very similar among the three ecoregions in the study area (Coast Range, Cascades and Klamath Mountains). On average, 55% of the individuals at each site were in the orders Ephemeroptera, Plecoptera or Trichoptera. Dipteran taxa (mostly chironomids) accounted for another 34%. At almost all sites, non‐insects made up <10% of the macroinvertebrate assemblage. Almost half (49%) of the assemblages were collectors; remaining individuals were about evenly divided among scrapers, shredders and predators. 3. There were 189 different macroinvertebrate taxa at the 167 sites with richness at individual sites ranging from 7 to 71 taxa. Ordination by non‐metric multidimensional scaling revealed a strong association between % Ephemeroptera, especially Baetis, and site scores along the first axis. This axis was also strongly related to % coarse substratum and fast water habitat. The second axis was strongly related to % intolerant individuals, site slope and altitude. No strong relationships were evident between any ordination axis and either logging activity, presence/absence of fish, catchment size or ecoregion. 4. Based on macroinvertebrate index of biotic integrity (IBI) scores, 62% of the sites had no impairment, 31% of the sites had slight impairment and only 6% of the sites had moderate or severe impairment. IBI scores were not strongly related to forest harvest history. All four severely impaired sites and five of the seven sites with moderate impairment were lower altitude, shallower slope stream reaches located in the Coast Range with evidence of agricultural activity in their catchment or riparian zone. % sand + fine substratum was the environmental variable most strongly related to macroinvertebrate IBI.     

The Influence of Environmental,Biotic and Spatial Factors on Diatom Metacommunity Structure in Swedish Headwater Streams

Stream assemblages are structured by a combination of local (environmental filtering and biotic interactions) and regional factors (e.g., dispersal related processes). The relative importance of environmental and spatial (i.e., regional) factors structuring stream assemblages has been frequently assessed in previous large-scale studies, but biotic predictors (potentially reflecting local biotic interactions) have rarely been included. Diatoms may be useful for studying the effect of trophic interactions on community structure since: (1) a majority of experimental studies shows significant grazing effects on diatom species composition, and (2) assemblages can be divided into guilds that have different susceptibility to grazing. We used a dataset from boreal headwater streams in south-central Sweden (covering a spatial extent of ∼14000 km²), which included information about diatom taxonomic composition, abundance of invertebrate grazers (biotic factor), environmental (physicochemical) and spatial factors (obtained through spatial eigenfunction analyses). We assessed the relative importance of environmental, biotic, and spatial factors structuring diatom assemblages, and performed separate analyses on different diatom guilds. Our results showed that the diatom assemblages were mainly structured by environmental factors. However, unique spatial and biological gradients, specific to different guilds and unrelated to each other, were also evident. We conclude that biological predictors, in combination with environmental and spatial variables, can reveal a more complete picture of the local vs. regional control of species assemblages in lotic environments. Biotic factors should therefore not be overlooked in applied research since they can capture additional local control and therefore increase accuracy and performance of predictive models. The inclusion of biotic predictors did, however, not significantly influence the unique fraction explained by spatial factors, which suggests low bias in previous assessments of unique regional control of stream assemblages.     

SPATIAL PATTERNS AND ECOLOGICAL DETERMINANTS OF BENTHIC ALGAL ASSEMBLAGES IN MID-ATLANTIC STREAMS, USA

We attempted to identify spatial patterns and determinants for benthic algal assemblages in Mid-Atlantic streams. Periphyton, water chemistry, stream physical habitat, riparian conditions, and land cover/use in watersheds were characterized at 89 randomly selected stream sites in the Mid-Atlantic region. Cluster analysis (TWINSPAN) partitioned all sites into six groups on the basis of diatom species composition. Stepwise discriminant function analysis indicated that these diatom groups can be best separated by watershed land cover/use (percentage forest cover), water temperature, and riparian conditions (riparian agricultural activities). However, the diatom-based stream classification did not correspond to Omernik's ecoregional classification. Algal biomass measured as chl a can be related to nutrients in habitats where other factors do not constrain accumulation. A regression tree model indicated that chl a concentrations in the Mid-Atlantic streams can be best predicted by conductivity, stream slope, total phosphorus, total nitrogen, and riparian canopy coverage. Our data suggest that broad spatial patterns of benthic diatom assemblages can be predicted both by coarse-scale factors, such as land cover/use in watersheds, and by site-specific factors, such as riparian conditions. However, algal biomass measured as chl a was less predictable using a simple regression approach. The regression tree model was effective for showing that ecological determinants of chl a were hierarchical in the Mid-Atlantic streams.     

Nitrogen Export from Forested Watersheds in the Oregon Coast Range: The Role of N<Subscript>2</Subscript>-fixing Red Alder

Variations in plant community composition across the landscape can influence nutrient retention and loss at the watershed scale. A striking example of plant species importance is the influence of N₂-fixing red alder (Alnus rubra) on nutrient cycling in the forests of the Pacific Northwest. To understand the influence of red alder on watershed nutrient export, we studied the chemistry of 26 small watershed streams within the Salmon River basin of the Oregon Coast Range. Nitrate and dissolved organic nitrogen (DON) concentrations were positively related to broadleaf cover (dominated by red alder: 94% of basal area), particularly when near-coastal sites were excluded (r ² = 0.65 and 0.68 for nitrate-N and DON, respectively). Nitrate and DON concentrations were more strongly related to broadleaf cover within entire watersheds than broadleaf cover within the riparian area alone, which indicates that leaching from upland alder stands plays an important role in watershed nitrogen (N) export. Nitrate dominated over DON in hydrologic export (92% of total dissolved N), and nitrate and DON concentrations were strongly correlated. Annual N export was highly variable among watersheds (2.4–30.8 kg N ha^–1 y^–1), described by a multiple linear regression combining broadleaf and mixed broadleaf–conifer cover (r² = 0.74). Base cation concentrations were positively related to nitrate concentrations, which suggests that nitrate leaching increases cation losses. Our findings provide evidence for strong control of ecosystem function by a single plant species, where leaching from N saturated red alder stands is a major control on N export from these coastal watersheds.     

Diatoms from Alto Adige/Südtirol (Northern Italy): characterization of assemblages and their application for biological quality assessment in the context of the Water Framework Directive

The Water Framework Directive 2000/60/EC (WFD) requires the analysis of biological elements of aquatic ecosystems to assess water quality. Diatoms are the component of the periphyton most commonly used to classify lotic environments. Within the context of the WFD the concept of ‘reference conditions’ was introduced and biological quality of watercourses is expressed as Ecological Quality Ratio (EQR). This study was carried out in Alto Adige/Südtirol (Province of Bolzano-Bozen, northern Italy), belonging to the Alpine eco-region, and to the hydro-ecoregion Inner Alps. During 2006–2009, epilithic diatoms were sampled from monitoring and reference sites of seven stream types. Diatom assemblages were analysed with TWINSPAN and CCA analyses to investigate species association and distribution in relation to stream characteristics. Altitude and geology resulted to be the most important factors influencing diatom assemblage composition, and were used to describe new stream types. Indicator species analysis was used to characterize reference assemblages. The biological quality of watercourses was assessed using different diatom indices: Specific Pollution sensitivity Index (IPS), Eutrophication and Pollution Index with Diatoms (EPI-D), Trophic Index (TI). We tested also the Intercalibration Common Metric index (ICM).     

Spatial factors contribute to benthic diatom structure in streams across spatial scales: Considerations for biomonitoring

Diatoms are widely used in the biological monitoring of streams because they are strong responders to environmental change, but dispersal and spatial factors can play important and potentially confounding roles in the presence, absence, and abundance of species along with characterizing species–environment relationships. To examine how spatial factors affect diatom community structure and biomonitoring, multiple scales were sampled including the Western Allegheny Plateau (n = 58), Leading Creek watershed (n = 18), and the adjacent Shade River watershed (n = 21) in southeast Ohio. Partitioning of spatial, environmental, and spatially-structured environmental variation was conducted on diatom assemblages and on diatom metrics used in biomonitoring. At the regional scale, diatom assemblages and metrics had strong relationships with agricultural (e.g., significant correlations with nutrients, conductivity, and pasture/row crops in the watershed) and alkalinity gradients. Diatom assemblages and metrics in both watersheds were strongly associated with acid mine drainage (AMD) impacts, and when spatial factors were set as covariables in CCAs, relationships with AMD gradients became even stronger, indicating the need to consider how spatial factors could reduce the strength of diatom-environment relationships. Metrics calculated at all scales had very little variation explained exclusively by spatial factors, likely because multiple species are combined into a simplified metric that reduces the effects of species dispersal. Local environmental variables accounted for 57, 42, and 42% of the total variation explained (TVE), and spatial variables accounted for 28, 31, and 37% of the TVE in the regional, Leading Creek, and Shade River datasets, respectively. The amounts of variation in diatom assemblages explained solely by spatial factors at these scales were substantial and similar to what has been reported at continental, national, and large regional (Level I Omernik ecoregions) scales (approximately 1/3 of TVE). Although amounts of variation explained are similar across scales, processes underlying the spatial structure likely differ. In addition to describing ecological patterns, recognizing the potential influence of spatial factors could improve the identification and management of environmental problems at a range of scales, as well as aid in the development of new research questions and hypotheses aimed at exploring factors that could explain portions of the spatially explicit variation.     

Environmental and spatial influences on benthic community composition in wooded headwater streams in Zoar Valley, New York, USA

This study determined the relative influences of environmental variation versus spatial autocorrelation on benthic macroinvertebrate community composition of temperate headwater streams. We enumerated fauna in riffle zones of 23 separate (i.e., not tributaries to each other) but closely grouped from first- to third-order woodland streams surrounding Zoar Valley Canyon, western New York State, USA, during spring, summer, and fall of 2006. Watershed geomorphology (stream order, catchment area, and forest cover) and semi-quantitative habitat characteristics, the latter of which were incorporated into a Qualitative Habitat Evaluation Index (QHEI), were also recorded for each stream. Non-metric Multidimensional Scaling (NMDS) ordination was used to establish patterns of biotic similarity among these streams. Matrices of biological and environmental Euclidean distances were constructed for all between-stream pairings. Additionally, a between-stream spatial matrix was constructed based on global position system coordinates of sampling sites. Non-significant partial Mantel coefficients indicated that biological distances were uncorrelated with spatial distances both among all 23 study streams and among 12 first-order streams only. In contrast, biological distances were positively correlated with environmental distances (r _M = 0.375 and 0.289 for all streams and for first-order streams only, respectively; P values < 0.05). Environmental and spatial distances were uncorrelated (partial Mantel P values > 0.05), indicating that the measured environmental characteristics were not spatially structured. Each of the geomorphological and habitat variables was statistically associated with NMDS community composition axes (stepwise multiple regression, one-factor MANOVA). These results suggest that environmental filters and niche-based species sorting may operate here between separate streams, and that study sites appear to be effectively insular in their biota despite close physical proximity. Handling editor: Sonja Stendera     

The roles of elevation and local environmental factors as drivers of diatom diversity in subarctic streams

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SURVEY OF BENTHIC DIATOM COMMUNITIES FROM LOTIC SYSTEMS WITHIN THE WESTERN ALLEGHENY PLATEAU

Over the course of 3 years (1997–1999), 72 stream sites were sampled for epilithic diatom communities. The analysis of these samples has led to the identification of over 325 species of diatoms. In addition to sampling the diatom community, selected physical and chemical parameters were recorded from each stream reach. These parameters included pH, specific conductance, current velocity, SRP, nitrate, silica, and total alkalinity. Canonical Correspondence Analysis (CCA) was used to identify influential environmental parameters and to assess the response of the diatom community to prominent anthropogenic inputs in the region (i.e. coal mine drainage, eutrophication). The initial analyses indicate that pH was the most influential environmental parameter along the first CCA axis. This shift was not unexpected, as acid mine drainage (AMD) in the region leads to a wide range of pH values (2.8–7.93). The highly acidic sites were characterized by species of the genus Eunotia (specifically E. exigua and E. steineckei), Frustulia rhomboides, and Pinnularia subcapitata. Furthermore, Achnanthidium minutissimum was the most widely distributed of the diatom species encountered, being found at 94% of the sites sampled. Streams that fluctuated between acidic and circumneutral pH (termed “teeter‐totter”) had greater abundances of Brachysira vitrea than other streams in this survey. Further implications for the use of these diatom communities as biomonitoring tools and the distribution of assemblages within the Western Allegheny Plateau will be discussed.     

Diatom biomonitoring of streams: Reliability of reference sites and the response of metrics to environmental variations across temporal scales

Benthic diatoms are widely used indicators of human impacts on stream ecosystems because they are very responsive to changing environmental conditions. However, little research has explicitly focused on their reliability with regards to temporal variation in assemblage structure and environmental conditions. We examined variability in diatom-environment relationships at bi-weekly, monthly, and yearly time scales from 7 reference, 7 agricultural, and 2 acid mine drainage (AMD)-impacted streams, and how nutrient and pH fluctuations may affect the interpretation of diatom metrics and the Diatom Model Affinity (DMA) index. Reference streams had less bi-weekly variability in NO₃-N concentrations than non-reference streams. The % eutraphentic diatoms and DMA scores were more strongly correlated with seasonal means of NO₃-N and PO₄-P concentrations than with same day concentrations. Most nutrient indicator metrics had strong correlations with watershed land use. All 14 non-AMD streams experienced substantial increases in NO₃-N and decreases in temperature from November to May, which were associated with high species turnover, substantial changes in community structure, reduced diversity and richness, increased relative abundances of high nutrient diatoms, and decreases in low nutrient diatoms and DMA scores. The % acidophilic diatoms and DMA scores were significantly correlated with increased pH associated with greater precipitation at AMD sites from December to April (r = ?0.77, r = 0.62, respectively; P < 0.01). Yearly, DMA scores for all reference streams were consistently in the minimally impaired category, whereas scores for non-reference streams varied among impairment categories. Reference sites serve as reliable benchmarks for diatom ecological integrity during the summer. In this region, June to October is a recommended time period for diatom sampling in monitoring programs because subsequent shifts in hydrologic regimes, nutrients, and diatom assemblages occurred, affecting all sites and masking among stream differences attributable to agricultural land uses.     

The roles of environment and space in shaping stream diatom communities

Diatoms are widely used in stream quality assessment due to their response to the local environment. Diatoms are also influenced by many large-scale processes and so the diatom communities of boreal streams incorporate a strong spatial component at a regional level. What is not properly known yet is whether the variation in diatom communities between regions is larger than the variation in measured environmental variables. We studied the roles of environment and space in accounting for variability in stream diatom communities across four regions in Finland. According to canonical correspondence analysis, geographical coordinates, nutrient concentrations (total N and P), and water conductivity were the most important factors affecting variation in diatom community composition. Of physical factors, depth and current velocity were also significant. According to Mantel tests, both environmental and geographical distances were related to dissimilarity in diatom community composition. Analysis of Similarities indicated that the regional differences in diatom community composition were larger than the regional differences in environmental variables. We also found many indicator species confined to certain regions. Our results suggest that the four study regions differ in their diatom species composition more than in their environmental features and that diatoms are structured not only by the local environment but also by large-scale processes, possibly related to history, climate and dispersal. These results imply that, while diatom species composition reflects well the environmental differences between regions, future bioassessments would benefit from regional stratification. Otherwise, relationships with environmental variables may be masked by trans-regional differences in species pools caused by the large-scale processes.     

Spatial and temporal variation of fish assemblages and their associations to habitat variables in a mountain stream of north Tiaoxi River,China

The spatial and temporal variations of the fish assemblages in mountain streams of China are poorly understood. The relationships between the fish assemblage and selected habitat features were examined in the North Tiaoxi River, one of headwaters of Taihu Lake. A total of 3,348 individuals belonging to 5 orders, 11 families, 25 genera and 34 species were collected including 33 native species and one invasive species. Among those, about 20 species were endemic to China. Non-metric Multidimensional Scaling (NMDS) was applied to compare fish assemblage structures from upstream to downstream during four seasons. Species assemblages differed along the stream continuum, but there was little apparent change associated with the seasons. Species richness and Shannon-Weaver index (H′) tended to increase along the stream continuum from the upstream to downstream and the proportion of invertivorous fish tended to significantly decrease along the continuum with a parallel significant increase in the percentage of omnivores. Fish assemblages were significantly related to both water quality and habitat structure variables. Canonical Correspondence Analysis ordinations (CCA) revealed that 6 of the 14 selected environmental variables had significant relationships with the fish assemblage such as distance to source, stream width, altitude, pH, water depth, and water velocity and different sampling sites were associated with different environmental variables in different seasons. The main differences in fish assemblage structure and diversity within the whole watercourse are probably related to large-scale factors such distance to source, altitude and stream width. Differences of instream characteristics are likely to be caused by natural variability of the ecosystems but also, in some case, by anthropogenic influence like human settlements, agriculture and river embankment and pollution from small factory.     

Distribution of benthic diatoms in relation to environmental variables in lowland streams

The composition (% relative abundance) of diatom assemblages from soft bottom sediments was studied at 75 sites situated in 46 rivers, brooks, and ditches in the islands of Hiiumaa and Saaremaa and in the lowland of West Estonia. Although the total number of recorded taxa was 205, the studied diatom assemblages consisted of 54, 55, 48, and 50 constant species in the drainage basin of Moonsund, the Gulf of Riga, Hiiumaa, and Saaremaa, respectively. The habitats of the dominating taxa were heterogenous and the most widespread species were Achnanthidium minutissimum, Martyana martyi, Meridion circulare, Cocconeis placentula, Planothidium lanceolatum, and Amphora pediculus. The Shannon–Weaver diversity (H) index varied from 2.09 to 4.63. Multivariate analyses were used to identify the environmental variables governing the composition and structure of the benthic diatom assemblage. Principal Component Analysis (PCA) and Redundancy Analysis (RDA), based on 56 most abundant taxa, indicated differences in the composition and structure of diatom assemblages between different drainage basins as well as between the upper and lower stream courses. In the headwaters there prevailed small epipsammic diatoms (Martyana, Planothidium, Staurosira, Staurosirella). Different motile epipelic species from the genera Amphora, Navicula, Nitzschia, etc. were distributed abundantly in the lower courses of the streams. There was a positive correlation between order of the stream site and trophic level of water (R=0.35; p<0.05). Along a river system, the increasing order of the stream was accompanied by higher trophic level of water.