Non-wadeable river bioassessment: spatial variation of benthic diatom assemblages in Pacific Northwest rivers,USA

Current bioassessment efforts are focused on small wadeable streams, at least partly because assessing ecological conditions in non-wadeable large rivers poses many additional challenges. In this study, we sampled 20 sites in each of seven large rivers in the Pacific Northwest, USA, to characterize variation of benthic diatom assemblages among and within rivers relative to environmental conditions. Analysis of similarity (ANOSIM) indicated that diatom assemblages were significantly different among all the seven rivers draining different ecoregions. Longitudinal patterns in diatom assemblages showed river-specific features. Bray–Curtis dissimilarity index values did not increase as a function of spatial distance among the sampled reaches within any river but the Malheur. Standardized Mantel r of association between assemblage similarity and spatial distance among sites ranged from a high of 0.69 (Malheur) to a low of 0.18 (Chehalis). In the Malheur River, % monoraphids, nitrogen-tolerant taxa, and beta-mesosaprobous taxa all decreased longitudinally while % motile taxa, especially Nitzschia, showed an opposite trend, reflecting a strong in-stream water quality gradient. Similar longitudinal trends in water quality were observed in other rivers but benthic diatom assemblages showed either weak response patterns or no patterns. Our study indicated that benthic diatom assemblages can clearly reflect among-river factors. The relationships between benthic diatom assemblages and water quality within each river may depend on the strength of the water quality gradients, interactive effects of water quality and habitat conditions, and diatom sampling design.     

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.     

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).     

Epilithic diatom assemblages and their relationship to environmental characteristics in an agricultural watershed (Guadiana River,SW Spain)

Large-scale patterns of benthic diatom assemblages were analyzed in an agricultural basin, the Guadiana River. The distribution patterns of epilithic diatom assemblages were analyzed at different spatial scales: the whole watershed, the upper calcareous subcatchment and the mid-lower siliceous subcatchment. At the whole watershed scale, two major ecological gradients were revealed. The first one summarized the diatom distribution throughout a nutrient concentration gradient, while the second gradient was related to the geological structure of the watershed. Variance partitioning allowed the effects of the different sets of environmental parameters related to every CCA gradient to be separated. Analyzing the subcatchment gradients with partial CCA allowed us to define specific key factors that affect diatom species composition. Although water chemistry consistently played the most important role in structuring diatom assemblages in the Guadiana, spatial factors such as altitude or geographic location also explained some variation in diatom distribution.     

How does water scarcity affect spatial and temporal patterns of diatom community assemblages in Mediterranean streams?

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Impact of Urban Pollution from the Hanoi Area on Benthic Diatom Communities Collected from the Red, Nhue and Tolich Rivers (Vietnam)

The effects of urban pollution from Hanoi city on the benthic diatom communities of the Nhue–Tolich river system were studied during the 2003 dry season. Benthic diatoms were allowed to grow on glass slides suspended in the water flow for 4 weeks. To reveal the relationship between water quality and diatom communities, Canonical correspondence analysis (CCA) was used on data concerning relative abundances of diatom species and environmental variables. Two diatom indices, IPS and DAIpo, were applied to evaluate water quality in the three rivers. A total of 291 diatom taxa were found in the Red, Nhue and Tolich Rivers. These were mainly cosmopolitan taxa, with some tropical, subtropical and endemic taxa. The most abundant taxa at the Red site were Aulacoseira granulata, Achnanthidium minutissimum, Encyonema minutum, Navicula recens and other halophilous taxa such as Nitzschia kurzii, Seminavis strigosa, Entomoneis paludosa, Bacillaria paradoxa. Diatom assemblages at the Tolich site consisted mainly of Nitzschia umbonata, Nitzschia palea and Eolimna minima. Diatom density ranged from 660 to 30,000 cells/cm². Environmental variables and diatom assemblage composition at all sites were significantly correlated. Two diatom indices gave similar results and indicate the Tolich River with the lowest values as a highly polluted site.     

Importance of scalar and riparian habitat effects for assessment of ecological status using littoral diatoms

The classification of waterbodies under the Water Framework Directive is dependent on the ability of monitoring programmes to reflect habitat quality using biotic elements including benthic diatom communities. This study investigated the influence of specific riparian habitats, of mixed woodland, grassland and lake artificial structures such as jetties and slipways, on benthic diatom assemblages in nine lakes across gradients of total phosphorus, alkalinity and in the presence or absence of Dreissena polymorpha. The heterogeneity of the benthic diatom assemblages at riparian and lake scale was assessed by taking three replicates per site category per lake, following standard European Union protocols. Canonical correspondence analysis (CCA) and mixed effect modelling was used to investigate the main environmental controls on assemblage structure. Non-metric Multidimensional Scaling (NMDS) was used to examine patterns in assemblage structure. No single environmental gradient was found to control benthic diatom composition, with differences among assemblages influenced both by riparian habitat type within lakes and interaction of multiple environment gradients, including presence of D. polymorpha. Greater control was exerted on community structure at the lake than local riparian scale. The influence of scalar factors on diatom assemblages increased with increasing scale. We recommend that for effective monitoring and assessment of ecological status, standard sampling protocols should include localised littoral habitats with individual samples pooled across riparian habitat types, thereby accounting for both multiple environmental and spatial controls on community structure.     

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.     

Diatom Assemblages and their Associations with Environmental Variables in Oregon Coast Range Streams, USA

The Oregon Coast Range, rich in natural resources, is under increasing pressure from rapid development. The purpose of this study was to examine diatom species patterns in relation to environmental variables in streams of this region. Diatoms, water quality, physical habitat and watershed characteristics were assessed for 33 randomly selected stream sites. Watershed size, elevation, geology, vegetation and stream morphology varied substantially among sites. Streams were characterized by dilute water chemistry and a low percent of fine substrate. A total of 80 diatom taxa were identified. Taxa richness was low throughout the region (median 15, range 10–26). Assemblages were dominated by two adnate species, Achnanthidium minutissimum and Achnanthes pyrenaicum. Diatoms sensitive to organic pollution dominated the assemblages at all sites (median 85%). Non-metric multidimensional scaling (NMDS) and correlational analysis showed quantitative relationships between diatom assemblages and environmental variables. NMDS axes were significantly correlated with watershed area, watershed geology, conductivity, total nitrogen, total solids and stream width. Diatom-based site classification (Two-way Indicators Species Analysis, (TWINSPAN)) yielded 4 discrete groups that displayed weak correlations with environmental variables. When stream sites were classified by dominant watershed geology, overall diatom assemblages between groups were significantly different (Analysis of Similarity (ANOSIM) global R = 0.19, p < 0.05). Our results suggest that streams in the coastal region are in relatively good condition. High natural variability in stream conditions in the Oregon Coast Range ecoregion may obscure quantitative relationships between environmental variables and diatom assemblages. A bioassessment protocol that classifies sites by major landscape variables and selects streams along the major human disturbance gradient might allow for detection of early signs of human disturbance in environmentally heterogeneous regions, such as the Pacific Northwest.     

Consistent patterns in diatom assemblages and diversity measures across water‐depth gradients from eight Boreal lakes from north‐western Ontario (Canada)

1. Until recently, the distribution of diatom species assemblages and their attributes (e.g. species richness and evenness) in relation to water depth have been identified but not quantified, especially across several lakes in a region. Here, we examined diatom assemblages in the surface sediment across a water‐depth gradient in eight small, boreal lakes in north‐western Ontario, minimally disturbed by human activities. 2. Surface‐sediment diatom assemblages were collected within each lake along a gentle slope from near‐shore to the centre deep basin of the lake, at a resolution of ～1 m water depth. Analysis of sedimentary samples provided an integrated view of assemblages that were living in the lake over several years and enabled a high‐resolution analysis of many lakes. The study lakes ranged in water chemistry, morphology and size and are located along an east–west transect approximately 250 km long in north‐western Ontario (Canada). 3. The majority of diatom species were distributed along a continuum of depth, with those taxa having similar habitat requirements forming distinct, though overlapping, assemblages. Three major zones of diatom assemblages in each lake were consistently identified: (i) a near‐shore assemblage of Achnanthes (sensu lato), Nitzschia, Cymbella (sensu lato) and other benthic species; (ii) a mid‐depth assemblage of small Fragilaria (sensu lato)/small Aulacoseira and various Navicula taxa; and (iii) a deep‐water assemblage of planktonic origin (mainly Discotella spp.). 4. The depth of the transition between assemblage zones varied between the eight lakes. The boundary between the deep‐water planktonic zone and the mid‐depth benthic zone varied according to water chemistry and was probably related to light attenuation. The boundary was deeper in lakes with the lower dissolved organic carbon and total phosphorus (TP) (i.e. less light attenuation) and vice versa. 5. Generally, species richness, species evenness and turnover rate of species as a function of depth were significantly lower in the planktonic assemblage zone in comparison with the two zones nearer the shore. Reproducibility of species and assemblage distributions across the depth gradient of the lakes illustrated that, despite potential for sediment transport, detailed ecological characterisation of diatom species can be gleaned from sedimentary data. Such data are often lacking, particularly for near‐shore benthic species.     

Benthic diatom assemblages and their response to human stress in small-sized volcanic-siliceous streams of central Italy (Mediterranean eco-region)

The river typology ‘volcanic rivers’ represents a unique typology in the Mediterranean eco-region in Europe but its type-specific diatom assemblages are still poorly investigated. The main aims of this study were (i) to analyse the diatom diversity of this river type, (ii) to investigate the relationships between diatom diversity and environmental variables and (iii) to identify the characterizing diatom community structure of different environmental groups. To this end, we selected 13 streams located in central Italy, belonging to a single river type and covering the whole quality range from human impacted sites to those in ‘reference conditions’. Most frequent and abundant diatom species found were Cocconeis placentula var. lineata, Planothidium lanceolatum, Rhoicosphenia abbreviata and Amphora pediculus. Species richness in reference and less impacted sites was not higher than in intermediate and heavily degraded ones, but non-metric multidimensional scaling (N-MDS) analysis revealed a substantial difference among their diatom community structure. The indicator species analysis defined the characteristic species of three environmental groups identified by N-MDS. The findings of this study represent a contribution to the knowledge on diatom communities of Mediterranean volcanic-siliceous rivers and provide information for a data-driven implementation of the Water Framework Directive in Italy for this specific river typology.     

Diatom, chrysophyte and protozoan distributions along a latitudinal transect in Fennoscandia

Diatoms, chrysophyte scales and cysts, and the siliceous plates of thecoamoebae were studied from the surface sediments of 20 freshwater habitats in Fennoscandia The study sites are distributed along a latitudinal transect extending from southern Finland to northern Norway, spanning boreal forest through arctic tundra vegetational zones Diatom assemblages were usually dominated by acidophilic, periphytic members of the genera Achnanthe, Fragilaria and Navicula Marked shifts in diatom assemblage composition were recorded along the latitudinal transect, whereas scaled chrysophytes were rare in all study sites Meanwhile, siliceous protozoa were common, but did not exhibit any noticeable trends in assemblage composition with changing latitude A comparison of the Fennoscandian diatom assemblages with those recorded from freshwater sites near Yellowknife (central Northwest Territories, Canada) revealed similar trends in diatom assemblage composition with changing ecoclimatic zones in both regions Moreover, canonical correspondence analysis showed that diatom assemblages in cold, dilute tundra sites were effectively separated from sites with forested catchments from both the Fennoscandian and Canadian transects The general similarity between the two regions suggests that autecological data and the resulting environmental transfer functions based on diatom assemblages may eventually be joined from North American and European regions     

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.     

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.     

Determinants of Benthic Diatom Community Structurein Boreal Streams: the Role of Environmental and Spatial Factors at Different Scales

Benthic diatoms were sampled at 197 sites in Finnish boreal streams. The diatom distribution patterns were related to environmental and spatial factors at three spatial scales using Canonical Correspondence Analysis (CCA). The results emphasized the predominance of chemical‐constituent concentration and ion composition on structuring benthic diatom communities in running waters. Within a river system, physical factors had notable influence on diatom community structure reflecting mainly only minor changes in water chemistry within a watershed. Partial CCA revealed that pure spatial component explained ca. 20% of explainable variation in diatom data at each of the three scales. Environmental factors captured 53–78% of explained variation in species data at ecoregion and river system level. According to Procrustean Randomization Test, spatial coordinates of the study sites and patterns in diatom community structure were strongly concordant (m² = 0.862, p = 0.001) across the largest spatial scale. Similarly, at smaller spatial scales in southern and central Finland, congruence was significant. These data support the view that diatom communities exhibit a rather strong spatial component especially at largest, national scale.     

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.     

Temporal trends in algae, benthic invertebrate, and fish assemblages in streams and rivers draining basins of varying land use in the south-central United States, 1993–2007

Site-specific temporal trends in algae, benthic invertebrate, and fish assemblages were investigated in 15 streams and rivers draining basins of varying land use in the south-central United States from 1993–2007. A multivariate approach was used to identify sites with statistically significant trends in aquatic assemblages which were then tested for correlations with assemblage metrics and abiotic environmental variables (climate, water quality, streamflow, and physical habitat). Significant temporal trends in one or more of the aquatic assemblages were identified at more than half (eight of 15) of the streams in the study. Assemblage metrics and abiotic environmental variables found to be significantly correlated with aquatic assemblages differed between land use categories. For example, algal assemblages at undeveloped sites were associated with physical habitat, while algal assemblages at more anthropogenically altered sites (agricultural and urban) were associated with nutrient and streamflow metrics. In urban stream sites results indicate that streamflow metrics may act as important controls on water quality conditions, as represented by aquatic assemblage metrics. The site-specific identification of biotic trends and abiotic–biotic relations presented here will provide valuable information that can inform interpretation of continued monitoring data and the design of future studies. In addition, the subsets of abiotic variables identified as potentially important drivers of change in aquatic assemblages provide policy makers and resource managers with information that will assist in the design and implementation of monitoring programs aimed at the protection of aquatic resources.     

PERIPHYTIC DIATOM ASSEMBLAGES FROM ULTRA‐OLIGOTROPHIC AND UV TRANSPARENT LAKES AND PONDS ON VICTORIA ISLAND AND COMPARISONS WITH OTHER DIATOM SURVEYS IN THE CANADIAN ARCTIC1

Periphytic diatoms are potentially powerful indicators of environmental change in climatically‐sensitive high latitude regions. However, only a few studies have examined their taxonomic and ecological characteristics. We identified and enumerated diatom assemblages from sediment, rock, and moss habitats in 34 ultra‐oligotrophic and highly transparent lakes and ponds on Victoria Island, Arctic Canada. The similar limnological characteristics of the sites allowed us to examine the influence of habitat, independent of water chemistry, on the diatom assemblages. As is typical in shallow arctic water bodies, benthic taxa, including species of Achnanthes, Caloneis, Cymbella, Navicula, and Nitzschia, were most widely represented. Minor gradients in our measured environmental variables did not significantly explain any variance in diatom species, but there were marked differences in diatom assemblages among sites. Pond ephemerality seems to explain some diatom variation, because aerophilic taxa such as Achnanthes kryophila Petersen and A. marginulata Grunow were dominant in shallow sites that had undergone appreciable reductions in volume. We identified several taxa that exhibited strong habitat preferences to sediment, moss, or rock substrates and also found significant differences (P < 0.01) in diatom composition among the three habitats. In comparisons with three similar diatom surveys extending over 1200 km of latitude, we determined that surface sediment assemblages differed significantly (P < 0.001) among all regions examined. Diatom species diversity was inversely related to latitude, a result likely explained by differences in the lengths of growing seasons. These data contribute important ecological information on diatom assemblages in arctic regions and will aid in the interpretation of environmental changes in biomonitoring and paleolimnological studies.     

Benthic Algae in High Altitude Streams of the Alps – a Neglected Component of the Aquatic Biota

This is a review on benthic algae from streams situated above the tree line in the Alps. It integrates published and unpublished data from alpine streams in Austria and in the Trento Province (Northern Italy). The main focus is on the structural and taxonomic composition of benthic algae including macro- and micro-algae and their contribution to the epilithic biofilm and the stream food-web. The environmental conditions relevant to algae within the two major stream types, the glacial (glacier-fed) and non-glacial krenal (spring-fed) stream are discussed. The paper considers both the maximum possible structural complexity of transverse algae zonation in cascading alpine/subalpine stream segments, and the effects of glacial water on species richness in the Central Alps in Austria. Autecological data are given for 46 common diatoms from 42 sites in the Central Eastern and Southern Central Alps and for 30 algae in addition to diatoms for 22 streams in the Central Eastern Alps. Since data on responses of benthic algae to the harsh conditions in high altitude Alpine streams are very scarce, results from literature and the author’s experiences from these and other mountain stream types are synthesized to formulate major objectives for future research in benthic high altitude Alpine stream ecology.