Isotopic variation in five species of stream fishes under the influence of different land uses

The aim of this study was to test if changes in land use alter the isotopic signature of fish species, promoting changes in the trophic position and food resource partitioning between these consumers. Three different systems were investigated: pasture streams (n = 3), streams in sugar cane plantations (n = 3) and reference streams (n = 3). Fish species Aspidoras fuscoguttatus, Astyanax altiparanae, Characidium zebra, Hisonotus piracanjuba and Knodus moenkhausii were selected, and their nitrogen and carbon isotopic compositions were estimated to assess changes in the trophic level and partitioning of food items consumed. The composition of δ¹³C (‰) only differed among the land use categories for A. altiparanae, H. piracanjuba and K. moenkhausii. Resource partitioning was different for all species, with changes in the sources or proportions they consumed in each land use category, but only A. altiparanae introduced new food sources in large quantity in altered land uses. It is important to note, however, that the results from the resource partitioning analysis are limited due to large overlapping of isotopic signatures between the analysed food resources. All fish species exhibited variation in δ¹⁵N (‰), with the highest values found in streams under sugar cane or pasture influence. Despite the variation in nitrogen isotopic values, only C. zebra and H. piracanjuba displayed changes in trophic level. Therefore, it is believed that the increase in the δ¹⁵N (‰) value of the individuals collected in streams under the influence of sugar cane or pasture was due to the greater influence of livestock dung and chemical and organic fertilizers. The results also highlight the importance of studying consumer species along with all forms of resources available at each location separately, because the signatures of these resources also vary within different land uses.     

Land-use effects on structural and functional composition of benthic and leaf-associated macroinvertebrates in four Andean streams

The replacement of native forests by pastures takes place widely in the Andes. The effects of such land-use change on aquatic assemblages are poorly understood. We conducted a comparative analysis of the effects of forest conversion to pastures on the taxonomic, structural, and functional composition of macroinvertebrates (benthic and leaf-associated) in montane and upper montane streams (ecosystem type) of the south Ecuadorian Andes. Taxonomic composition of benthic and leaf-associated macroinvertebrates was different between ecosystem type and land use. Also, major differences in the structural and functional composition of benthic and leaf-associated macroinvertebrates were mainly promoted by land use in both ecosystem types. Forested streams showed higher diversity than pasture streams, sustaining more shredder, scraper, and predatory invertebrates. We also observed differences in the macroinvertebrate communities between benthic and leaf-bag samples. Leaf bags had lower diversity and more collector invertebrates than benthic samples. This study highlights the large effect of riparian forest conversion to pasture land on macroinvertebrate communities, and the importance of using appropriate sampling techniques to characterize aquatic assemblages. We also recommend the maintenance and restoration of riparian vegetation to mitigate the effects of deforestation on stream communities and ecosystem processes.     

Fish determine macroinvertebrate food webs and assemblage structure in Greenland subarctic streams

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The effects of land use on environmental features and functional organization of macroinvertebrate communities in Patagonian low order streams

Benthic invertebrates, water quality variables, chlorophyll a and particulate organic matter (POM) were studied in 18 sites of mountain streams in Patagonia (Argentina) subjected to six different land uses: native forest, pine plantation, pasture, harvest forest, urban and reference urban. Three streams of each land use were studied in March 2006. Macroinvertebrates were sampled in three riffles and three pools (n = 108) and biomass of detrital fractions of POM were quantified. Overall benthic POM biomass was higher at native and harvest forest than pastures, whereas fine fraction (FPOM) was higher in harvest forest than in pastures. Regarding to autotrophic subsidies bryophytes were the only that changed consistently among uses. These differences in energy resources were correlated with changes in community attributes. Shredder richness was clearly higher at native and harvest forest than exotic pine plantations, collector gatherers density was consistently high at harvest sites, and total density was significantly higher at urban and harvest forest. Multidimensional scaling ordination based on macroinvertebrate density data showed a clear separation of forested (either native or exotic species) from riparian modified areas (pasture, urban and harvest sites). Environmental variables having explanation power on macroinvertebrate assemblages were mostly related with: detritus availability (wood and leaves biomass) and impairment (total phosphorous and % sand). These results confirm that macroinvertebrate assemblage structure in Patagonian low order streams can be altered by land use practices. Among guild structure measures, those indicators based on benthic community functional attributes, shredders richness and collectors density, resulted good candidates to assess land use impacts. On account of riparian corridor management may be critical to the distribution of benthic taxa, the maintenance of good conditions of vegetation adjacent to rivers will enhance water quality and the environment for highly endemic headwater communities of Patagonian streams.     

Catchment land cover influences macroinvertebrate food‐web structure and energy flow pathways in mountain streams

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Influence of functional feeding groups and spatiotemporal variables on the δ<Superscript>15</Superscript>N signature of littoral macroinvertebrates

The δ¹⁵N trophic enrichment in littoral food webs is not well known despite the importance of macroinvertebrates in lacustrine energy fluxes. We wanted to assess the influence of functional feeding group (grazer, collector, shredder, predator, predator–hematophagous, predator–sucker) and spatiotemporal variables (year, month, station of sampling) on littoral macroinvertebrate δ¹⁵N signatures. For 2 years, during the plant growth period phytophilous littoral macroinvertebrates were sampled in Lake St. Pierre, a large fluvial lake of the St. Lawrence River, Québec, Canada. The δ¹⁵N analyses showed that station was the most important factor for explaining δ¹⁵N variation, followed by sampling month and functional feeding group. The organisms sampled in the stations of the south shore, which experienced greater macrophyte abundance, slower currents, and stronger NO₃ depletion exhibited higher δ¹⁵N values than those sampled on the north shore. Grazer-to-predator δ¹⁵N enrichment valued 1.6‰, which is inferior to the 3.4‰ generally admitted in food-web research. Shredders exhibited the lowest δ¹⁵N values and predators–hematophagous the highest. δ¹⁵N signature of invertebrates increased 3‰ through the summer between May and September. Only samples collected within a short period should be pooled to avoid an error value equivalent to one trophic level (1.6) enrichment. Furthermore, it is recommended not to pool macroinvertebrate samples collected at stations with differing watershed land uses.     

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.     

Does rapid glacial recession affect feeding habits of alpine stream insects?

1. Glacial retreat, accompanied by shifts in riparian vegetation and glacier meltwater inputs, alters the energy supply and trophic structure of alpine stream food webs. Our goal in this study was to enhance understanding of dietary niches of macroinvertebrates inhabiting different alpine streams with contrasting glacial and non‐glacial (groundwater, precipitation, snowmelt) water inputs in conjunction with seasonal and habitat‐specific variation in basal resource availability.
2. We measured a range of stream physico‐chemical attributes as well as carbon and nitrogen isotopes (δ¹³C, δ¹⁵N) of macroinvertebrates and primary food sources at seven sites across seasons within a Swiss glaciated catchment (Val Roseg) undergoing rapid glacial retreat (1–2 km between 1997 and 2014). Sampling sites corresponded to streams used in a previous (1997/1998) study within the same alpine catchment.
3. Physico‐chemical attributes showed wide variation in environmental conditions across streams and seasons. Significant correlation among physico‐chemical proxies of glacier meltwater (phosphate‐P, total inorganic carbon, conductivity, turbidity) and macroinvertebrate δ¹³C, δ¹⁵N, and size‐corrected standard ellipse area (a proxy for feeding niche width) values showed that the extent of glacial water input shapes the energy base among alpine streams. Feeding niche differences among common alpine stream insect taxa (Chironomidae, Baetidae, Heptageniidae) were not significant, indicating that these organisms probably are plastic in feeding behaviour, opportunistically relying on food resources available in a particular stream and season.
4. Seasonal trends in macroinvertebrate δ¹³C largely followed patterns in periphyton δ¹³C values, indicating that autochthonous resources were the main consumer energy source within the stream network, as shown previously. The overall range in macroinvertebrate δ¹³C (?33.5 to ?18.4‰) and δ¹⁵N (?6.9 to 6.7‰) values also corresponded to values measured in the previous study, suggesting that macroinvertebrates altered diets in line with changes in environmental conditions and food resources during a period of rapid glacial retreat. Our results suggest that environmental changes brought on by rapid glacial retreat have not yet caused a profound change in the trophic structure within these fluvial networks.

     

Riparian land use and the relationship between the benthos and litter decomposition in tropical montane streams

1. Although stream–catchment interactions have been analysed in some detail in temperate environments, little is known about the effects of land‐use changes in the tropics. Here, we analyse differences in benthic communities (macroinvertebrates and fungi) under two contrasting land uses (mature secondary forest and pasture) in montane streams in north‐western Ecuador and their influence on the rates of litter processing. 2. Between 2005 and 2006, we used a combination of coarse and fine mesh bags to study the relative contribution of macroinvertebrates and fungi to processing of two types of litter, Alnus acuminata and Inga spectabilis, in three‐first‐order streams running through mature secondary montane forests and adjacent downstream reaches running through pastures. At the same time, we characterised the assemblages of shreddering macroinvertebrates and fungi communities and the litter processing rates in stream reaches under both vegetation types. 3. Litter processing rates attributable to invertebrate feeding (coarse mesh bags) were significantly slower in streams running through pastures. Nevertheless, shredder diversity and richness were similar between pasture and forest sections, while shredder abundance was significantly higher in forest streams (mainly Phylloicus sp. :Trichoptera). Fungal reproductive activity and litter processing rates were low (fine mesh bags) and did not differ significantly between pasture and forest stream reaches. 4. Phylloicus sp. abundance was the best predictor of the percentage of litter remaining in coarse mesh bags across pasture and forest sites. Neither shredder diversity nor their species richness was a significant predictor of mass loss, as most of the decomposition was performed by a single keystone species. Although litter decomposition by microbial decomposers was low, fungal biomass (but not diversity) was the best variable explaining the percentage of litter remaining in fine mesh bags. 5. Our data suggest that, in these Neotropical montane streams, land use can have a significant impact on the rates of critical ecosystem processes, such as litter decomposition. In this study, this effect was not mediated by a major shift in the structure of the benthos, but by a decrease in the abundance and relative representation of a single species whose life history makes it critical to litter processing. 6. This study highlights the significant role that macroinvertebrate fauna can have in the processing of litter in Neotropical streams and the predominant role that single species can have in terms of controlling stream ecosystem‐level processes. Understanding the extent to which these patterns affect the long‐term and large‐scale functioning of stream ecosystems still needs further research and will become increasingly important in terms of managing lotic ecosystems in the context of rapid land‐use change.     

Seasonal dynamics of macroinvertebrate assemblages in the benthos and associated with detritus packs in two low-order streams with different riparian vegetation

• 1 The seasonal dynamics of the benthic macroinvertebrate assemblage, and the subset of this assemblage colonising naturally formed detritus accumulations, was investigated in two streams in south‐west Ireland, one draining a conifer plantation (Streamhill West) and the other with deciduous riparian vegetation (Glenfinish). The streams differed in the quantity, quality and diversity of allochthonous detritus and in hydrochemistry, the conifer stream being more acid at high discharge. We expected the macroinvertebrate assemblage colonising detritus to differ in the two streams, due to differences in the diversity and quantity of detrital inputs.
• 2 Benthic density and taxon richness did not differ between the two streams, but the density of shredders was greater in the conifer stream, where there was a greater mass of benthic detritus. There was a significant positive correlation between shredder density and detritus biomass in both streams over the study period.
• 3 Detritus packs in the deciduous stream were colonised by a greater number of macroinvertebrates and taxa than in the conifer stream, but packs in both streams had a similar abundance of shredders. The relative abundance of taxa colonising detritus packs was almost always significantly different to that found in the source pool of the benthos.
• 4 Correspondence analysis illustrated that there were distinct faunal differences between the two streams overall and seasonally within each stream. Differences between the streams were related to species tolerances to acid episodes in the conifer stream. Canonical correspondence analysis demonstrated a distinct seasonal pattern in the detrital composition of the packs and a corresponding seasonal pattern in the structure of the detritus pack macroinvertebrate assemblage.
• 5 Within‐stream seasonal variation both in benthic and detritus pack assemblages and in detrital inputs was of similar magnitude to the between‐stream variation. The conifer stream received less and poorer quality detritus than the deciduous stream, yet it retained more detritus and had more shredders in the benthos. This apparent contradiction may be explained by the influence of hydrochemistry (during spate events) on the shredder assemblage, by differences in riparian vegetation between the two streams, and possibly by the ability of some taxa to exhibit more generalist feeding habits and thus supplement their diets in the absence of high quality detritus.

     

Geometry of biodiversity patterning: assemblages of benthic macroinvertebrates at tributary confluences

We assessed whether tributaries in upland catchments (=watersheds) affected assemblages of benthic macroinvertebrates in mainstems, as has been reported in northern hemisphere systems. Eight confluences of small to medium streams (stream orders 1–4, 2.2–10.8 m wide) were studied in the Acheron River basin in Victoria, Australia. For each confluence, two transects were sampled at each of five zones relative to the confluence: two zones upstream in the mainstem, one zone upstream in the tributary, one zone at the confluence and one zone downstream in the mainstem. Surveys were conducted in both high-flow and low-flow conditions. In mainstems, there was no change in macroinvertebrate density, taxonomic richness or functional feeding group composition downstream relative to upstream of the confluences. While tributaries statistically had distinctive benthic macroinvertebrate assemblages compared to mainstems, these distinctions were small. In low flows, densities in tributaries were substantially lower than in mainstems, but densities during high flows were more similar (albeit only about one-third as high as in low flow) in tributaries and mainstems. An inverse pattern was evident for taxonomic richness, where richness in tributaries and mainstems was similar in low flows but was greater in mainstems than in tributaries in high flows. We found little evidence of tributary effects in macroinvertebrate assemblages in this basin, which is at odds with some previous results from other continents. To explain this divergence, we suggest a conceptual model outlining factors that control variation in effects of tributaries on assemblages of benthic macroinvertebrates in mainstems.     

Trophic basis of production in tropical headwater streams,Puerto Rico: an assessment of the importance of allochthonous resources in fueling food webs

The relative importance of allochthonous and autochthonous resources in fueling tropical headwater streams remains an open topic. We combined estimates of secondary production and assessment of its trophic basis to determine which resources were responsible for animal production. We studied benthic insect assemblages in two streams in the Luquillo Experimental Forest, Puerto Rico. Habitat-weighted production estimates were similar in both streams (528.5 and 591.5 mg m⁻² year⁻¹), but production was over twice as high in pool versus riffle habitats. The mayfly Neohagenulus (Leptophlebiidae) was a major contributor to total production (259.1 and 352.2 mg m⁻² year⁻¹). All taxa relied heavily on amorphous detritus and plant tissue. Aquatic insect production was similar to that reported for shrimp assemblages in the same study area, but low relative to temperate region estimates. The trophic basis of production appears to be allochthonous organic matter, which agrees with the small size and closed canopy cover over the study streams. This is the first study quantifying the production and trophic basis of the non-shrimp macroinvertebrate assemblage in tropical island streams. We also provide support for the importance of riparian vegetation as the main energy sources for stream tropical stream food webs.

     

Beyond cool: adapting upland streams for climate change using riparian woodlands

Managed adaptation could reduce the risks of climate change to the world's ecosystems, but there have been surprisingly few practical evaluations of the options available. For example, riparian woodland is advocated widely as shade to reduce warming in temperate streams, but few studies have considered collateral effects on species composition or ecosystem functions. Here, we use cross‐sectional analyses at two scales (region and within streams) to investigate whether four types of riparian management, including those proposed to reduce potential climate change impacts, might also affect the composition, functional character, dynamics and energetic resourcing of macroinvertebrates in upland Welsh streams (UK). Riparian land use across the region had only small effects on invertebrate taxonomic composition, while stable isotope data showed how energetic resources assimilated by macroinvertebrates in all functional guilds were split roughly 50:50 between terrestrial and aquatic origins irrespective of riparian management. Nevertheless, streams draining the most extensive deciduous woodland had the greatest stocks of coarse particulate matter (CPOM) and greater numbers of ‘shredding’ detritivores. Stream‐scale investigations showed that macroinvertebrate biomass in deciduous woodland streams was around twice that in moorland streams, and lowest of all in streams draining non‐native conifers. The unexpected absence of contrasting terrestrial signals in the isotopic data implies that factors other than local land use affect the relative incorporation of allochthonous subsidies into riverine food webs. Nevertheless, our results reveal how planting deciduous riparian trees along temperate headwaters as an adaptation to climate change can modify macroinvertebrate function, increase biomass and potentially enhance resilience by increasing basal resources where cover is extensive (>60 m riparian width). We advocate greater urgency in efforts to understand the ecosystem consequences of climate change adaptation to guide future actions.     

Influences of fish on food web structure and function in mountain lakes

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Impacts of acidification on macroinvertebrate communities in streams of the western Adirondack Mountains,New York,USA

Limited stream chemistry and macroinvertebrate data indicate that acidic deposition has adversely affected benthic macroinvertebrate assemblages in numerous headwater streams of the western Adirondack Mountains of New York. No studies, however, have quantified the effects that acidic deposition and acidification may have had on resident fish and macroinvertebrate communities in streams of the region. As part of the Western Adirondack Stream Survey, water chemistry from 200 streams was sampled five times and macroinvertebrate communities were surveyed once from a subset of 36 streams in the Oswegatchie and Black River Basins during 2003–2005 and evaluated to: (a) document the effects that chronic and episodic acidification have on macroinvertebrate communities across the region, (b) define the relations between acidification and the health of affected species assemblages, and (c) assess indicators and thresholds of biological effects. Concentrations of inorganic Al in 66% of the 200 streams periodically reached concentrations toxic to acid-tolerant biota. A new acid biological assessment profile (acidBAP) index for macroinvertebrates, derived from percent mayfly richness and percent acid-tolerant taxa, was strongly correlated (R² values range from 0.58 to 0.76) with concentrations of inorganic Al, pH, ANC, and base cation surplus (BCS). The BCS and acidBAP index helped remove confounding influences of natural organic acidity and to redefine acidification-effect thresholds and biological-impact categories. AcidBAP scores indicated that macroinvertebrate communities were moderately or severely impacted by acidification in 44–56% of 36 study streams, however, additional data from randomly selected streams is needed to accurately estimate the true percentage of streams in which macroinvertebrate communities are adversely affected in this, or other, regions. As biologically relevant measures of impacts caused by acidification, both BCS and acidBAP may be useful indicators of ecosystem effects and potential recovery at the local and regional scale.     

Inter-annual patterns in macroinvertebrate communities of wilderness streams in Idaho, U.S.A.

This study examined the inter-annual variation in macroinvertebrate assemblages in six wilderness streams in central Idaho over a 6-year period (1990–1995). Benthic macroinvertebrates and associated environmental correlates were sampled during baseflow each summer. Little environmental change, as assessed using coefficients of variation (CVs) for substrate size and embeddedness, width, depth and periphyton standing crops, occurred in the streams over the period of study. There was also little temporal change in macroinvertebrate assemblages based on the relative abundance of the 10 most abundant taxa, all shredder taxa and all plecopteran taxa. CVs for individual taxa were substantially greater than those of most community measures, with rare taxa contributing 30–50% of the variation for any one stream. Frequency distributions for taxa CVs excluding rare taxa were more normally distributed. Differences in assemblage structure among streams were attributed to stream size (shift in shredder assemblages) and temperature (shift in plecopteran taxa). These data indicate a long-term (multi-year) persistence in the macroinvertebrate composition of these pristine streams, thus supporting the premise that such streams are excellent references for use in long-term biomonitoring programs.     

The Effect of Forest Type on Benthic Macroinvertebrate Structure and Ecological Function in a Pine Plantation in the North Carolina Piedmont

We examined the impact of small-scale commercial forestry on the structure and function of 6 headwater streams in the North Carolina Piedmont. During 2001–2003 terrestrial organic matter inputs, temperature, macroinvertebrate community composition and tolerance, leaf breakdown rate, and food web structure were quantified for 2 streams draining mature stands of managed loblolly pine, 2 streams draining mature hardwood forests, and 2 streams draining 3-year-old clear cuts, which had been replanted with loblolly pine. Streams in the clear-cuts and pine plantations were bordered by a 15 m hardwood buffer. Despite differences in watershed land-use, there were no significant differences in the organic matter supply or temperature between streams draining different forest types. However, algal biomass was significantly higher in clear-cut sites than forested sites, and was also higher in hardwood sites than pine sites. Streams draining the clear-cut sites contained lower macroinvertebrate richness and diversity, and fewer intolerant species, than streams draining pine and hardwood stands. Despite the differences in macroinvertebrates community composition, there was no difference among forest types in leaf-pack breakdown rates. Analysis of δ¹⁵N and δ¹³C natural abundance of functional feeding group indicated that the shredders and predators collected from streams draining clear-cuts had a δ¹⁵N value that was enriched relative to the macroinvertebrates of streams draining pine and hardwood forests. This difference in δ¹⁵N signature appears to be the result of the incorporation of riparian grass species in the clear-cuts, which have a higher δ¹⁵N, into the diet of shredders. Pine sites had similar food webs to natural hardwood sites. Our results suggest that clear-cutting changes both the trophic dynamics and macroinvertebrate composition of low-order Piedmont streams in North Carolina despite the presence of hardwood buffers. However, large differences were not found between older pine and hardwood stands, indicating rapid recovery following re-growth of forest vegetation, when hardwood buffer strips were present.     

Land use change affects macroinvertebrate community size spectrum in streams: the case of Pinus radiata plantations

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Relationship between Riparian Vegetation and Stream Benthic Communities at Three Spatial Scales

We examined the influence of riparian vegetation on macroinvertebrate community structure in streams of the Upper Thames River watershed in southwestern Ontario. Thirty-three μ-basins (129–1458 ha) were used to identify land cover variables that influenced stream macroinvertebrates. Micro-basins represented the entire drainage area of study streams and were similar in stream order (first, second) and land cover (agricultural or forest; no urban). We described the structure and composition of riparian vegetation and benthic macroinvertebrate communities at the outflow reach. The nature of the land cover was quantified for the stream network buffer (30 m) and the whole μ-basin. The objective of this study was to measure the magnitude and nature of the relationship between the riparian vegetation and benthic macroinvertebrate community at the outflow reach, stream network buffer, and whole μ-basin scales. Taxon richness (including total number of Ephemeroptera, Plecoptera, and Trichoptera taxa) and Simpson’s diversity of the macroinvertebrate community all increased with increased tree cover in the riparian zone at the outflow reach scale. Simpson’s equitability was lower with greater agricultural land cover in the stream network buffer. No relationship between the macroinvertebrate community and land cover was found at the whole μ-basin scale. Analysis of the influence of land cover on stream communities within a spatial hierarchy is important for understanding the interactions of stream ecosystems with their adjacent landscapes.     

Macroinvertebrate species and assemblages in the headwater streams of the River Tyne,northern England in relation to land cover and other environmental variables

The macroinvertebrate species and assemblages of headwater streams of the River Tyne catchment in northern England were classified and their relationship with environmental variables based on stream structure, water acidity, distance from source and land cover investigated using constrained ordination and logistic regression. Fuzzy classification of data from 322 stream sites generated five assemblages. Stream structure, quantified as an exposure index, was found to be the most important environmental variable, with water acidity also important. Distance from source and land cover had less influence on species and assemblage distribution. A considerable amount of variation in assemblage distribution was explained using a two-variable logistic regression with stream structure (exposure index) and water acidity (pH) in a template. Site structure and water acidity appeared to be related to drift, geology and topography with little anthropogenic influence. The applicability of the habitat template concept for explaining the distribution of stream macroinvertebrates is discussed.