Inferring the minimum pH of streams from macroinvertebrates using weighted averaging regression and calibration

1. Data on macroinvertebrates and stream chemistry were collected from sixty-four streams in Finland. Weighted averaging (WA) regression and calibration models were constructed to infer the minimum pH of streams from their invertebrate assemblages. The purpose was to develop an instrument for biological assessment and monitoring of stream acidification. The WA method was compared with simpler approaches, based on qualitative invertebrate data and pH tolerance limits, that are widely used.
2. Performance of the two approaches was assessed in terms of correlation between the inferred and observed minimum pH within the 'training set', and in terms of root mean squared differences (predicted – observed) (RMSEP) estimated by cross-validation or bootstrap resampling techniques. The models were further tested using independent data from the literature representative of a wide geographical range.
3. The predictive power of the WA models was reasonable (RMSEP 0.40–0.44 pH units) in the training set and consistently better than that of the tolerance limit method. In contrast to the latter, the WA models were able to infer a minimum pH above 5.5, suggesting they could detect the early stages of acidification.
4. The WA models performed better than the tolerance limit method in inferring pH from the independent literature, further demonstrating the superiority and generality of the WA approach.
5. The weighted averaging technique could be an effective and widely applicable tool for contemporary biological monitoring and assessment using aquatic invertebrates.     

Inferring the minimum pH of streams from macroinvertebrates using weighted averaging regression and calibration

1. Data on macroinvertebrates and stream chemistry were collected from sixty-four streams in Finland. Weighted averaging (WA) regression and calibration models were constructed to infer the minimum pH of streams from their invertebrate assemblages. The purpose was to develop an instrument for biological assessment and monitoring of stream acidification. The WA method was compared with simpler approaches, based on qualitative invertebrate data and pH tolerance limits, that are widely used.
2. Performance of the two approaches was assessed in terms of correlation between the inferred and observed minimum pH within the 'training set', and in terms of root mean squared differences (predicted – observed) (RMSEP) estimated by cross-validation or bootstrap resampling techniques. The models were further tested using independent data from the literature representative of a wide geographical range.
3. The predictive power of the WA models was reasonable (RMSEP 0.40–0.44 pH units) in the training set and consistently better than that of the tolerance limit method. In contrast to the latter, the WA models were able to infer a minimum pH above 5.5, suggesting they could detect the early stages of acidification.
4. The WA models performed better than the tolerance limit method in inferring pH from the independent literature, further demonstrating the superiority and generality of the WA approach.
5. The weighted averaging technique could be an effective and widely applicable tool for contemporary biological monitoring and assessment using aquatic invertebrates.     

Concordance between ecotypes and macroinvertebrate assemblages in Mediterranean streams

1. According to the guidelines of the European Water Framework Directive, assessment of the ecological quality of streams and rivers should be based on ecotype-specific reference conditions. Here, we assess two approaches for establishing a typology for Mediterranean streams: a top-down approach using environmental variables and bottom-up approach using macroinvertebrate assemblages.
2. Classification of 162 sites using environmental variables resulted in five ecotypes: (i) temporary streams; (ii) evaporite calcareous streams at medium altitude; (iii) siliceous headwater streams at high altitude; (iv) calcareous headwater streams at medium to high altitude and (v) large watercourses.
3. Macroinvertebrate communities of minimally disturbed sites ( n  = 105), grouped using UPGMA (unweighted pair-group method using arithmetic averages) on Bray–Curtis similarities, were used to validate four of the five ecotypes obtained using environmental variables; ecotype 5, large watercourses, was not included as this group had no reference sites.
4. Analysis of similarities ( anosim ) showed that macroinvertebrate assemblage composition differed among three of the four ecotypes, resulting in differences between the bottom-up and top-down classification approaches. Siliceous streams were clearly different from the other three ecotypes, evaporite and calcareous ecotypes did not show large differences in macroinvertebrate assemblages and temporary streams formed a very heterogeneous group because of large variability in salinity and hydrology.
5. This study showed that stream classification schemes based on environmental variables need to be validated using biological variables. Furthermore, our findings indicate that special attention should be given to the classification of temporary streams.     

Non-interactive fish communities in the coastal streams of North-western France

1. Spatial patterns of freshwater fish species at regional and local scales were investigated to explore the possible role of interspecific interactions in influencing distribution and abundance within communities occupying coastal streams of North-Western France.
2. Nine sites from nine streams situated in the same biogeographical region were sampled annually over the 6-year period from 1990 to 1995.
3. Similar habitats (sites) with richer regional colonization pools exhibited proportionally richer local communities in terms of number of species, total density and total biomass of individuals. Furthermore, no negative relationships were found between density and biomass of each of the most common species and local species richness.
4. Results of dynamic regression models (applied to the above-mentioned species) suggest an absence of strong competition between all pairs of species.
5. The evidence on lack of density compensation for species-poor communities and absence of perceptible interspecific competition between species suggest that the communities studied are non-interactive.
6. Two main explanations can be advanced. First, the local abundance of species in the communities studied could be determined through differential responses to unpredictable environmental changes, rather than through biological interactions. Second, as a result of historical events, the communities studied are reduced in congeneric species which can limit, in turn, the influence of interspecific competition in structuring these communities.
7. These results underline the strong influence of regional processes in shaping local riverine fish communities and minimize the possible influence of species interactions in governing these communities.     

Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Colonization and ecological development of new streams in Glacier Bay National Park, Alaska

SUMMARY 1. Colonization and ecological development of postglacial freshwater communities was investigated in Glacier Bay National Park, south-eastern Alaska, following the rapid recession of a Neo-glacial ice sheet within the last 250 years.
2. Environmental variables shown to be most significant in stream development were temperature, flow regime and sedimentation.
3. The Chironomidae (Diptera) were the pioneer invertebrate colonizers of newly emergent streams arising as meltwater from receding ice sheets and displayed a distinct pattern of succession with stream maturity.
4. Ephemeroptera and Plecoptera colonized warmer clearwater streams, but Trichoptera had a minimal role in invertebrate community development.
5. Establishment and production of salmonid fish populations in the new streams related principally to stream flow and sediment characteristics.
6. Future pathways along which the streams may develop is probably dependent on the degree of large organic debris input.
7. Stream development, structure and function are summarized including reference to theories of ecosystem development, ecological succession and community stability.     

Fish habitat preferences in large streams of southern France

1. Relationships between fish and their habitat over whole geographic regions, which are evident from studies of many streams and species, can improve understanding of lotic communities and provide reliable management tools. Nevertheless, most habitat preference studies have been based on single sites, and confined to small streams and to game species.
2. Regional habitat preference models, based on local velocity, depth and roughness, were developed for twenty-four species and their size classes commonly found in large European streams. Fish surveys were conducted in six large streams in southern France over an 8-year period. To limit the influences of habitat variables other than those studied, we estimated fish preferences within each survey and averaged this information across surveys. Preferences were fitted with confidence intervals and their sensitivity to field uncertainty was evaluated.
3. Most species and size classes had significant preferences for local habitat conditions which were consistent across the region. Habitat preferences predominant in the region overall were not always observed at any one site, but habitat conditions preferred on average in the region were never actually avoided locally. These results support the use of regional preference models for fish and the development of similar models for other lotic groups whose sensitivity to local habitat conditions has been reported elsewhere.     

Do stream macroinvertebrates use instream refugia in response to severe short-term flow reduction in New Zealand streams?

1. Demand for water is increasing and water managers need to know how much they can remove from a stream before there are significant detrimental effects on its biological integrity. Flow reduction alters a number of habitat variables known to be important to aquatic invertebrates such as depth, velocity, temperature and fine sediment accumulation. Some taxa may attempt to use instream refugia to mitigate the effects of flow reduction.
2. We experimentally manipulated flows by constructing weirs and diversions in three small New Zealand streams. Discharge was reduced by 88–96%. We tested the hypothesis that macroinvertebrates would use pools and the hyporheic zone as refugia during short-term (1-month) periods of reduced flow.
3. We sampled hyporheic invertebrates with colonization chambers and pool invertebrates with kick nets within a before-after, control-impact (BACI) experimental design. A suite of physicochemical parameters was measured concurrently including surface and hyporheic temperatures.
4. Flow reduction significantly decreased velocity (60–69%) in all streams. Depth (18–61%) and wetted width (24–31%) tended to decrease but these changes were not always significant. Sediment cover increased the most in farmland streams (10–80%). Apart from decreasing temperature range (18–26%), flow reduction had little impact on the surface water temperatures.
5. Flow reduction had no impact on the abundance of common pool macroinvertebrates or on the abundance, vertical distribution or community composition of hyporheic macroinvertebrates.
6. Our results suggest that aquatic macroinvertebrates are resistant to short-term, severe flow reduction as long as some water remains.     

The search for reference conditions for stream vegetation in northern Europe

1. The European Water Framework Directive provides a framework for improving the ecological quality of stream ecosystems, with deviation from reference used as a measure of ecological status.
2. Here we examine the possibility of using less impacted stream sites from Latvia, Lithuania and Poland to establish a Danish reference network for macrophyte assemblages, and as a guiding image for identification of possible references sites within Denmark. Both approaches were evaluated using historical Danish records.
3. Four different macrophyte assemblages were identified for mid-sized streams in the Central and Eastern Lowland ecoregions. Macrophyte assemblages could not be delineated using physical stream site characteristics; however a gradual change in assemblage composition was attributed to differences in alkalinity and human impact.
4. Assemblages of contemporary vegetation in Denmark were quite similar to those found in Polish, Latvian and Lithuanian streams (26–35%). However, more importantly, from species-based predictions we noted higher similarity, particularly with Latvian and Lithuanian streams, before intensive land use commenced in Denmark ( c. 1900). These results show that stream sites from these three countries can be considered in a Danish reference network.
5. Two of the four macrophyte assemblages comprised species such as Fontinalis antipyretica, Myriophyllum spicatum , Nuphar lutea , Potamogeton alpinus and P. perfoliatus that have a very scattered occurrence in the contemporary vegetation in Denmark. These groups were closely associated with the predictions from historic records, thereby lending support the conjecture that these assemblages could be part of the guiding image for the identification of potential reference sites within Denmark.     

Macroinvertebrate diversity in headwater streams: a review

1. Headwater streams are ubiquitous in the landscape and are important sources of water, sediments and biota for downstream reaches. They are critical sites for organic matter processing and nutrient cycling, and may be vital for maintaining the 'health' of whole river networks.
2. Macroinvertebrates are an important component of biodiversity in stream ecosystems and studies of macroinvertebrate diversity in headwater streams have mostly viewed stream systems as linear reaches rather than as networks, although the latter may be more appropriate to the study of diversity patterns in headwater systems.
3. Studies of macroinvertebrate diversity in headwater streams from around the world illustrated that taxonomic richness is highly variable among continents and regions, and studies addressing longitudinal changes in taxonomic richness of macroinvertebrates generally found highest richness in mid-order streams.
4. When stream systems are viewed as networks at the landscape-scale, α-diversity may be low in individual headwater streams but high β-diversity among headwater streams within catchments and among catchments may generate high γ-diversity.
5. Differing ability and opportunity for dispersal of macroinvertebrates, great physical habitat heterogeneity in headwater streams, and a wide range in local environmental conditions may all contribute to high β-diversity among headwater streams both within and among catchments.
6. Moving beyond linear conceptual models of stream ecosystems to consider the role that spatial structure of river networks might play in determining diversity patterns at the landscape scale is a promising avenue for future research.     

Microhabitat availability in Welsh moorland and forest streams as a determinant of macroinvertebrate distribution

SUMMARY. 1 Eighteen streams in mid-Wales were sampled for macro-invertebrates in both riffles and margins in April 1985–87. Stream macro-flora, substrata and marginal habitats were surveyed in May 1988.
2. TWINSPAN classification of the macroinvertebrate data indicated three major stream groups. One was distinguished by circumneutral pH and had a flora and fauna typical of such conditions. The other two groups consisted of acidic streams with moorland and conifer afforested catchments respectively. The forest streams were the more acidic but the two groups also differed significantly in the composition of their marginal habitats.
3 The acidic moorland streams had more vegetation ('soft' features) in the margins and supported several invertebrate taxa which were relative more abundant there than in the riffles. These taxa may be excluded from forest streams because the margins are 'hard' due to greater erosiveness and shading.
4. In view of the increasing cover by conifer afforestation in Britain, it is clearly necessary to elucidate all its effects on stream ecosystems, which include changes to the physical environment.     

Dispersal and recruitment of Tasiagma ciliata (Trichoptera: Tasimiidae) in rainforest streams, south-eastern Australia

1. This study examined genetic variation within and among populations of the caddis fly Tasiagma ciliata (Tasimiidae: Trichoptera) from rainforest streams in south-east Queensland, Australia.
2. Very low levels of genetic differentiation at large spatial scales, between subcatchments and between catchments, indicated that dispersal by the winged adults is widespread. However, significant genetic differentiation at the smallest spatial scale examined, within reaches in a single stream, suggested limited movement by larvae within streams.
3. A patchy distribution of deviations from Hardy–Weinberg equilibrium and differences in patterns among allozyme loci suggested that populations in particular reaches were the result of only a few matings.
4. These results are surprising, given the large numbers of larvae present within a single reach. We suggest that stochastic effects of recruitment may underlie much of the spatial and temporal variation in population numbers in these rainforest streams.     

The distribution of Ephemerella ignita (Ephemeroptera) in streams: the role of pH and food resources

SUMMARY. 1. The mayfly Ephemerella ignita does not occur in upland streams of the River Duddon catchment which have waters with low pH values of 4.8-5.2, negative alkalinities and low ionic contents. However, it does occur in lowland streams of the catchment which have waters with higher pH values of 6.6 and above, large positive alkalinities and high ionic contents. This study addresses the question of whether the absence of the animal in the acid water streams is due directly to the water chemistry or to the absence of suitable food.
2. In comparative experiments using unfed animals of several species it was shown that E. ignita has a considerable tolerance of low pH, low ion water. In this it is similar to Amphinemura sulcicollis (Plecoptera), a stonefly which occurs in the acid water streams of the River Duddon catchment.
3. Growth rates in laboratory experiments with E. ignita were equally good whether the food supplied was that available in the low pH water streams (bryophyte Nardia compressa plus filamentous alga Hormidium subtile) or typical of that available in high pH water streams (moss Rhynchostegium riparioides with the epiphytic diatom Cocconeis placentula). However, a consideration of the natural habits of the animal suggests that it is the inaccessibility of the H. subtile algal food, rather than its unsuitability, which is relevant to the exclusion of E. ignita from the low pH streams. In addition, the very limited diversity of other algae in the low pH water streams, in the summer season, may well be influential in inhibiting the establishment of E. ignita.
4. The comparative experiments using unfed animals suggested that, contrary to the findings for E. ignita , the mayflies Baetis muticus and Beatis rhodani are probably excluded from the acid water streams of the Upper Duddon because of water chemistry.     

Genetic differentiation among populations of Caridinazebra (Decapoda: Atyidae) in tropical rainforest streams, northern Australia

1. Caridina zebra is a common atyid shrimp in some tropical rainforest streams in far north Queensland, Australia. Genetic variation at five allozyme loci was used to estimate the level of dispersal among populations of this species, within and between stream systems. Shrimps were sampled from nine streams in the Tully River catchment and two headwater streams in the adjacent Herbert River catchment in an area under consideration for extensive hydroelectric development.
2. High levels of genetic differentiation were recorded among most populations which suggests that, like other fully aquatic species, movement is limited to a very small spatial scale.
3. In the Tully catchment, populations of shrimp from streams with confluences at high altitude showed less genetic differentiation than those from streams which directly entered the lower river. Dispersal between the latter streams is clearly limited by the presence of large waterfalls and cascades.
4. Adjacent stream populations were often highly differentiated, despite their close proximity, suggesting that overland dispersal is unlikely. However, populations of shrimp in the two streams in the Herbert catchment were strikingly similar in genetic structure to those in adjacent headwater streams of the Tully. Such similarity may reflect relatively recent changes in drainage patterns.     

Detritivores in Kenyan highland streams: more evidence for the paucity of shredders in the tropics?

1. The relationship between coarse particulate organic matter (CPOM) standing stock and benthic macroinvertebrate assemblages in Kenyan highland streams was determined by sampling seven sites on three rivers (2000–2700 m a.s.l.). Taxa recorded were allocated to functional feeding groups using published literature, mouthpart analysis and examination of gut contents. Patterns were compared with five structurally similar streams in three areas of Europe (south-west France, south-east England, north-east England).
2. Number of individuals and proportion of detritivores in Kenyan streams were equivalent to, or greater than, those in European sites. Shredders were, however, almost completely absent from Kenyan sites, despite high standing stocks of CPOM. Shredders were abundant in all European sites.
3. The phenomenon of low shredder abundance has been observed in other tropical streams in south-east Asia and Central and South America but, in contrast to these regions, the African rivers studied were devoid of shrimps or fish which may occupy the shredding niche elsewhere.
4. These preliminary data suggest that shredder-mediated detritus processing, which is a key functional component of streams in the North Temperate Zone, does not operate in East African streams. There are three possible reasons for this. The first is that tropical African rivers are functionally different to those in temperate regions. This could be because of enhanced microbial activity replacing shredder activity at high temperatures. Alternatively, it could be a result of low palatability of detrital inputs from dominant riparian trees in the region. The second and third are methodological: that our allocation to functional feeding groups is incorrect, and that our sampling methods missed a potentially key shredding taxon – the freshwater crab Potamonautes sp.     

Response of aquatic hyphomycete communities to enhanced stream retention in areas impacted by commercial forestry

1. Aquatic hyphomycetes are an important component of detritus processing in streams. Their response to enhanced stream retentiveness was tested by manipulating three streams located in Kielder Forest (northern England), a large plantation of exotic conifers, and two streams in Montagne Noire (south-west France) dominated by native broadleaf woodland. Treatment was by placement of logs or plastic litter traps into a 10–20 m stream section. Fungal spores were collected from stream water upstream and downstream of the treated sections over 1–2 years.
2. The average concentration of fungal spores in reference sections was nearly 10× greater in the French streams than in the English streams. The number of hyphomycete species was also higher in the French streams. These differences between regions were probably a consequence of the much lower standing stock and diversity of leaf litter in the English streams.
3. Despite these large regional differences, the treatment had a clear effect in all streams. Detrital standing stocks were enhanced in treated sections by up to 90% in French streams and 70% in English streams.
4. Mean spore density below treated sections increased by 1.8–14.8% in French streams and 10.2–28.9% in the naturally less retentive English streams. The number of fungal species increased significantly below the treated sections of the English streams, although not the French ones.
5. In biologically impoverished plantation streams, input of woody debris can increase detritus retention and enhance hyphomycete diversity and productivity. This may have consequent benefits for detritus processing and macroinvertebrate production.     

A method for estimating larval life spans of aseasonal aquatic insects from streams on Bougainville Island, Papua New Guinea

1. The larval life spans of twelve species of aseasonal aquatic insects (eight Trichoptera, three Ephemeroptera, one Odonata) were investigated in two tropical streams with near constant water temperatures.
2. Life spans were estimated from equations relating larval life span to the ratio between cohort production (estimated from the size–frequency technique) and annual production (estimated from regressions in the literature).
3. Life spans varied from 40 to 250 days and were positively correlated with maximum larval dry weight ( r = 0.73, P < 0.001). Ephemeroptera had the shortest larval life spans (40–110 days), Trichoptera intermediate (95–185 days) and Odonata the longest (250 days).     

A method for estimating larval life spans of aseasonal aquatic insects from streams on Bougainville Island, Papua New Guinea

1. The larval life spans of twelve species of aseasonal aquatic insects (eight Trichoptera, three Ephemeroptera, one Odonata) were investigated in two tropical streams with near constant water temperatures.
2. Life spans were estimated from equations relating larval life span to the ratio between cohort production (estimated from the size–frequency technique) and annual production (estimated from regressions in the literature).
3. Life spans varied from 40 to 250 days and were positively correlated with maximum larval dry weight ( r = 0.73, P < 0.001). Ephemeroptera had the shortest larval life spans (40–110 days), Trichoptera intermediate (95–185 days) and Odonata the longest (250 days).