Factors Affecting Macroinvertebrate Richness and Diversity in Portuguese Streams: a Two‐Scale Analysis

We analysed the spatial patterns in macroinvertebrate taxon richness and abundance at two scales: sampling unit and basin. We sampled 12 stream sites in three zones of Portugal, differing in climate geomorphology and water chemistry. At a sampling unit scale, substratum organic matter content, depth and the dominant size of substratum particles were correlated with numbers of taxa and individuals. We propose that the number of taxa at a small scale depends on the number of individuals, which in turn is the result of organic matter accumulation, hydrologic and substratum characteristics. The environmental parameters better explaining the large‐scale biological data were temperature, minimum size of substratum particles and pH. Regardless of the relative importance of variable types and mechanisms regulating stream invertebrates along the climatic gradient, rivers from the North and Centre appeared to be richer in taxa than the typically Mediterranean streams in the South.     

Role of transported particulate organic matter in the macroinvertebrate colonization of litter bags in streams

1. The exposure of mesh litter bags has been widely used to investigate the role of benthic macroinvertebrates in leaf processing in freshwaters. In this sense, several studies have related litter bag breakdown rates to the presence of colonizing invertebrates. A possible confounding factor in such experiments is that the litter bags trap suspended particulate organic matter that can itself attract invertebrate colonists, irrespective of the intended experimental treatment.
2. We attempted to quantify the accumulation of particulate organic matter (POM) within litter bags and to investigate its possible impact on macroinvertebrate density and richness. In seven headwater forested streams we exposed mesh bags filled either with beech leaves ( Fagus sylvatica ) or with plastic strips of an equal surface area.
3. Principal component analysis (PCA) showed that bag type and stream were the main explanatory variables for invertebrate colonization and POM accumulation within the bags. In contrast, there was little variation among sampling dates (6.4% of the total inertia).
4. The accumulated POM within the bags was substantial (up to 8.83 g ash-free dry mass (AFDM)) but highly variable among sites (mean from 0.32 to 4.58 g AFDM). At each of the seven sites, both richness and abundance of invertebrates were positively correlated with the mass of accumulated POM in bags. Macroinvertebrate colonization (notably taxon richness) was directly linked with the quantity of POM accumulated.
5. Our findings provide evidence of a potential pitfall in linking invertebrates to litter processing in mesh bags, particularly when large amounts of POM, entrained in stream flow, accumulate within the bags. An evaluation of the POM mass trapped in litter bags could account for the erratic patterns sometimes observed in their colonization by invertebrates.     

Disturbance history influences the distribution of stream invertebrates by altering microhabitat parameters: a field experiment

1. We investigated the effects of local disturbance history and several biotic and abiotic habitat parameters on the microdistribution of benthic invertebrates after an experimental disturbance in a flood‐prone German stream. 2. Bed movement patterns during a moderate flood were simulated by scouring and filling stream bed patches (area 0.49 m²) to a depth of 15–20 cm. Invertebrates were investigated using ceramic tiles as standardized substrata. After 1, 8, 22, 29, 36 and 50 days, we sampled one tile from each of 16 replicates of three bed stability treatments (scour, fill and stable controls). For each tile, we also determined water depth, near‐bed current velocity, the grain size of the substratum beneath the tile, epilithic algal biomass and standing stock of particulate organic matter (POM). 3. Shortly after disturbance, total invertebrate density, taxon richness and density of the common taxa Baetis spp. and Chironomidae were highest in stable patches. Several weeks after disturbance, by contrast, Baetis spp. and Hydropsychidae were most common in fill and Leuctra spp. in scour patches. The black fly Simulium spp. was most abundant in fill patches from the first day onwards. Community evenness was highest in scour patches during the entire study. 4. Local disturbance history also influenced algal biomass and POM standing stock at the beginning of the experiment, and water depth, current velocity and substratum grain size throughout the experiment. Scouring mainly exposed finer substrata and caused local depressions in the stream bed characterized by slower near‐bed current velocity. Algal biomass was higher in stable and scour patches and POM was highest in scour patches. In turn, all five common invertebrate taxa were frequently correlated with one or two of these habitat parameters. 5. Our results suggest that several ‘direct’ initial effects of local disturbance history on the invertebrates were subsequently replaced by ‘indirect’ effects of disturbance history (via disturbance‐induced changes in habitat parameters such as current velocity or food).     

Influences of habitat complexity on the diversity and abundance of epiphytic invertebrates on plants

SUMMARY 1. The compound influence of habitat complexity and patch size on stream invertebrate assemblages associated with submerged macrophytes was investigated through field sampling of two natural macrophyte species with contrasting leaf morphologies (complex, Ranunculus yezoensis; simple, Sparganium emersum) and an experiment with two artificial plants with different levels of morphological complexity. 2. The artificial plant experiment was designed to separate the effects of habitat area (patch size) and habitat complexity, thus enabling a more rigorous assessment of complexity per se than in previous studies where only a single patch size was used. Simple and complex artificial plants were established with five different patch sizes corresponding to the range found in natural plants. 3. Invertebrates occurred on both complex and simple forms of natural and artificial plants at similar abundances with dipterans and ephemeropterans being predominant. Taxon richness was higher on structurally complex Ranunculus than on simple Sparganium and was similarly higher on the complex artificial plant than on the simple one, over the entire range of habitat patch sizes. Thus, architectural complexity affected the taxon richness of epiphytic invertebrates, independently of habitat scale. 4. On the natural plants there was no difference in the abundance (both number of individuals and biomass) of invertebrates between simple and complex forms, while on artificial plants more invertebrates occurred on complex than on simple forms. The amount of particulate organic matter, >225 μm (POM) and chlorophyll a showed mixed patterns on natural and artificial plants, suggesting that the availability of these resources is not an overriding proximate factor controlling invertebrate abundance on plants. The difficulty of extrapolating from experimental results involving use of artificial plants is discussed, especially when considering the relationship between habitat structure and the occurrence of epiphytic invertebrates on natural plants.     

The influence of chemistry and habitat features on the microcrustacea of some upland Welsh streams

SUMMARY. 1. Microcrustaceans were sampled during September 1990 from three microhabitats (margins, stony riffles and gravel) at thirteen stream sites in mid-Wales.
2. Patterns in the distribution, abundance and community structure of microcrustacean taxa were related to stream chemistry, physiography, substratum composition and marginal habitat structure. Patterns of distribution between microhabitats were also assessed.
3. Chemical variables were the strongest correlates with microcrustacean parameters. Total numbers of microcrustaceans and harpacticoid copepods were reduced at sites with high aluminium concentrations, and ostracods were scarce or absent at the most acidic sites. In contrast, cyclopoid copepods were more abundant and species rich at low pH.
4. Microcrustacean communities were most species rich at sites with a high percentage cover of macrophytes on the substratum. Cladoceran species richness and abundance, and the abundance of several microcrustacean species, were also positively correlated with percentage macrophyte cover.
5. Three site groupings produced by TWINSPAN classification were primarily related to percentage macrophyte cover and marginal habitat structure, with chemistry and land use also of importance.
6. Total microcrustacean abundance was highest in margins, but of all the species, only the harpacticoid Bryocamptus cuspidatus showed a strong microhabitat preference.     

Anthropogenic modification disrupts species co‐occurrence in stream invertebrates

The question of whether species co‐occurrence is random or deterministic has received considerable attention, but little is known about how anthropogenic disturbance mediates the outcomes. By combining experiments, field surveys and analysis against null models, we tested the hypothesis that anthropogenic habitat modification disrupts species co‐occurrence in stream invertebrates across spatial scales. Whereas communities in unmodified conditions were structured deterministically with significant species segregation, catchment‐scale conversion to agriculture and sediment deposition at the patch‐ or micro‐habitat scale apparently randomized species co‐occurrences. This shift from non‐random to random was mostly independent of species richness, abundance and spatial scale. Data on community‐wide life‐history traits (body size, dispersal ability and predatory habits) and beta‐diversity indicated that anthropogenic modification disrupted community assembly by affecting biotic interactions and, to a lesser extent, altering habitat heterogeneity. These data illustrate that the balance between predictable and stochastic patterns in communities can reflect anthropogenic modifications that not only transcend scales but also change the relative forces that determine species coexistence. Research into the effects of habitat modification as a key to understanding global change should extend beyond species richness and composition to include species co‐occurrence, species interactions and any functional consequences.     

Effects of land cover on sediment regime and fish assemblage structure in four southern Appalachian streams

SUMMARY 1. We examined the relationship between catchment land cover, sediment regime and fish assemblage structure in four small streams in the upper Little Tennessee River basin of North Carolina. Study streams drained similar sized catchments (17–31 km²) with different fractions of non-forested land cover. Non-forested land cover was <3% in two 'reference' streams, whereas it was 13 and 22% in two 'disturbed' streams. Land cover data were compared with sediment transport data (suspended and bedload), benthic habitat data (embeddedness, substratum composition and coverage of fines) and fishes collected in autumn 1997.
2. Suspended sediment concentration was significantly higher in disturbed streams during both baseflow and stormflow. During baseflow disturbed streams nearly always exceeded 10 nephelometric turbidity units (NTU), whereas reference streams never exceeded this threshold. The difference in suspended sediment concentration between reference and disturbed streams was more consistent at baseflow than at stormflow. Therefore, baseflow turbidity may be a useful indicator of potential stream degradation.
3. Disturbed sites had five- to nine-fold more bedload transport than reference sites. Both embeddedness and streambed instability increased with increasing non-forested land cover.
4. Relative abundance of fishes requiring clean cobble/gravel substratum for spawning was lower in disturbed streams, whereas relative abundance of mound-building cyprinids, their nest associates and fishes that excavate nests in soft sediments (centrarchids) was higher. Relative abundance of fishes spawning in benthic crevices and gravel (BC + G) declined as the proportion of non-forested land cover increased. This study supports growing evidence that human-induced sedimentation alters stream fish assemblages.     

The stability–diversity relationship in stream macroinvertebrates: influences of sampling effects and habitat complexity

1. Theory predicts that the stability of a community should increase with diversity. However, despite increasing interest in the topic, most studies have focused on aggregate community properties (e.g. biomass, productivity) in small‐scale experiments, while studies using observational field data on realistic scales to examine the relationship between diversity and compositional stability are surprisingly rare. 2. We examined the diversity–stability relationship of stream invertebrate communities based on a 4‐year data set from boreal headwater streams, using among‐year similarity in community composition (Bray–Curtis coefficient) as our measure of compositional stability. We related stability to species richness and key environmental factors that may affect the diversity–stability relationship (stream size, habitat complexity, productivity and flow variability) using simple and partial regressions. 3. In simple regressions, compositional stability was positively related to species richness, stream size, productivity and habitat complexity, but only species richness and habitat complexity were significantly related to stability in partial regressions. There was, however, a strong relationship between species richness and abundance. When abundance was controlled for through re‐sampling, stability was unrelated to species richness, indicating that sampling effects were the predominant mechanism producing the positive stability–diversity relationship. By contrast, the relationship between stability and habitat complexity (macrophyte cover) became even stronger when the influence of community abundance was controlled for. Habitat complexity is thus a key factor enhancing community stability in headwater streams.     

Reach- and catchment-scale determinants of the distribution of freshwater mussels (Bivalvia: Unionidae) in south-eastern Michigan, U.S.A.

1. We investigated the diversity and distribution of freshwater mussels at 40 sites in an agricultural catchment, the River Raisin in south‐eastern Michigan, to relate mussel assemblages and individual taxa to reach and catchment‐scale variables. Unionids were surveyed by timed searches in 100‐m reaches, and in‐stream and riparian habitat were quantified as well as flow, water chemistry and channel morphology. Land use/cover and surficial geology were determined for site subcatchments and riparian buffers. 2. Some 21 mussel species were found overall; richness ranged from 0 to 12 living species per site. From the upper to middle to lower catchment, the number of individuals, number of species, Shannon–Weaver diversity and relative abundance of intolerant unionids all declined significantly. 3. Four groupings based on overall mussel diversity and abundance were significantly related to reach‐scale habitat variables. The richest mussel assemblages were associated with sites with higher overall habitat quality, greater flow stability, less fine substratum, and lower specific conductance. 4. Stepwise multiple regressions revealed that the distribution and abundance of the total mussel assemblage, as well as the most common species, could be predicted from a combination of reach‐ and catchment‐scale variables (R² = 0.63 for total mussels, R² = 0.51–0.86 for individual species). 5. Flow stability, substratum composition and overall reach habitat quality were the most commonly identified reach‐scale variables, and measures of surficial geology were the most effective catchment‐scale variables. The spatial pattern of geology is likely to be responsible for the diversity gradient from the upper to the lower catchment. 6. Prior studies, attempting to explain mussel distributions from local habitat features alone, have found relatively weak relationships. By employing a combination of reach‐ and catchment‐scale habitat variables, this study was able to account for a substantial amount of the spatial variability in mussel distributions.     

Physico-chemistry and aquatic insects of a glacier-fed and a spring-fed alpine stream

1. Physico-chemical conditions and benthic macroinvertebrates were studied in two adjacent alpine streams in the Tyrolean Alps, Austria, for 2 years, and aquatic insect emergence was recorded for 1 year.
2. In the spring-fed system, maximum discharge and increased concentrations of suspended solids, nitrate and particulate phosphorus occurred during snowmelt in June. In the glacier-fed stream, high discharge and strong diel fluctuations in flow and concentrations of suspended solids created a harsh and unstable environment during summer. Glacial ablation, variation in groundwater inflow, and water inputs from tributaries draining calcareous rocks caused water chemistry to vary both seasonally and longitudinally in glacier-fed Rotmoosache.
3. A total of 126 aquatic or semi-aquatic invertebrate taxa were collected, 94 of which were found in the glacier-fed stream and 120 in the spring-fed stream. Chironomid abundance was 2–8 times and taxa richness 2–3 times lower in the glacier-fed stream than in the spring-fed stream, as was the number of chironomid taxa (72 versus 93 total).
4. These results broadly support the conceptual model by Milner & Petts (1994) concerning glacier-fed stream systems. However, single samples and seasonal means showed relatively high invertebrate abundance and richness, especially during winter, indicating a considerable degree of spatial and temporal variability.
5. We suggest that the seasonal shifts from harsh environmental conditions in summer to less severe conditions in autumn and a rather constant environment in winter are an important factor affecting larval development, life-history patterns and the maintenance of relatively high levels of diversity and productivity in glacier-fed streams.     

Diversity at hydrothermal vents

Aim To describe patterns of hydrothermal vent community diversity and dispersion at the intersegment scale (> 100 km). Location The area discussed is an approximately 170 km portion of the Juan de Fuca Ridge, a mid‐ocean ridge in the north‐east Pacific Ocean. Methods Samples of benthic invertebrates from hydrothermal vents on three segments of the Juan de Fuca are examined for community characteristics such as diversity, abundance and distribution. Results Species richness (55 species) and evenness are low. If the macrofauna only are considered, species richness is about 30% lower than when meiofauna are also considered. The geometric series describes the species‐abundance distribution. The relationship between vent species’ distribution and abundance is significantly positive (r² = 0.818; P < 0.001). Alpha diversity is lower in patchy habitat than continuous habitat and gamma diversity is similar for both habitat types. Beta diversity is higher in patchy habitat. Local diversity is linearly related to regional diversity. Main conclusions Species richness is comparable to other highly disturbed systems. The geometric series species abundance model implies some degree of niche pre‐emption in the vent community and is consistent with the suggestion that the geometric series distribution can be found in species‐poor environments that experience harsh conditions and are structured by relatively few environmental factors. Species distribution and abundance are highly correlated. The regional species pool affects local vent diversity. Vent diversity studies should be conducted on at least the ridge scale.     

Colonisation of Heliconia caribaea by aquatic invertebrates: resource and microsite characteristics

Abstract.  1. Colonisation of ephemeral aquatic habitats via oviposition by invertebrates may be influenced by a variety of factors, such as the quality of aquatic habitat and the characteristics of the surrounding terrestrial environment. The water-holding bracts of Heliconia caribaea , a subtropical herb that produces ephemeral aquatic habitats, are colonised by a variety of aquatic invertebrates. To date, no experiments have been conducted to identify the cues that affect colonisation patterns via oviposition selection in Heliconia .
2. Artificial bracts were used to assess the influence of two types of resources found in bracts (plant produced carbohydrates and terrestrial snail faeces) on oviposition site-selection by invertebrate taxa via a replicated factorial design at four locations in the Luquillo Experimental Forest of Puerto Rico, U.S.A. Eleven microsite characteristics thought to affect oviposition were measured for each experimental container.
3. Most taxa responded in a minor way to microsite characteristics, whereas site selection by the most numerically dominant groups (e.g. Syrphidae) were influenced principally by resources within artificial bracts. Overall, the greatest response by particular taxa was to the presence of snail faeces. At the community level, total abundance, richness, and evenness of invertebrates increased with increasing biomass of faeces. Variation in sugar produced a more complex response.
4. In general, the terrestrial matrix surrounding these aquatic habitats was only a secondary determinant of population and community attributes; the principal factor affecting site selection was the quality of the aquatic habitat.     

Effects of prey dispersal on predator–prey interactions in streams

1. We studied the effect of mesh size (6 and 3 mm) on interactions between brown trout ( Salmo trutta ) and benthic invertebrates in enclosures placed in a stream in southern Sweden. We also compared how different prey exchange rates affected interactions between trout and invertebrates.
2. Trout had strong impacts on some benthic taxa, and different mesh sizes produced different patterns. Trout affected the abundance of 10 of the 21 taxa examined, six in enclosures with 3 mm mesh and six in enclosures with 6 mm mesh. The abundance of nine of the prey taxa was lower in the presence of trout, only leptocerids were more numerous in the presence of trout.
3. Our measurements of prey immigration/emigration, together with trout diet data, suggest that direct consumption by trout, rather than avoidance behaviour by prey, explains most decreases in prey abundance. There was avoidance behaviour by only two of the twenty-one prey taxa, with trout inducing emigration of the mayflies Baetis rhodani and Paraleptophlebia sp.
4. Trout indirectly increased periphyton biomass in both 3 and 6 mm enclosures. The effect of trout on periphyton was probably due to strong effects of trout on the grazer, Baetis rhodani , Heptagenia sp. and Paralepthoplebia sp.
5. Our results suggest that mesh size, through its effects on exchange rates of prey, may affect interactions between predators and prey in running waters, but that the effects of dispersal and predation on invertebrates are taxon specific.     

Assessing the influence of water and substratum quality on benthic macroinvertebrate communities in a metal-polluted stream: an experimental approach

SUMMARY 1. Field and laboratory experiments were conducted to assess the relative influence of water quality and substratum quality on benthic macroinvertebrate communities in the Animas River, a metal-polluted stream in south-western Colorado (U.S.A.).
2. A community-level in situ toxicity test measured direct effects of Animas River water on benthic invertebrates collected from a reference stream (Elk Creek). The effects of metal-contaminated biofilm were examined by comparing macroinvertebrate colonisation of clean and contaminated substrata placed in Elk Creek. A feeding experiment with the mayfly Baetis tricaudatus Dodds (Ephemeroptera: Baetidae) examined metal bioaccumulation and effects of metal-contaminated biofilm on growth and survival.
3. Animas River water was acutely toxic to most taxa, with greatest effects observed on mayflies (Heptageniidae, Ephemerellidae) and stoneflies (Taeniopterygidae and Capniidae).
4. Although Animas River biofilm was characterised by high concentrations of metals and low algal biomass, most taxa colonised substratum from the reference stream and the Animas River equally. The exceptions were Ephemerellidae, Taeniopterygidae and Simuliidae, which were less abundant on Animas River substratum. Mayflies grazing Animas River biofilm accumulated significantly more metals and showed reduced growth compared with organisms feeding on Elk Creek biofilm.
5. Results of our experiments demonstrated that effects of heavy metals on benthic community structure in the Animas River were complex, and that responses to metals in water and contaminated substratum were species-specific. Predicting recovery of benthic communities following remediation requires an understanding of these species-specific responses.     

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.     

Stream channel geomorphology influences mussel abundance in southern Appalachian streams, U.S.A.

1. Studies of North American streams have shown that hydraulic parameters and stream geomorphology can explain unionid mussel abundance at both the reach and catchment scale. However, few studies have examined applicability of hydrogeomorphic variables across broader spatial scales, such as across whole catchments, or have elucidated conditions under which spates can affect mussel populations in streams. 2. We quantified freshwater mussel abundance and species richness and their physical habitat at 24 sites in eight streams in southern Appalachian catchments in 2000 and 2001. In addition, we modelled site‐specific hydraulic parameters during summer baseflow and bankfull stages to estimate high‐ and low‐discharge conditions, respectively. 3. Mussel abundance was related to stream geomorphology, whereas richness was related to stream size. Baseflow habitat parameters explained only minor variation in abundance or richness, and both measures were highly correlated with mean current velocity or stream size. Bankfull shear stress composed a relatively low proportion of overall mussel habitat variability, but it accounted for significant variation in abundance and richness. 4. Mussel abundance was highly variable at sites subject to low‐shear stress during spates, whereas abundance always was low at sites subject to high‐shear stress. These data suggest that habitat conditions during floods, rather than those at summer baseflow, limit the abundance of mussels in Appalachian streams. These data also suggest that mussel abundance and assemblage structure may be sensitive to any changes in channel geomorphology and hydraulic conditions that might result from land use in the catchment.     

Disentangling the effects of local and regional processes on biodiversity patterns through taxon‐contingent metacommunity network analysis

Metacommunity theory, which has gained a central position in ecology, accounts for the role of migration in patterns of diversity among communities at different scales. Community isolation has a main role in this theory, but is difficult to estimate empirically, partly due to the taxon‐dependent nature of dispersal. Landscapes could be perceived as either fragmented or connected for organisms with contrasting dispersal abilities. Indeed, the dispersal ability of a taxon, and the spatial scale at which eco‐evolutionary processes shape local diversity, determine a taxon‐dependent metacommunity network. In this paper, we introduce a methodology using graph theory to define this taxon‐dependent metacommunity network and then to estimate the isolation of local communities. We analyzed the relative importance of local conditions versus community isolation as determinants of community richness for 25 taxa inhabiting 18 temporary ponds. Although local factors have been the foci of most previous empirical and theoretical considerations, we demonstrate that the metacommunity network is an equally important contributor to local diversity. We also found that the relative effect of local conditions and the metacommunity network depend on body size and taxon abundance. Local diversity of larger species was more affected by patch isolation, while taxon abundances were associated with positive or negative effects of isolation. Our results provide empirical support for the proposed role of metacommunity networks as determinants of community diversity and show the taxon‐dependent nature of these networks.     

Trends of macroinvertebrate community structure in glacier-fed rivers in relation to environmental conditions: a synthesis

1. Generalized additive models (GAMs) were used to predict macroinvertebrate taxonomic richness and individual taxon diversity at the reach level across seven European glacier-fed river sites from a set of 11 environmental variables. Maximum water temperature and channel stability were found to explain the most deviance in these models.
2. Using this information, and data from other recent studies of glacier-fed rivers, a modified conceptual model based on Milner & Petts (1994) is presented which predicts the occurrence of macroinvertebrate families and subfamilies as determined by maximum water temperature ( T _max) and channel stability. This deterministic model only applies to the summer meltwater period when abiotic variables drive community structure.
3. Where maximum water temperature is below 2 °C, Diamesinae chironomids are typically the sole inhabitants, but where T _max >2 °C but <4 °C Orthocladiinae are found and, where channels are more stable, Tipulidae and Oligochaeta also occur. Above 4 °C Perlodidae, Taeniopterygidae, Baetidae, Simuliidae and Empididae can be expected to be part of the glacier-fed river community, particularly in Europe.
4. At other times of the year when environmental conditions ameloriate, glacial rivers support higher macroinvertebrate abundance and diversity, with a number of taxa present that are not found during the summer melt period.
5. Dispersal constraints influence macroinvertebrate assemblages of many glacier-fed rivers located on islands and in some alpine areas.     

Stream community structure in relation to spatial variation: the influence of mesohabitat characteristics

Community structure of benthic macroinvertebrates was studied in six first- through fourth-order streams in northeast France, to elucidate changes in richness, abundance, diversity and evenness of mesohabitat assemblages as a function of environmental conditions. Patch samples were subjected to multivariate analyses to determine: (i) relationships among seven indices describing community structure (structure parameters); (ii) relationships among seven environmental variables; (iii) the relationship between community structure and environmental characteristics of patches. Faunal data showed that indices measuring the distribution of individuals among taxa (evenness, dominance) and richness are prominent in describing the structure of macroinvertebrate communities of mesohabitats. The analysis of environmental data demonstrated a major differentiating ability of current velocity and strong inter-relations among in-stream hydraulic-dependent parameters in structuring the mesohabitat environment. The co-structure (= relationship) between community organization and environmental variables indicated that substrate may be a primary determinant of community structure. Current velocity and water depth emerged as secondary factors. Trends in community structure were closely related to the spatial variability of mesohabitats. Species richness increased with habitat heterogeneity. Total abundance increased with trophic potentialities of patches. Equitability and diversity seemed to increase with patch stability. This revised version was published online in July 2006 with corrections to the Cover Date.