Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

Landscape variables influence taxonomic and trait composition of insect assemblages in Neotropical savanna streams

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Taxonomic and biological trait differences of stream macroinvertebrate communities between mediterranean and temperate regions: implications for future climatic scenarios

Streams in mediterranean regions have highly seasonal discharge patterns, with predictable torrential floods and severe droughts. In contrast, discharge is less variable in temperate regions and intermittent flow conditions are uncommon. Hydroclimatic models predict that climate change would increase frequency and severity of floods and droughts across Europe, thus increasing the proportion of streams with mediterranean characteristics in actually temperate areas. Correspondingly, understanding actual ecological differences between mediterranean and temperate streams may help to anticipate large‐scale ecological impacts of climate change. Given that large‐scale factors determine local community composition, we hypothesized that climatic differences between mediterranean and temperate regions should affect the taxonomic and biological trait composition in streams. We assembled the abundance of stream macroinvertebrate genera of 265 sites each from the Mediterranean Basin and from temperate Europe and linked these abundances to published information on 61 categories of 11 biological traits reflecting the potential of resilience from and resistance to disturbances. Although regional taxonomic richness was higher in the mediterranean than in the temperate region, local taxonomic richness and diversity did not significantly differ between regions. Local trait richness and diversity were significantly higher in the mediterranean region. Both local taxonomic and trait‐community composition differed between regions, but the former varied much more than the latter, highlighting that climate change could produce large changes in the taxonomic but rather weak changes in the trait composition. The mediterranean region was characterized by macroinvertebrates with higher dispersion and colonization capabilities, suggesting that species loss in the temperate region, by extinction or northward emigration of taxa, would be compensated for by immigration of southern mediterranean taxa. Thus, climate change would likely have stronger implications for the local conservation of taxa than for the trait composition of stream macroinvertebrate communities.     

Habitat preferences,seasonal abundance and diets of rodents in Alage,Southern Ethiopia

To understand habitat preferences, seasonal abundance and diets of rodents in wet and dry season surveys were conducted in Alage, Southern Ethiopia. Sherman and snap traps were used to capture rodents from the four habitats: bushland, Acacia woodland, maize and wheat farmlands. A total of 3312 trap nights, from the four trapping habitats, yielded 776 individuals that represented 11 species of rodents. The distribution of rodents varied between habitats and seasons. Wet season rodent abundance was 52.3% while in the dry season it was 47.7%. Seasonal differences in species abundance were insignificant. Bushland habitat had high wet and dry season abundances with 137 and 211 individuals, respectively. Abundance was low in maize farm (57 individuals) in the wet season and wheat farm (10 individuals) in the dry season. Stomach content composition analysis of snap‐trapped rodents from different habitats showed differences between species and across seasons. Six rodent species were recorded as pests on the farmlands in this study area. In conclusion, variation in habitat preferences and diet of rodents in different habitats and across seasons might be due to the role of ground cover and food sources.     

Environmental predictability of taxonomic and functional community composition in high‐latitude streams

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Macroinvertebrate communities in streams with contrasting water sources in the Japanese Alps

Alpine streams are typically fed from a range of water sources including glacial meltwater, snowmelt, groundwater flow, and surface rainfall runoff. These contributions are projected to shift with climate change, particularly in the Japanese Alps where snow is expected to decrease, but rainfall events increase. The overarching aim of the study was to understand the key variables driving macroinvertebrate community composition in groundwater and snowmelt‐fed streams (n = 6) in the Kamikochi region of the northern Japanese Alps (April–December 2017). Macroinvertebrate abundance, species richness, and diversity were not significantly different between the two stream types. Community structure, however, was different between groundwater and snowmelt‐fed streams with macroinvertebrate taxa specialized for the environmental conditions present in each system. Temporal variation in the abundance, species richness, and diversity of macroinvertebrate communities was also significantly different between groundwater and snowmelt streams over the study period, with snowmelt streams exhibiting far higher levels of variation. Two snowmelt streams considered perennial proved to be intermittent with periodic drying of the streambed, but the macroinvertebrates in these systems rebounded rapidly after flows resumed with no reduction in taxonomic diversity. These same streams, nevertheless, showed a major reduction in diversity and abundance following periods of high flow, indicating floods rather than periodic drying was a major driver of community structure. This conclusion was also supported from functional analyses, which showed that the more variable snowmelt streams were characterized by taxa with resistant, rather than resilient, life‐history traits. The findings demonstrate the potential for significant turnover in species composition with changing environmental conditions in Japanese alpine stream systems, with groundwater‐fed streams potentially more resilient to future changes in comparison to snowmelt‐fed streams.     

Seasonal flow variation allows 'time-sharing' by disparate aquatic insect communities in montane desert streams

1. Flow variation can drive major abiotic changes in stream environments between seasons. Theoretically, disparate biotic communities could be maintained during different seasons at a single site if suitable refuges and colonist sources were available. Using isolated montane desert streams in south‐east Arizona as a model system, we hypothesised that two disparate aquatic insect faunas (montane temperate and neotropical) could be maintained at the same sites through strong seasonal variation in abiotic conditions. 2. We collected aquatic insects representing 59 families from seven streams during high‐flow (March–April) and low‐flow (June) sampling periods across two years. We assessed changes in aquatic insect community and functional feeding group composition by habitat (riffle, pool) and season (high flow, low flow). 3. Within sites, wetted stream area decreased by an average of 97% between high‐flow (predominately riffles) and low‐flow (predominately pools) seasons. Community composition likewise showed strong seasonal patterns; the montane temperate fauna was strongly associated with the high‐flow season while neotropical hemipterans and coleopterans were associated with the low‐flow season. Increased water temperature was significantly associated with this shift from temperate to neotropical assemblages. 4. Functional feeding group composition shifted dramatically by season. The proportion of predators increased from 24.5% (high flow) to 75.2% (low flow) while collector–filterers and shredders declined from 38.4% (high flow) to 1.7% (low flow). 5. We suggest that habitat ‘time‐sharing’ by disparate communities is facilitated via strong seasonal variation in temperature and flow and the presence of high elevation refuges or diapause stages for temperate montane taxa to survive the dry season.     

Macroinvertebrate community structure and biological traits related to flow permanence in a Mediterranean river network

In Mediterranean climate areas, the great seasonal variability in temperature and rainfall is considered to be an evolutionary pressure that constrains plant and animal communities and their biological traits. Droughts alter habitat availability (changes of flow alter riffle-pool sequences), although habitat characteristics may also exacerbate drought to some extent. Using a simple quantitative index based on the proportion of conglomerate bedrock versus gravel and cobbles, pools versus riffles and winter versus summer flow, we show how habitat characteristics (in terms of substratum and flow) may influence the permanency of a stream site and how flow permanence constrains macroinvertebrate community structure and biological traits. Annual and seasonal macroinvertebrate richness, and the EPT (Ephemeroptera, Plecoptera and Trichoptera) and OCH (Odonata, Coleoptera and Hemiptera) metrics differed between permanent, intermittent and ephemeral sites, but not between permanent and intermittent sites. In contrast, distinct biological traits were observed in the three flow categories, although permanent sites presented few significant traits which was attributed to the stability of the habitat. Intermittent sites were dominated by taxa with pool-like strategies, while ephemeral sites were characterized by fauna with life-history adaptations to floods and droughts. In contrast to most traits (e.g., dissemination, reproduction, substrate relation), which were more constrained by local flow and substrate characteristics, life-cycle characteristics did not differ significantly among flow categories. This pattern can be explained by the features of the Mediterranean climate and particularly its high seasonal predictability, which serves as a large-scale filter of life-cycle traits, independently of local hydromorphological characteristics. Our findings indicate that drought is related to habitat characteristics and that local habitat variability favours organisms with certain traits, while other traits are independent of habitat variability on this scale and are probably affected by other large-scale habitat characteristics. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: S. Declerck     

Seasonal differences in population‐, ensemble‐ and community‐level responses of bats to landscape structure in Amazonia

The amount (composition) and spatial arrangement (configuration) of forest patches in fragmented landscapes influence the accessibility, as well as the abundance and diversity of resources available to bats. Moreover, tropical fruit and insect abundance differ seasonally in response to changes in precipitation, and many bats in the family Phyllostomidae employ seasonal reproductive strategies. Because reproductive activities involve constraints on time and energy as well as increased nutritional demands, foraging behavior and home range size may differ between wet and dry seasons. Nonetheless, seasonal variation in response to landscape structure by bats has not been examined previously. Consequently, population‐, ensemble‐ and assemblage‐level responses of phyllostomids to landscape composition and configuration were quantified separately during the wet and dry season at three circular focal scales (1, 3 and 5 km radii) for 14 sites in fragmented lowland Amazon forest. Responses to landscape characteristics were scale‐dependent, species‐specific, and seasonal. Abundances of frugivores responded to landscape composition in the dry season and to landscape configuration in the wet season. Conversely, abundances of animalivores responded to landscape configuration in the dry season and to landscape composition in the wet season. Divergent responses to landscape structure between seasons suggest that variation in resource abundance and diversity play a significant role in structuring population‐, ensemble‐ and assemblage‐level patterns. As such, considerations of the effects of dietary flexibility and reproductive constraints on foraging strategies and habitat use may be important when designing management plans that successfully promote long‐term persistence of biodiversity in fragmented landscapes.     

Invertebrate assemblages of pools in arid‐land streams have high functional redundancy and are resistant to severe drying

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Interannual variability in species composition explained as seasonally entrained chaos

The species composition of plankton, insect and annual plant communities may vary markedly from year to year. Such interannual variability is usually thought to be driven by year-to-year variation in weather conditions. Here we examine an alternative explanation. We studied the effects of regular seasonal forcing on a multi-species predator–prey model consisting of phytoplankton and zooplankton species. The model predicts that interannual variability in species composition can easily arise without interannual variability in external conditions. Seasonal forcing increased the probability of chaos in our model communities, but squeezed these irregular species dynamics within the seasonal cycle. As a result, the population dynamics had a peculiar character. Consistent with long-term time series of natural plankton communities, seasonal variation led to a distinct seasonal succession of species, yet the species composition varied from year to year in an irregular fashion. Our results suggest that interannual variability in species composition is an intrinsic property of multi-species communities in seasonal environments.     

Season‐modulated responses of Neotropical bats to forest fragmentation

Seasonality causes fluctuations in resource availability, affecting the presence and abundance of animal species. The impacts of these oscillations on wildlife populations can be exacerbated by habitat fragmentation. We assessed differences in bat species abundance between the wet and dry season in a fragmented landscape in the Central Amazon characterized by primary forest fragments embedded in a secondary forest matrix. We also evaluated whether the relative importance of local vegetation structure versus landscape characteristics (composition and configuration) in shaping bat abundance patterns varied between seasons. Our working hypotheses were that abundance responses are species as well as season specific, and that in the wet season, local vegetation structure is a stronger determinant of bat abundance than landscape‐scale attributes. Generalized linear mixed‐effects models in combination with hierarchical partitioning revealed that relationships between species abundances and local vegetation structure and landscape characteristics were both season specific and scale dependent. Overall, landscape characteristics were more important than local vegetation characteristics, suggesting that landscape structure is likely to play an even more important role in landscapes with higher fragment‐matrix contrast. Responses varied between frugivores and animalivores. In the dry season, frugivores responded more to compositional metrics, whereas during the wet season, local and configurational metrics were more important. Animalivores showed similar patterns in both seasons, responding to the same group of metrics in both seasons. Differences in responses likely reflect seasonal differences in the phenology of flowering and fruiting between primary and secondary forests, which affected the foraging behavior and habitat use of bats. Management actions should encompass multiscale approaches to account for the idiosyncratic responses of species to seasonal variation in resource abundance and consequently to local and landscape scale attributes.     

Effects of floods on fish assemblages in an intermittent prairie stream

1. Floods are major disturbances to stream ecosystems that can kill or displace organisms and modify habitats. Many studies have reported changes in fish assemblages after a single flood, but few studies have evaluated the importance of timing and intensity of floods on long‐term fish assemblage dynamics. 2. We used a 10‐year dataset to evaluate the effects of floods on fishes in Kings Creek, an intermittent prairie stream in north‐eastern, Kansas, U.S.A. Samples were collected seasonally at two perennial headwater sites (1995–2005) and one perennial downstream flowing site (1997–2005) allowing us to evaluate the effects of floods at different locations within a watershed. In addition, four surveys during 2003 and 2004 sampled 3–5 km of stream between the long‐term study sites to evaluate the use of intermittent reaches of this stream. 3. Because of higher discharge and bed scouring at the downstream site, we predicted that the fish assemblage would have lowered species richness and abundance following floods. In contrast, we expected increased species richness and abundance at headwater sites because floods increase stream connectivity and create the potential for colonisation from downstream reaches. 4. Akaike Information Criteria (AIC) was used to select among candidate regression models that predicted species richness and abundance based on Julian date, time since floods, season and physical habitat at each site. At the downstream site, AIC weightings suggested Julian date was the best predictor of fish assemblage structure, but no model explained >16% of the variation in species richness or community structure. Variation explained by Julian date was primarily attributed to a long‐term pattern of declining abundance of common species. At the headwater sites, there was not a single candidate model selected to predict total species abundance and assemblage structure. AIC weightings suggested variation in assemblage structure was associated with either Julian date or local habitat characteristics. 5. Fishes rapidly colonised isolated or dry habitats following floods. This was evidenced by the occurrence of fishes in intermittent reaches and the positive association between maximum daily discharge and colonisation events at both headwater sites. 6. Our study suggests floods allow dispersal into intermittent habitats with little or no downstream displacement of fishes. Movement of fishes among habitats during flooding highlights the importance of maintaining connectivity of stream networks of low to medium order prairie streams.     

Flow intermittency alters longitudinal patterns of invertebrate diversity and assemblage composition in an arid‐land stream network

1. Temporary streams comprise a large proportion of the total length of most stream networks, and the great majority of arid‐land stream networks, so it is important to understand their contribution to biotic diversity at both local and landscape scales. 2. In late winter 2010, we sampled invertebrate assemblages in 12 reaches of a large arid‐land stream network (including perennial and intermittent headwaters, intermittent middle reaches and perennial rivers) in south‐east Arizona, U.S.A. Intermittent reaches had then been flowing for c. 60 days, following a dry period of more than 450 days. We sampled a subset of the perennial study reaches three more times between 2009 and 2011. Since intermittent reaches were dry during these additional sampling periods, we used assemblage data from two other intermittent streams in the study network (sampled in 2004–05 and 2010) to explore interannual variability in intermittent stream assemblage composition. 3. Invertebrate richness was lowest in intermittent reaches, despite their often being connected to species‐rich perennial reaches. The assemblages of these intermittent reaches were not simply a subset of the species in perennial streams, but rather were dominated by a suite of stoneflies, blackflies and midges with adaptations to intermittency (e.g. egg and/or larval diapause). On average, 86% of individuals in these samples were specialists or exclusive to intermittent streams. Predators were 7–14 times more abundant in perennial than in intermittent reaches. 4. Despite being separated by long distances (12–25 km) and having very different physical characteristics, the assemblages of perennial headwaters and rivers were more similar to one another than to intervening intermittent reaches, emphasising the prime importance of local hydrology in this system. 5. The duration and recurrence intervals of dry periods, and the relative importance of dispersal from perennial refuges, probably influence the magnitude of biological differences between neighbouring perennial and temporary streams. Although perennial headwaters supported the highest diversity of invertebrates, intermittent reaches supported a number of unique or locally rare species and as such contribute to regional species diversity and should be included in conservation planning.     

Seasonal variation in the composition of fish assemblages in small Amazonian forest streams: evidence for predictable changes

1. The effects of seasonal inundation on the biology of fishes on floodplains of large Amazonian rivers are well studied. However, the small seasonal changes in headwater streams are generally considered to have little effect on fish assemblages. 2. In this study, we analysed seasonal changes in the species composition and abundance of fish in small Amazonian forest streams. We sampled fish with hand and seine nets in headwater streams in a 10 000 ha terra‐firme forest reserve near Manaus, Brazil. Each stream was surveyed at the end of the 2005 dry season, at the beginning of the 2006 rainy season and at the beginning of the 2006 dry season, by means of a standardized sampling effort. 3. The numbers of individuals and species caught were higher in the dry season, but rarefaction analyses indicated that greater species numbers could have been due simply to the larger number of individuals caught. 4. Between the dry and rainy season, the direction of changes in species composition in multivariate space varied among sites, especially for quantitative (abundance) data. However, the observed variation among sites was the less than expected if the directions of change were random. 5. Fish assemblages in the second dry season were more similar to those in the previous dry season than expected if changes in species composition among seasons were random. This indicates that a general seasonal pattern in fish assemblages can be detected, despite the existence of some erratic site‐specific changes. 6. Most of the species that showed large seasonal variations in density occupy temporary ponds during the rainy season, when much of the valley is inundated and pond networks form adjacent to streams. Short‐duration lateral migrations to these ponds may play an important role in the seasonal fish‐assemblage dynamics in Amazonian headwater streams. 7. Our results contrast with previous studies on small Amazonian streams, which have found little seasonal change in fish assemblages, and highlight the importance of the abundance of common species as an indicator of general fish assemblage structure in biological monitoring programmes.     

The sensitivity of annual grassland carbon cycling to the quantity and timing of rainfall

Global climate models predict significant changes to the rainfall regimes of the grassland biome, where C cycling is particularly sensitive to the amount and timing of precipitation. We explored the effects of both natural interannual rainfall variability and experimental rainfall additions on net C storage and loss in annual grasslands. Soil respiration and net primary productivity (NPP) were measured in treatment and control plots over four growing seasons (water years, or WYs) that varied in wet‐season length and the quantity of rainfall. In treatment plots, we increased total rainfall by 50% above ambient levels and simulated one early‐ and one late‐season storm. The early‐ and late‐season rain events significantly increased soil respiration for 2–4 weeks after wetting, while augmentation of wet‐season rainfall had no significant effect. Interannual variability in precipitation had large and significant effects on C cycling. We observed a significant positive relationship between annual rainfall and aboveground NPP across the study (P=0.01, r²=0.69). Changes in the seasonal timing of rainfall significantly affected soil respiration. Abundant rainfall late in the wet season in WY 2004, a year with average total rainfall, led to greater net ecosystem C losses due to a ～50% increase in soil respiration relative to other years. Our results suggest that C cycling in annual grasslands will be less sensitive to changes in rainfall quantity and more affected by altered seasonal timing of rainfall, with a longer or later wet season resulting in significant C losses from annual grasslands.     

A macroecological perspective of trait patterns in stream communities

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Rainfall reliability, a neglected factor in explaining convergence and divergence of plant traits in fire-prone mediterranean-climate ecosystems

Aim Rainfall reliability has been neglected as a determinant of plant trait convergence and divergence in mediterranean‐climate ecosystems. This paper reports on patterns of rainfall reliability — quantified as interannual variation in monthly and seasonal rainfall, and as the frequency of individual events in terms of their size, duration and intensity — for four fire‐prone mediterranean‐climate ecosystems. Location The four mediterranean‐climate regions of the world with fire‐prone ecosystems, namely SW Cape (South Africa), SW Australia, California and the Mediterranean Basin (Andalusia, Spain). Methods Using long‐term monthly rainfall data from stations dispersed across the four regions, we computed monthly means and interannual variation for each month of the year — the latter quantified as the coefficient of variation (CV) — and divided these into winter and summer seasons. We also computed the mean number of rainfall events, the mean frequency in various categories of event duration (days), the amount of rainfall per event (mm) and the rainfall intensity per event (mm/day) per year for winter and summer seasons for a subset of the rainfall stations. Results The fraction of rain falling in summer was lowest in California (5%) and similarly low (c. 25%) in the other three regions. The hierarchy of values of coefficient of variation (CV) of monthly rainfall during the winter period was as follows: California > Andalusia >> SW Cape > SW Australia; results for summer were: California > > Andalusia >> SW Australia ～ SW Cape. SW Australian sites experienced the greatest frequency of short, small and low‐intensity rainfall events in both seasons; patterns in the SW Cape were intermediate between Australia and the two northern hemisphere sites which both received fewer, larger and more intense events. Overall, the two southern hemisphere regions (SW Australia and the SW Cape) had significantly more reliable regimes than the two northern hemisphere ones (Mediterranean Basin and California). Main conclusions These differences in rainfall reliability regimes may provide a novel perspective on the distribution of certain plant life‐history traits in mediterranean‐climate ecosystems. Less reliable regimes would select for germination and seedling survival traits that enable persistence of genets in the face of uncertain moisture conditions during the winter and spring establishment phase. Study systems that accommodate for phylogenetic constraints, namely invasive species derived from mediterranean‐climate ecosystems, as well as shared lineages, provide good opportunities to develop and test hypotheses on the implication of different rainfall reliability regimes. One of the novel implications of this study is that the distinctive trait of assemblages in the southern hemisphere regions may be a consequence not so much of their shared nutrient‐poor soils as of their similarly reliable rainfall regimes.     

Dynamics of habitats and macroinvertebrate assemblages in rivers of the Australian dry tropics

1. The dry tropics are characterised by episodic summer rainfall such that the majority of annual river flow occurs in a short period of time. This dryland hydrological cycle leads to variably connected channels and waterholes along the length of a river bed. 2. We investigated the seasonal changes in biophysical characteristics and macroinvertebrate assemblage composition in dry‐tropics rivers at 15 sites on four rivers, each sampled five times (representing one annual hydrological cycle), in the Burdekin catchment, north Queensland, Australia. 3. Assemblages and their temporal trajectories differed among seasons, sites and habitats, even within the same habitat and/or river. Wet season flooding did not appear to ‘reset’ assemblages, with post‐wet season assemblages differing between years. 4. We found no consistent pattern in taxonomic richness over time, and sites within rivers showed no consistent convergence or divergence (i.e. turnover) in macroinvertebrate assemblage composition. However, biophysical variables associated with the rigours of the late dry season had significant effects on macroinvertebrate assemblages, highlighting the variable and often harsh conditions of dry‐tropics rivers. Underlying these patterns were different resistance and resilience traits of invertebrates (such as colonisation and establishment abilities), as well as the local‐scale effects of biophysical variables. 5. The dynamic nature of dryland rivers presents major challenges to monitoring programmes, and our results suggest a more complex scenario for monitoring and management than previously described.     

Traits of benthic macroinvertebrates in semi-natural French streams: an initial application to biomonitoring in Europe

• 1 The methods used to indicate the biological state of streams are often based on taxonomic composition, and the abundance of species or other taxa. This ‘taxonomic structure’ varies among ecoregions and cannot be applied to wider geographical areas. Therefore, we assessed the species traits of benthic macroinvertebrates from semi‐natural reference sites as a potential benchmark for large‐scale biomonitoring. Our purpose was to assess the stability of community structure, based on the representation of taxa and of traits, across large gradients of geology (sedimentary to granitic), altitude (65–1982 m), geographical coordinates (0° 48′ W to 7° 20′ E and 42° 52′ to 48° 44′ N), stream order (1–5) and slope (0.5–60‰).
• 2 We used invertebrate abundance data from the 62 most natural French stream sites available. These abundance data served to weight the occurrence of ‘biological’ traits, such as reproductive characteristics, mobility, resistance forms, food, feeding habits, respiration, and ‘ecological’ traits, such as preferences for temperature, trophic level, saprobity, biogeographic distribution, longitudinal zonation, substratum and current velocity.
• 3 Multivariate analyses of taxonomic composition demonstrated a clear site gradient from lowlands to uplands and from calcareous to granitic geology. In contrast, community structure based on both biological and ecological traits was stable across environmental gradients.
• 4 The frequency distribution of biological traits indicated that the stream benthos of the ‘reference sites’ had a mixture of categories which confirmed theoretical predictions for temporally stable and spatially variable habitats. A mixture of ecological trait categories also occurred at our reference sites. Thus, semi‐natural benthic macroinvertebrate communities are functionally diverse. Moreover, we included an initial application of these traits to a case of slightly to moderately polluted sites to show that the impact of humans significantly changes this natural functional diversity.
• 5 Future studies should focus on the potential for various biological and ecological traits to discriminate different human impacts on the benthic macroinvertebrates of running waters, and on the integration of this functional application into a general ‘reference‐condition’ approach.