A fuzzy coding approach for the analysis of long-term ecological data

• 1 We present an unconventional procedure (fuzzy coding) to structure biological and environmental information, which uses positive scores to describe the affinity of a species for different modalities (i.e. categories) of a given variable. Fuzzy coding is essential for the synthesis of long-term ecological data because it enables analysis of diverse kinds of biological information derived from a variety of sources (e.g. samples, literature).
• 2 A fuzzy coded table can be processed by correspondence analysis. An example using aquatic beetles illustrates the properties of such a fuzzy correspondence analysis. Fuzzy coded tables were used in all articles of this issue to examine relationships between spatial-temporal habitat variability and species traits, which were obtained from a long-term study of the Upper Rhône River, France.
• 3 Fuzzy correspondence analysis can be programmed with the equations given in this paper or can be performed using ADE (Environmental Data Analysis) software that has been adapted to analyse such long-term ecological data. On Macintosh Apple^TM computers, ADE performs simple linear ordination, more recently developed methods (e.g. principal component analysis with respect to instrumental variables, canonical correspondence analysis, co-inertia analysis, local and spatial analyses), and provides a graphical display of results of these and other types of analysis (e.g. biplot, mapping, modelling curves).
• 4 ADE consists of a program library that exploits the potential of the HyperCard^TM interface. ADE in an open system, which offers the user a variety of facilities to create a specific sequence of programs. The mathematical background of ADE is supported by the algebraic model known as ‘duality diagram’.

     

协惯量分析与典范对应分析在植物群落排序中的应用比较

采用协惯量分析(PCA-CA COIA)和典范对应分析(CCA)两种排序方法, 对北京小龙门林场的黄檗 (Phellodendron amurense)群落进行了分析, 并用Spearman秩相关系数检验了对应排序轴的相关性。两种排序方法得出的结果基本一致, 两者的第一排序轴都反映了海拔高度和坡向对群落分布的影响, 而各自第二、第三排序轴所代表的环境意义有所差异, 并出现了交叉, 但是两者的前3个排序轴均反映了海拔、坡位、土壤厚度和凋落物层厚度的变化趋势, 说明在环境因子个数较少或共线性效应不明显的情况下, 协惯量分析也能达到CCA的分析效果, 并且在排序轴特征值解释量上高于典范对应分析。     

Multiple co-inertia analysis: a tool for assessing synchrony in the temporal variability of aquatic communities

Multitable techniques are rarely used for investigating patterns in ecological data surveys despite their ability to deal with the spatial and/or temporal stability of assemblages. Based on a covariance optimisation criterion, Multiple Co-inertia analysis (MCOA) enables the simultaneous ordination of several tables. Such analysis allows the representation of the stable vs. unstable part of the assemblage structure in comparison to a reference derived from each table. We used MCOA on multiple time series of invertebrate sampling to show that synchrony in the temporal variability of communities can establish between geographically distant locations despite the spatial and temporal plasticity of the faunistic responses to long-term changes in environmental conditions.     

Effect of sediment quality on benthic macroinvertebrate communities in streams in the north of France

The Nord/Pas-de-Calais region located in the North of France has been submitted, for several decades, to important pollution due to intensive industrialization. The present study reports on the effects of the metal and organic content of sediments in different streams on the benthic fauna, using multivariate analyses (PCA, CA, co-inertia analysis). Benthic macroinvertebrate communities present different assemblages along the Canal à Grand Gabarit and the river Aa, both in its natural and in its canalized stretches. These differences are more specifically due to anthropic factors such as the regulation of waterways and urban and industrial contamination. Co-inertia analysis enables to bring to the light the existence of a significant co-structure between environmental and faunistic data. This revised version was published online in September 2006 with corrections to the Cover Date.     

Floristics and environmental relations in two extreme desert zones of western Egypt

• 1 The present study provides quantitative estimates of soil, vegetation structure and species distribution in 144 stands along two transects in two extreme desert zones of western Egypt. This study had two aims: (1) to describe the floristic composition of the accidental type of vegetation growing in parts of the Western Desert of Egypt; and (2) to analyse the distribution of species in relation to certain environmental factors by multivariate data analysis.
• 2 Eight environmental variables were recorded: salinity, pH, calcium carbonate, soil moisture, organic matter content, and percentages of sand, silt and clay. A total of 60 species belonging to 19 families of the angiosperms and Ephedra alata was recorded. Chamaephytes and therophytes were the most frequent, denoting a typical desert life‐form spectrum, mainly of Saharo‐Arabian distribution.
• 3 The samples were classified into eight twinspan groups. Detrended correspondence analysis (DCA) showed that these groups could be distinguished by the first two DCA axes. They could also be effectively interpreted and explained with canonical correspondence analysis (CCA). Based on forward regression, salinity, fine sediments, organic matter and soil moisture content related closely to the first three canonical axes, and accounted for 84.1% of the species–environment relationship among the stands.
• 4 Both classification and ordination resulted in a clear demonstration of the vegetation pattern in the study area. Other twinspan groups were characterized in addition to those identified previously. In conclusion, the study area has a transitional character where the Nubian Desert and the Northern Mediterranean vegetation meet.

     

Temporal and spatial environmental variability in the Upper Rhône River and its floodplain

• 1 This paper develops a framework of spatial and temporal variability for a habitat typology of the Upper Rhône River (France) and its alluvial floodplain that is based on about 17 years of data collection and analysis. The aim was to provide a scale of spatial-temporal variability for river habitat templet predictions on trends in species traits and species richness.
• 2 In developing this framework, eight physical-chemical variables were available and could be considered for twenty-two habitat types: seventeen superficial (surface) and five interstitial (0.5 m below the substrate surface). These habitat types were selected in two areas (Jons and Brégnier-Cordon) after geomorphological considerations and because of differences in their biological characteristics.
• 3 The data sets used were processed by a ‘fuzzy coding’ method using, for each variable, the frequency distribution (by modalities = categories) of all measurements and monthly means over an annual scale. Two tables were produced; the first corresponded to an expression of the total variability, and the second represented an evaluation of the temporal variability.
• 4 Each of these tables was analysed by correspondence analysis, which provided factorial scores that were used to calculate, by habitat type and by variable, a total variability and a temporal variability in terms of cumulated variability of factorial scores for the eight physical–chemical variables. The rationale in describing variability from these two tables is that total variability equals temporal variability plus spatial variability. The spatial variability was then determined by the difference between total and temporal variability. From this procedure, a positioning of the twenty-two habitat types on the spatial and temporal variability axes was obtained.
• 5 The estimate of spatial variability did not consider any error term that may have occurred in the above model; it was then tested by an independent assessment of the spatial variability using thirteen variables in nine major habitat types. A high correlation between the two ways of assessing spatial variability (r = 0.85, P < 0.004) underscored the reliability of the spatial variability that was calculated previously.
• 6 The river habitat templet obtained for the Upper Rhône and its alluvial floodplain appears to be appropriate to test the predictions on patterns of species traits and species richness in the framework of spatial and temporal variability.

     

Using multiviriate analyses for separating spatial and temporal effects within species-environment relationships

We present a multiviriate approach for the analysis of contigency tables involved in the study of species-environment relationships. The first table (species x sample) contains the abundance of p species collected in n samples. The second table (environment variables x sample) contains values for q environmental variables measured in the n samples. The third table contains the indication of where and when samples were taken.In this paper we demonstrate: (1) how to match an environmental table and a faunistic table using co-inertia analysis; (2) how to take into account a spatial effect using between-class analyses; and (3) how to combine point (1) and (2) to determine the spatial structure shared by fauna and habitat.We illustrate such an approach by using a set of hydrobiological data concerning 13 Ephemeroptera species and ten physical and chemical variables which were collected in the same site at the same dates in a small river of the Prealps.     

Theoretical habitat templets, species traits, and species richness: aquatic oligochaetes in the Upper Rhône River and its floodplain

• 1 This paper summarizes twenty years of ecological research on aquatic oligochaetes of the Upper Rhône River and its alluvial floodplain (France). Species traits of fifty species of the ninety taxa recorded from two areas Gons and Brégnier-Cordon) were used to examine the relationships among species traits, habitat utilization of these species, whether a relationship exists between species traits and habitat utilization, and the applicability of predictions from the river habitat templet and the patch dynamics concept in the framework of spatial and temporal habitat variability. We used fourteen habitat types and sixteen species traits in this analysis.
• 2 When examined by correspondence analysis, species traits separate the Naididae (with a higher potential for reproduction, small size, high mobility, and opportunistic diet) from all other families.
• 3 Habitat utilization by oligochaetes demonstrates two gradients: a vertical gradient that arranges species by their affinity for interstitial habitats (stygophily) and a transversal gradient that arranges them by their affinity for main channel habitats (rheophily).
• 4 No significant relationship was found between species traits and habitat utilization in a co-inertia analysis.
• 5 Trends observed for species traits within the framework of spatial-temporal habitat variability show only minor agreement with predictions of the river habitat templet.
• 6 Species richness is generally higher in superficial and interstitial habitats that are permanently connected with the main channel, and peaks in the superficial parapotamons (backwaters that are permanently connected with the main channel) characterized by intermediate levels of spatial as well as temporal variability; this pattern only partially fits with predictions of the patch dynamics concept.

     

Species assemblages and community structure of adult caddisflies along a headwater stream in southeastern Ghana (Insecta: Trichoptera)

Adult caddisflies were collected at 12 sites along a 5 km stretch of a forested headwater stream in southeastern Ghana in March and November 1993 and the catches are related to environmental variables. Some 34,000 specimens, belonging to 178 morphospecies in 43 genera and 11 families, were caught with Malaise traps and light traps. Many species demonstrate differences between sampling method, seasonal occurrence, and sex-ratios, with the light trap in the second wet sunny season (November) being most efficient with respect to the diversity of the catches. A Moran index analysis for global structures indicates a major transition in the fauna from the ravine with waterfalls to the forest, and secondly, a gradient through the forest. A Geary index analysis for local structures indicates further faunal turnovers in the lower reaches of the stream. Species-environment relationships were analyzed through correspondence analysis and co-inertia analysis, allowing ordination of both species and the environment into three zones. The first co-inertia axis reveals a transition between the waterfalls in the open ravine (sites 1–3), and the stream through the riverine forest (sites 4–12), while the second co-inertia axis reveals a gradient from the lotic stream in the moist semi-deciduous forest (sites 4–8) to more lentic stretches of the stream in a gradually more depleted forest (sites 9–12). The most important environmental factors related to the species transition are: riparian moss growth (r=0.94), leaves in the stream substratum (r=0.91), forest type (r=0.90), hygropetric surfaces (r=0.88), bedrock (r=0.87), lotic [riffle] (r=0.85) and lentic [pool] (r=0.85) stretches. The most important environmental factors related to the species gradient are: lotic [riffle] stretches (r=0.92), gravel (r=0.88) and leaves (r=0.85) in the stream substratum, forest type (r=0.81), and agricultural use (r=0.76). Three clusters representing three major environmental zones (zone I, II and III) along the stream are identified by projecting the average species positions on the co-inertia plane. By analyzing the Malaise trap samples significant indicator values are found for 29 species in zone I, 16 species in zone II, and 18 species in zone III. By analyzing the light trap samples significant indicator values are found for 17 species in zone I, 57 species in zone II, and 38 species in zone III. The high diversity of caddisflies in the sanctuary might be reflected both in the geological and climatological history of West Africa, as well as in the recognition of three major environmental zones, and the clean water with high diversity of stream microhabitats favorable for Trichoptera larvae. The closed forest seems to play an important role for the Trichoptera community, and the study demonstrates the importance of protecting forested headwater streams in order to maintain a sustainable aquatic biodiversity in tropical Africa.     

Theoretical habitat templets, species traits, and species richness: fish in the Upper Rhône River and its floodplain

• 1 Recent developments in ecological theory concerned with habitat templets, species assemblages, and life history traits were examined for the riverine fish communities of the Upper Rhône River, France, in the context of spatial–temporal habitat variability. Relationships among species traits, habitat utilization of species, the relationship between species traits and habitat utilization, and trends of species traits and species richness in the spatial–temporal variability of the habitat types were analysed.
• 2 Relationships among twelve species traits, and utilization of eight habitats were examined for twenty-five fish species using correspondence analysis; the relationship between species traits and habitat utilization was investigated by co-inertia analysis.
• 3 Positive relationships among species traits were observed for size, fecundity, and the number of reproductive cycles per individual. However, species were not well differentiated according to the habitat utilization, except for habitats rarely connected with the main channel (i.e. two types of oxbow lakes).
• 4 No significant relationship was found between species traits and habitat utilization, nor for either species traits or species richness when examined in the framework of spatial–temporal habitat variability. Only two species traits corresponded (with slight trends) to predictions in a river habitat templet: (i) the number of descendants per reproductive cycle increased along with temporal variability; and (ii) the number of reproductive cycles per individual was either low or high at low temporal variability and intermediate at elevated temporal variability.
• 5 The discrepancy between the predictions of the river habitat templet as well as of the patch dynamics concept and the results observed for the fish in the Upper Rhône was explained in terms of scale problems, the evolutionary ecology of the European fish fauna, and the history of the Rhône River.

     

Theoretical habitat templets, species traits, and species richness: amphibians in the Upper Rhône River and its floodplain

• 1 The purpose of this study was to test predictions of the habitat templet and the patch dynamics concept by analysing the relationship between either the species traits or species richness of amphibians and the spatial–temporal variability of eight habitat types of the Upper Rhône River and its floodplain, France.
• 2 The information on species traits of the twelve amphibian species was accumulated primarily from the literature; that on habitat utilization was based on field surveys. The information was ‘fuzzy coded’, analysed by ordination techniques, and finally linked to the spatial–temporal variability of habitat types elaborated elsewhere in this issue.
• 3 After elimination of the variance caused by the differences between urodeles and anurans, correspondence analysis of the matrix of species traits demonstrated that the flexibility of the temporal pattern of reproduction was the major source of variation among amphibian species. A less significant source of variation within anurans was related to traits usually linked to the concept of r–K selection (potential number of descendants per reproductive cycle and ratio of egg size to female size).
• 4 A correspondence analysis of habitat utilization by amphibian species separated four groups of habitat types, corresponding to different degrees of influence that the active channel had on adjacent waters in the floodplain (i.e. from habitats that are most frequently disturbed by floods, to habitats of braided sections, old oxbow lakes, and the more terrestrialized habitats).
• 5 A co-inertia analysis of the relationship between species traits and habitat utilization separated two habitat types (terrestrialized former meanders and oxbow lakes) from the others. Species of terrestrialized habitats had a reproductive display, a low number of descendants per reproductive cycle, and a high ratio of egg size to female size. Species of oxbow lakes were early or synchronous breeders, and laid a single clutch.
• 6 Neither the predictions of the habitat templet concept nor those of the patch dynamics concept were validated. Species traits were not significantly correlated to the axes of the spatial–temporal variability templet. The highest species richness was observed in habitat types with the highest temporal variability; in addition, richness did not peak at highest spatial variability.
• 7 Deviation from the predictions of the river habitat templet could be explained by: (i) the over-simplification of this model with regard to the evolution of the complex life cycles of the amphibians; and (ii) an underestimation of the importance of phylogenetic constraints and the evolution of community interactions.

     

A biotic index using benthic macroinvertebrates for Italian lakes

The relationship between benthic macroinvertebrates and environmental variables from 42 Italian lakes were analysed with the aim of developing a biotic index.In the investigated lakes, 570 species were found of which 373 belonged to the family of Chironomidae and 85 to the class of oligochaeta. Rare species, those present in less than 10 samples, were excluded from the analysis, leaving 57 species for data analysis.Multivariate analysis: canonical correlation analysis (CANON) and multivariate analysis of variance and covariance (MANCOVA) were carried out on a large database of 1060 sampling points, for which both environmental data (16 chemical and morphometric variables) and 57 species counts from soft bottom samples were available. In addition, a second dataset of 94 sites from small lakes sampled in 2005 in Northern Italy (Lombardy), was analysed for comparison.The data analysis (CANON) from the large database gave the following results:

• (1)the first canonical variate was related to conductivity, pH and alkalinity and accounted for 17% of the total variation;
• (2)the second canonical variate was related to total phosphorus, N-NH₄ and dissolved oxygen, and accounted for 15% of the total variation;
• (3)lake maximum depth, volume and water temperature were related to the third canonical variate, which accounted for 14% of the source of variation.

The analysis of the database from small lakes revealed that in the plane of the first two canonical variates dissolved oxygen was inversely correlated with the depth of sampling and that the second canonical variate showed an inverse relationship between transparency and nutrients.MANCOVA found evidence for differences between lake types and sampling years, confirmed the CANON results and allowed the quantification of the contribution of different target variables on species composition.A trophic status index was calculated ranking percentage oxygen saturation (O₂ %sat), transparency (transp) and total phosphorus (TP). The means of O₂ %sat, transp and TP weighted by the species abundances were calculated to have optimum values for each species; a benthic quality index was then obtained by multiplying the weighted means of O₂ %sat, transp and TP by species abundances and dividing the product by the total number of specimens present at each station. A significant correlation was observed between trophic status and the benthic quality index. A good agreement was also observed between the indexes calculated using the large database and the second dataset on small lakes.     

Environmental and spatial controls of taxonomic versus trait composition of stream biota

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Long‐term and contemporary environmental conditions as determinants of the species composition of bog organisms

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Fishes of the southeastern Brazil Atlantic Forest

The largest preserved areas of Atlantic Forest are located in the southeastern region of Brazil. Estuaries, mangroves, rivers and streams are home to a community of fish that is still poorly understood. The composition of fish species and their relationships with environmental conditions were used to identify homogeneous areas within a landscape of 33,000 km². A total of 6,542 specimens, representing 124 species and 13 orders of bony fishes were collected at 102 sampling stations. Resulting scores from a canonical correspondence analysis applied to biological and environmental data were used in a cluster analysis to identify fish species with similar spatial distributions and localities with similar fish fauna and abiotic conditions. Five micro-regions were identified, with homogeneous compositions of species, allowing better planning of conservation of aquatic biodiversity. Despite a high degree of preservation, impacts due to deforestation, drainage of wetlands and overfishing were detected, as well as the introduction of 41 exotic species.     

Distribution of benthic algae in the upper Illinois River basin in relation to geology and land use

• 1 Benthic‐algal distributions in the upper Illinois River basin, IL, U.S.A., were examined in relation to geology, land use, water chemistry and stream habitat using (detrended) (canonical) correspondence analysis, autecological metrics and indicator‐species analysis in order to identify the major environmental gradients influencing community variation.
• 2 Ionic composition and major nutrient [i.e. nitrogen (N) and phosphorus (P)] concentration of surface waters, salinity (Na‐Cl type), substratum type and physiognomic form of dominant species were primary factors contributing to variation in benthic‐algal assemblages of the basin. Basin geology was a significant contributing factor, but the explained variance associated with this factor was less than that related to land use.
• 3 Proportions of algal biomass consisting of cyanophytes, filamentous chlorophytes, halophilic diatoms and diatoms which utilize nitrogen heterotrophically were greater in eutrophic river segments than in less nutrient‐enriched segments. Composition of the benthic flora indicated meso‐eutrophic or eutrophic conditions throughout the basin; there were few diatoms indicative of hypertrophic waters. Shifts in diatom‐assemblage structure in response to nutrient loading provided an incomplete representation of the community‐response curve.
• 4 A weighted‐averages regression model based on total P and benthic‐algal abundances (all divisions included) yielded a highly significant correlation (r² = 0.83) between species‐inferred [WA_(tol)] and observed total P, with systematic bias (increased deviation of residuals) occurring only at concentrations greater than ～ 1.0 mg L^?1 total P. This result indicates that total P regression and calibration models can be predictable for a river basin receiving excessive loadings of phosphorus.

     

Mountain summer farms in Røldal,western Norway –; vegetation classification and patterns in species turnover and richness

This paper discusses vegetation types and diversity patterns in relation to environment and land-use at summer farms, a characteristic cultural landscape in the Norwegian mountains. Floristic data (189 taxa) were collected in 130 4-m² sample plots within 12 summer farms in Røldal, western Norway. The study was designed to sample as fully as possible the range of floristic, environmental, and land-use conditions. Vegetation types delimited by two-way indicator species analysis were consistent with results from earlier phytosociological studies. Detrended correspondence analysis and canonical correspondence analysis show that rather than being distinct vegetation types, the major floristic variation is structured along a spatial gradient from summer farm to the surrounding heathland vegetation. Species richness (alpha diversity) was modelled against environmental variables by generalized linear modelling and compositional turnover (beta diversity) by canonical correspondence analysis. Most environmental factors made significant contributions, but the spatial distance-to-farm gradient was the best predictor of both species richness and turnover. While summer farms reduce mean species richness at the plot scale, the compositional heterogeneity of the upland landscapes is increased, thereby creating ‘ecological room’ for additional vegetation types and species. Within an overall similarity across scales, soil variables (pH, base saturation, LOI, phosphate and nitrogen) differed considerably in their explanatory power for richness and turnover. A difference between ‘productivity limiting’ factors and ‘environmental sieves’ is proposed as an explanation. Species turnover with altitude is relatively low in grasslands as compared to heaths.     

Species richness and assemblage structure of Trichoptera in Danish streams

• 1 Species richness and assemblages of Trichoptera from 157 ‘pristine’ Danish lowland stream sites were analyzed, for dependence on geographical position of the sites and simple physical variables, using two way indicator species analysis and detrended correspondence analysis.
• 2 A total of 106 species were recorded, representing ≈ 90% of the species pool known from Danish streams. Only seven species occurred at more than half the sites, whereas an additional 11 species were found at more than a quarter of the sites.
• 3 Although sites showed significant regional differences in environmental variables (stream order, width, slope and presence/absence of riparian forest), species richness and assemblages were primarily correlated with stream order, width and slope. Maximum richness was found at the largest (5th order) stream sites.
• 4 Regional differences in species assemblages were found, with several species being absent from the islands Funen and Bornholm. Species assemblages also differed between forested and non‐forested stream sites.
• 5 We found evidence that stream temperature may be of only minor importance in determining Trichoptera species richness and assemblage composition in Danish streams compared to other size‐related physical factors.

     

Identification of predominant environmental factors structuring stream macroinvertebrate communities within a large agricultural catchment

• 1 Patterns of macroinvertebrate community composition were examined in streams within a 40000-km² catchment in central Michigan, U.S.A., to identify the major environmental gradients influencing community variation. Agriculture and associated clay and sandy soils predominated in much of the region.
• 2 Eighty macroinvertebrate taxa were collected from stream surveys conducted during May and August 1990. Community composition varied primarily by the proportions of Plecoptera and Ephemeroptera. Benthic communities from the heaviest agricultural zones were most different from those at other sites.
• 3 Chemical composition among the sites varied most in relation to nutrients (NH₃, NO3, PO₄). Other parameters were relatively similar. Physical characteristics of the sites were scored in six habitat categories: (i) substrate characteristics, (ii) instream cover, (iii) channel morphology, (iv) riparian zone and stream-bank conditions, (v) riffle/run quality, (vi) pool quality. Most physical habitat scores were lowest in the intense agriculture zones.
• 4 The relative importance of physical and chemical variables in explaining variation in macroinvertebrate communities was quantified using canonical correspondence analysis. Substrate characteristics were most important in both surveys. Significant correlations (P<0.05 and P<0.10) were observed between substrate quality and total numbers of Ephemeropteran, Plecopteran, and Trichopteran taxa. These relationships reflected correlations from sites in the clay soil-type region (P<0.01 and P<0.10) which contrasted with non-significant results from the less impacted, sandy soil-type region.
• 5 Effective stream restoration efforts in this region will require the alteration of local land-use activities that influence the physical habitat. Further development of empirical relationships between catchment activities and substrate and channel characteristics within natural geomorphic regions is essential for the evaluation of restoration projects.

     

Regional and local drivers of macroinvertebrate assemblages in boreal springs

Aim To identify the most important environmental drivers of benthic macroinvertebrate assemblages in boreal springs at different spatial scales, and to assess how well benthic assemblages correspond to terrestrially derived ecoregions. Location Finland. Methods Benthic invertebrates were sampled from 153 springs across four boreal ecoregions of Finland, and these data were used to analyse patterns in assemblage variation in relation to environmental factors. Species data were classified using hierarchical divisive clustering (twinspan ) and ordinated using non‐metric multidimensional scaling. The prediction success of the species and environmental data into a priori (ecoregions) and a posteriori (twinspan ) groups was compared using discriminant function analysis. Indicator species analysis was used to identify indicator taxa for both a priori and a posteriori assemblage types. Results The main patterns in assemblage clusters were related to large‐scale geographical variation in temperature. A secondary gradient in species data reflected variation in local habitat structure, particularly abundance of minerogenic spring brooks. Water chemistry variables were only weakly related to assemblage variation. Several indicator species representing southern faunistic elements in boreal springs were identified. Discriminant function analysis showed poorer success in classifying sites into ecoregions based on environmental than on species data. Similarly, when classifying springs into the twinspan groups, classification based on species data vastly outperformed that based on environmental data. Main conclusions A latitudinal zonation pattern of spring assemblages driven by regional thermal conditions is documented, closely paralleling corresponding latitudinal patterns in both terrestrial and freshwater assemblages in Fennoscandia. The importance of local‐scale environmental variables increased with decreasing spatial extent. Ecoregions provide an initial stratification scheme for the bioassessment of benthic macroinvertebrates of North European springs. Our results imply that climate warming, landscape disturbance and degradation of spring habitat pose serious threats to spring biodiversity in northern Europe, especially to its already threatened southern faunistic elements.