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.     

Environmental factors accounting for benthic macroinvertebrate assemblage structure at the sample scale in streams subjected to a gradient of cattle grazing

Macroinvertebrate assemblages were related to environmental factors that were quantified at the sample scale in streams subjected to a gradient of cattle grazing. Environmental factors and macroinvertebrates were concurrently collected so assemblage structure could be directly related to environmental factors and the relative importance of stressors associated with cattle grazing in structuring assemblages could be assessed. Based on multivariate and inferential statistics, measures of physical habitat (% fines and substrate homogeneity) had the strongest relationships with macroinvertebrate assemblage structure. Detrital food variables (coarse benthic and fine benthic organic matter) were also associated with assemblage structure, but the relationships were never as strong as those with physical habitat measures, while autochthonous food variables (chlorophyll a and epilithic biomass) appeared to have no association with assemblage structure. The amount of variation explained in taxa composition and macroinvertebrate metrics is within values reported from studies that have examined macroinvertebrate metric–sediment relationships. The % Coleoptera and % crawlers had consistent relationships with % fines during this study, which suggests they may be useful metrics when sediment is a suspected stressor to macroinvertebrate assemblages in Blue Ridge streams. Findings from this study also demonstrate the importance of quantitative sampling through time when research goals are to identify relationships between macroinvertebrates and environmental factors.     

Changing ecology of Lake Victoria cichlids and their environment: evidence from C<Superscript>13</Superscript> and N<Superscript>15</Superscript> analyses

Springs are spatially restricted and insular ecotones. In the Alps, topography enhances this isolation. Springs are to a certain extent inhabited by organisms which are adapted to the relatively constant environmental conditions in springs. We examined thirty-six springs in eight different areas in the Swiss National Park (SNP) to understand if the macroinvertebrate assemblages of high-elevation springs are isolated or interconnected communities. We quantitatively and qualitatively sampled the macroinvertebrate assemblages of the springs and monitored environmental parameters. The similarity of the macroinvertebrate assemblages of the springs within and the dissimilarities between the different areas were relatively high; a clear spatial isolation was not evident. The differences of macroinvertebrate assemblages in different areas were explained by substrate parameters: springs in areas at lower altitudes were characterized by organic substrates and many water mite species. High-elevation springs were characterized by coarse inorganic substrates and Trichoptera of the genus Drusus. For spring conservation, it is important to decide on an individual basis if the loss of a single spring will have severe consequences for spring biodiversity.     

Land‐use legacy and the differential response of stream macroinvertebrates to multiple stressors studied using in situ experimental mesocosms

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Lentic macroinvertebrate assemblage structure along gradients in spatial heterogeneity, habitat size and water chemistry

Littoral zones of small water bodies are spatially heterogeneous habitats, harbouring diverse biotic communities. Despite this apparent heterogeneity, many studies have stressed the importance of water chemistry in determining the structure of littoral macroinvertebrate assemblages. The purpose of this study was to consider the relative importance of several spatial and water chemistry variables in explaining the patterns in the structure of macroinvertebrate assemblages in 21 lentic water bodies in northeastern Finland. Water bodies were selected to represent various habitat conditions ranging from small permanent bog ponds to small forest lakes. According to canonical correspondence analysis (CCA), the most important environmental factors related to assemblage composition were water body area, moss cover, total nitrogen and water hardness. In general, species composition in small bog ponds tended to differ from that in larger lakes with forested shoreline. Total species richness was best explained by a composite variable (PCA) describing physical habitat heterogeneity, species richness being lowest in small bog lakes with simple bottom structure and low amount of aquatic plants. Species numbers in dominant functional feeding groups were related to different environmental factors. Shredder species richness was best explained by a regression model incorporating total nitrogen and the amount of organic matter, both of which were negatively related to the number of shredder species. The number of gatherer species increased with mean substratum particle size. Scraper species richness was negatively affected by the abundance of detritus and positively affected by depth, and a model including both variables explained most of the variation. Variation in the number of predatory species was best explained by a regression model including moss cover and lake area.     

Scale-related patterns in the spatial and environmental components of stream macroinvertebrate assemblage variation

Aim We examined the relative contributions of spatial gradients and local environmental conditions to macroinvertebrate assemblages of boreal headwater streams at three hierarchical extents: bioregion, ecoregion and drainage system. We also aimed to identify the environmental variables most strongly related to assemblage structure at each study scale, and to assess how the importance of these variables is related to regional context and spatial structuring at different scales. Location Northern Finland ( 62 – 68° N, 25–32° E). Methods Variation in macroinvertebrate data was partitioned using partial canonical correspondence analysis into components explained by spatial variables (nine terms from the cubic trend surface regression), local environmental variables (15 variables) and spatially structured environmental variation. Results The strength of the relationship between assemblage structure and local environmental variables increased with decreasing spatial extent, whereas assemblage variation related to spatial variables and spatially structured environmental variation showed the opposite pattern. At the largest extents, spatial variation was related to latitudinal gradients, whereas spatial autocorrelation among neighbouring streams was the likely mechanism creating spatial structure within drainage systems. Only stream size and water acidity were consistently important in explaining assemblage structure at all study scales, while the importance of other environmental variables was more context‐dependent. Main conclusions The importance of local environmental factors in explaining macroinvertebrate assemblage structure increases with decreasing spatial extent. This scale‐related pattern is not caused solely by changes in study extent, however, but also by variable sample sizes at different regional extents. The importance of environmental gradients is context‐dependent and few factors are likely to be universally important correlates of macroinvertebrate assemblage structure. Finally, our results suggest that bioassessment should give due attention to spatial structuring of stream assemblages, because important assemblage gradients may not only be related to local factors but also to biogeographical constraints and neighbourhood dispersal processes.     

A continental‐scale analysis of fish assemblage functional structure in European rivers

The theory of traits (life‐history, ecological and biological traits) states that a species’ characteristics might enable its persistence and development in given environmental conditions. If environment is the major factor controlling functional assemblage structure, species with similar attributes are expected to inhabit a similar environment. This study uses trait states in 849 European riverine fish assemblages to analyze the influence of environment, phylogeny and biogeography on the functional structure of these assemblages. European fish assemblages were highly structured and two main syndromes (a suite of coevolved traits) were observed: 1) assemblages dominated by stenothermal intolerant individuals and 2) assemblages dominated by eurythermal, eurytopic and tolerant individuals. Temperature and stream physical structure were the two main environmental factors explaining the diversity of fish assemblage functional structures, while the influence of biogeographic factors was weak, once environment was taken into account. This suggests that, whatever the regional species pool, similar assemblage functional structures will be found in similar environmental conditions. The phylogenetic relatedness between species might also explain to some extent the associations between the species traits observed among European fish assemblages.     

The Effect of Riffle-Scale Environmental Variability on Macroinvertebrate Assemblages in a Tropical Stream

In a tropical stream (at the Soberaní;a National Park, Panama), different environmental factors were quantified in riffle habitats (water characteristics: velocity, depth, turbulence, and direction; stone characteristics: surface area, sphericity, and degree of burial; and others: substrate type, and canopy cover). Characteristics of macroinvertebrate assemblages (mean density of individuals, mean taxon richness, and cumulative taxon richness in three stones at each riffle) were related to both mean values and variability of these environmental factors at riffle scale. Macroinvertebrate density was higher in shallow, fast flowing, stony riffles, with low variability in dominant substrate type. Taxon richness was also higher in shallow riffles with loose, not buried stones, and water direction more or less parallel to the bank. Environmental variability resulted as important as mean values of environmental factors to explain variation in macroinvertebrate assemblages. This is the first study, to our knowledge, that quantifies substratum variability and demonstrates its influence on macroinvertebrate assemblages in a tropical stream.     

Factors influencing macroinvertebrate assemblages in artificial subtropical ponds of Taiwan

Macroinvertebrate assemblages and its association with environmental factors at the 11 artificial subtropical ponds of Taiwan were examined using the multivariate analysis software STATICO. The aims of the study were to determine whether spatial and seasonal variation of macroinvertebrate assemblages changed seasonally, to examine which environmental factors determined the spatial and temporal structure of maroinvertebrate assemblages, and to compare between-pond variations in the taxon composition of macroinvertebrates. Macroinvertebrates were collected seasonally by a corer and a sweep net in 2007, and 13 physical and chemical factors were measured at the same time. A total of 31 macroinvertebrate taxa were collected during the sampling period, and the most dominant taxa were Chironomidae (31.7% of total animal abundance) and Tubificidae (22.4%). STATICO identified pond size, pond depth, sediment depth, and altitude as the major abiotic factors and Bufo melanostictus (Amphibia) as the major biotic factor to influence macroinvertebrate assemblages at these ponds. These factors changed with seasonality. For example, the abundance of B. melanostictus was the most important factor during the spring but became much less important in other seasons. According to the spatial distribution patterns of macroinvertebrate assemblages, macroinvertebrates could be split into two groups based on their dispersal. The active dispersers, such as insect taxa, were strongly associated with pond size and the passive dispersers, such as non-insect taxa, were strongly associated with the pond depth and/or sediment depth. The results of this study suggested that pond size might influence macroinvertebrate assemblages through their dispersal mechanisms and that the environmental factors which influenced the macroinvertebrate assemblages most changed with seasons in this study area.     

Comparative Environmental Assessment in the Studies of Benthic Diatom,Macroinvertebrate, and Fish Communities

Community response to environmental gradients operating at hierarchical scales was assessed in studies of benthic diatoms, macroinvertebrates and fish from 44 stream sites in the New York City watershed. Hierarchical cluster analysis (TWINSPAN) of diatoms and fish partitioned the study sites into four groups, i.e., acid streams, reservoir outlets and wetland streams, large eutrophic streams, and small eutrophic streams; macroinvertebrate TWINSPAN distinguished an additional group of silty eutrophic streams. The correspondence among the three assemblage TWINSPAN groupings was moderate, ranging from 51 to 57%. The similarity across the four major group types was the highest among large eutrophic stream and acid stream assemblages, and the lowest among small eutrophic stream assemblages. Stepwise discriminant function analysis revealed that environmental factors discriminated most effectively the diatom grouping and least effectively the fish grouping. The best environmental predictors for diatom and macroinvertebrate grouping were conductance and percent surface water, while population density was most powerful in separating the fish groups. Carbaryl was the only pesticide that correlated with macroinvertebrate grouping. Partial redundancy analyses suggested a differential dependence of freshwater communities on the scale of the environmental factors to which they respond. The role of small‐scale habitat and habitatland cover/land use interaction steadily increased across the diatom, macroinvertebrate, and fish assemblages, whereas the effect of large‐scale land cover/land use declined.     

Different roles of environmental variables and spatial factors in structuring stream benthic diatom and macroinvertebrate in Yangtze River Delta,China

Exploring the relative contribution of spatial factors and environmental variables in shaping communities is of widespread interest in biodiversity conservation and environmental management. Stream communities are hierarchically regulated by environmental variables over multiple spatial scales, and the reaction of different organisms to stressors are still equivocal. We sampled both macroinvertebrates and diatom at 80 sites and additional 10 sites for macroinvertebrates, field measured and laboratory analyzed environmental variables, from the tributaries of Qiantang River, Yangtze River Delta China in 2011. We used PCNM (principal coordinates of neighbor matrices) to generate spatial predictors. We applied redundancy analysis and variation partitioning procedures to identify key spatial and environmental factors, and to quantify their relative roles in shaping diatom and macroinvertebrate assemblages. Our results demonstrated the role of spatial and environmental variables differed in shaping benthic diatom and macroinvertebrate. Diatom assemblage variations were better explained by spatial factors, however macroinvertebrate assemblage variations were better explained by environmental variables. In terms of environmental variables, catchment scale variables (e.g., land use estimators, land use diversity) played the primary role in determining the patterns of both diatom and macroinvertebrate assemblages, whereas the influence of reach-scale variables (e.g., pH, substrates, and nutrients) appeared less. However, nutrients were the stronger factors influencing benthic diatom, whereas physical habitat (e.g., substrates) played more important role than water chemistry in structuring macroinvertebrates. Our results provided more evidence to the incorporation of spatial factors interpreting spatial patterns of stream organisms, and highlighted the useful of multiple organisms and environmental variables at different spatial scales in diagnosing mechanism of stream degradation and in building a sound stream conditions monitoring program for Yangtze River Delta.     

春季长江口及其邻近海域鱼类浮游生物群落特征

根据1999年5月和2001年5月长江口及其邻近海域的鱼类浮游生物和环境调查资料,应用双向指示种分析（TWINSPAN）和典范对应分析（CCA）方法探讨了长江口及其邻近海 域鱼类浮游生物的群落特征.结果表明：采获的11 540 个鱼类浮游生物隶属11目18科32种,其中凤鲚、鳀、六丝矛尾鰕虎鱼、白氏银汉鱼、松江鲈为优势种,占总丰度的92.49％. TWINSPAN分类将鱼类浮游生物群落划分为3个类群：1）以凤鲚为优势种的河口群组;2）以 松江鲈和白氏银汉鱼为优势种的沿岸群组;3）以鳀和六丝矛尾鰕虎鱼为优势种的近海群组.CCA排序给出了各群组与环境因子的相互关系,表明影响这一海域鱼类浮游生物分布的主 要环境因子为盐度、深度、溶解氧和悬浮物.     

Benthic macroinvertebrates in Swedish streams: community structure, taxon richness, and environmental relations

Spatial scale, e.g. from the stream channel, riparian zone, and catchment to the regional and global scale is currently an important topic in running water management and bioassessment. An increased knowledge of how the biota is affected by human alterations and management measures taken at different spatial scales is critical for improving the ecological quality of running waters. However, more knowledge is needed to better understand the relationship between environmental factors at different spatial scales, assemblage structure and taxon richness of running water organisms. In this study, benthic macroinvertebrate data from 628 randomly selected streams were analysed for geographical and environmental relationships. The dataset also included 100 environmental variables, from local measures such as in-stream substratum and vegetation type, catchment vegetation and land-use, and regional variables such as latitude and longitude. Cluster analysis of the macroinvertebrate data showed a continuous gradient in taxonomic composition among the cluster groups from north to south. Both locally measured variables (e.g. water chemistry, substratum composition) and regional factors (e.g. latitude, longitude, and an ecoregional delineation) were important for explaining the variation in assemblage structure and taxon richness for stream benthic macroinvertebrates. This result is of importance when planning conservation and management measurements, implementing large-scale biomonitoring programs, and predicting how human alterations (e.g. global warming) will affect running water ecosystems.     

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

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

Associations between fish assemblages and environmental factors for Mediterranean-type rivers during summer

The structure of summer fish assemblages was examined along longitudinal gradients in 31 Mediterranean-type rivers of the middle Guadiana basin (south-west Iberian Peninsula), using data from 157 sites including small streams to deep rivers. An ordination analysis, based on 16 variables, was applied to species presence, using principal component and canonical correspondence analysis. The results for the habitat data were compared with those for the biological data using a Mantel analysis, and the agreement was highly significant. Spatial structure was considered by partitioning the total variability among the environmental and geographical variables. The fish assemblages showed longitudinal zoning during the summer, with species distributed over gradients of habitat size (depth), water quality (current and physico-chemical variables), and cover (substratum and vegetation), according to their adult size and life history. The size of the habitat that remained available in summer had the greatest biological effect, being the most important factor explaining fish species distribution and assemblage structure during this stressing season. Strictly spatial variation was low, but there was still a high residual variation. Habitat associations and life-history strategies are discussed for native and exotic species.     

Space–time interactions and invertebrate assemblage change in stream networks

Streams form hierarchical, dendritic physical networks, but relatively little is known about how this spatial structure affects community assembly. We investigated interactions between changes over time in macroinvertebrate assemblages and their distribution in space (the space–time interaction) in stream networks. Assemblages were sampled from every tributary, and every reach between tributaries, to determine effects of network position on assemblage composition, in four West Coast, South Island, New Zealand, headwater networks. Using canonical redundancy analysis, we found that macroinvertebrate assemblages were significantly spatially structured and species assemblages changed significantly between two sampling periods. The most important environmental variables (averaged over all AIC models) explaining change in assemblage composition were related to disturbance, local habitat/resources and habitat size. The lack of a significant interaction between space and time, however, indicated the spatial pattern of assemblages remained the same over time, regardless of changes in assemblage composition. Consistent spatial structuring could be the result of unchanging processes such as those arising from the universal nature of stream topology and hydrology acting both on habitat‐ and dispersal‐ related community processes. Thus, we conclude that although community assemblages changed over time, the spatial arrangement of communities could potentially be predicted from stream network topology and hydrology.     

Fishes of Indiana streams: current and historic assemblage structure

We examined Indiana fish assemblages using taxonomy and ecological categories to assess temporal shifts in community structure and recent environmental relationships. Historic (1945) and recent (1996–2007) presence/absence data were compiled by subbasin and analyzed with Nonmetric Multidimensional Scaling (NMS) ordination and by species richness. Canonical Correspondence Analysis (CCA) was used to test taxonomic identity and ecological category abundance data for explanation with recent (1996–2007) environmental variables. We found a decrease in assemblage heterogeneity for recent assemblages and an increase in the number of tolerant species per subbasin. Recent Indiana streams are dominated by tolerant fishes with generalist life history strategies and low functional variation. The use of ecological categories resulted in weaker relationships with environmental variables than analyses with taxonomic identities. Analyses using taxonomy resulted in strong assemblage explanation from stream size and flow variation, while analyses using ecological categories resulted in strong assemblage explanation from habitat variation in silt substrates and flow. Analyses of recent assemblage structure using ecological categories resulted in decreased assemblage variation among subbasins than in analyses using taxonomic identities. We found that fish assemblages of Indiana streams are structured primarily by habitat complexity and have been altered during the past 50 years through multiple disturbances including fragmentation, siltation, and species introductions.     

Macroinvertebrate succession during leaf litter breakdown in a perennial karstic river in Western Brazil

Leaf litter is a major basal resource to stream ecosystems, but few studies addressed their role in karst systems, mainly in intermittent springs and lakes. Patterns of resource use in perennial rivers are poorly known, although the input of leaf litter strongly influences macroinvertebrate assemblage structure. In this study, we evaluated the structure of macroinvertebrate assemblages along the decomposition of leaf litter in a tropical karst river, using leaf litter cages made of coarse nylon mesh (25 mm) to allow colonization by macroinvertebrates. The experiment was followed weekly for 10 weeks. The assemblages were dominated by snails (90.5% of total fauna), hyalellid amphipods, and larval chironomid midges, with highest abundances in the intermediate stages of the experiment, resulting in a gradient in assemblage structure. The large abundance of snails, which are common in other karst systems, suggest that this group may have an important role in decomposer food webs, facilitating or directly contributing to leaf breakdown. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Temporal and spatial macroinvertebrate variance compared: crucial role of CPOM in a headwater stream

The aim of the present study was to estimate the extent to which macroinvertebrate assemblages in a headwater stream are determined by environmental conditions and temporal dynamics. Six mesohabitats defined by substratum were sampled monthly throughout one year, environmental conditions at each sampling point being precisely described. Environmental variables could be truncated into two main gradients related to (a) the availability of food and space resources (CPOM) and (b) hydraulic conditions. The response of the macroinvertebrate assemblage to the environmental gradients and temporal dynamics were analyzed using GAM and (p)RDA. Twice as high portion of variance in the faunal data was attributed to environmental gradients compared to temporal dynamics. Total abundance, as well as the abundance of almost all feeding groups, was dependent on the availability of food and space resources (CPOM), while their proportions were determined by the quality of food resources driven by hydraulic conditions. Temporal dynamics was of lower importance. Our results suggest the role of CPOM be crucial in woodland headwater streams, as it greatly enhances habitat quality, serving both as a food source to dominant shredders and a space source to most macroinvertebrates.     

Is assemblage variability related to environmental variability? An answer for riverine fish

We examine the variability of riverine fish assemblages in terms of assemblage stability (i.e. variability of numbers of individuals within species over time and variability of assemblage total density), assemblage persistence, and assemblage species richness using data from a 9-yr survey of 27 sites within 18 coastal streams of North-western France. To do so, we test a hypothesized directional model for the expected relationships between environmental variability, assemblage variability, assemblage persistence, and assemblage species richness: 1) environmental variability within a given system is likely to generate variable local population size within this system, thus increasing local assemblages variability; 2) environmental variability should increase extinction rates (or, under constant colonization rates, decrease persistence), because the more population sizes vary within an assemblage, the more likely they are to become zero in some period of time; 3) assemblage variability should reduce assemblage species richness by increasing extinction rates within populations composing these assemblages. Results are compatible with our starting hypotheses and show that assemblage variability increased with environmental variability (i.e. discharge variability), that assemblage persistence decreased with environmental variability, and that species richness decreased with assemblage variability after environmental factors were controlled for. Thus, disturbance regimes, in our case, can alter the stability properties of assemblages and extrinsic determinants of assemblage variability may be an important determinant of assemblage species richness. These results have important conservation and management implications, due to the strong impact of river regulation on flow regimes.