A multi-scale comparison of trait linkages to environmental and spatial variables in fish communities across a large freshwater lake

Species present in communities are affected by the prevailing environmental conditions, and the traits that these species display may be sensitive indicators of community responses to environmental change. However, interpretation of community responses may be confounded by environmental variation at different spatial scales. Using a hierarchical approach, we assessed the spatial and temporal variation of traits in coastal fish communities in Lake Huron over a 5-year time period (2001–2005) in response to biotic and abiotic environmental factors. The association of environmental and spatial variables with trophic, life-history, and thermal traits at two spatial scales (regional basin-scale, local site-scale) was quantified using multivariate statistics and variation partitioning. We defined these two scales (regional, local) on which to measure variation and then applied this measurement framework identically in all 5 study years. With this framework, we found that there was no change in the spatial scales of fish community traits over the course of the study, although there were small inter-annual shifts in the importance of regional basin- and local site-scale variables in determining community trait composition (e.g., life-history, trophic, and thermal). The overriding effects of regional-scale variables may be related to inter-annual variation in average summer temperature. Additionally, drivers of fish community traits were highly variable among study years, with some years dominated by environmental variation and others dominated by spatially structured variation. The influence of spatial factors on trait composition was dynamic, which suggests that spatial patterns in fish communities over large landscapes are transient. Air temperature and vegetation were significant variables in most years, underscoring the importance of future climate change and shoreline development as drivers of fish community structure. Overall, a trait-based hierarchical framework may be a useful conservation tool, as it highlights the multi-scaled interactive effect of variables over a large landscape.     

Benthic diatom communities in boreal streams: community structure in relation to environmental and spatial gradients

An important goal for community ecology is the characterization and prediction of changes in community patterns along environmental gradients. We aimed to identify the major environmental correlates of diatom distribution patterns in boreal running waters. We classified 197 stream sites based on their diatom flora. Direct ordination methods were then used to identify the key environmental determinants of this diatom-based stream typology. Finally, we tested whether a regional classification scheme based on terrestrial landscapes (ecoregions) provides a reasonable framework for a regional grouping of streams based on their diatom flora. Two-way indicator species analysis produced 13 site groups, which were primarily separated by chemical variables, mainly conductivity, total P and water colour. In partial CCA, the environmental and spatial factors accounted for 38% and 24%, respectively, of explained variation in community composition. A high proportion (almost 40%) of variation explained by the combined effect (spatially-structured environmental) indicated that diatom communities of boreal streams incorporate a strong spatial component. At the level of subecoregions, classification strength was almost equally strong for all sites as for near-pristine reference sites only. Procrustes analysis indicated that spatial factors and patterns in diatom community structure were strongly concordant. Our data support the argument that diatom communities are strongly spatially structured, with distinctly different communities in different parts of the country. Because of the strong spatial patterns of community composition, bioassessment programs utilising lotic diatoms would clearly benefit from regional stratification. A combination of regional stratification and the prediction of assemblage structure from local environmental features might provide the most robust framework for diatom-based assessment of the biological integrity of boreal streams.     

Environmental variables interact across spatial scales to structure trichopteran assemblages in Ouachita Mountain rivers

An important goal in aquatic ecology is to determine the interacting variables that regulate community structure; however, complex biotic and abiotic interactions coupled with the significance of scale have confounded the interpretation of community data. We evaluated stream and riparian habitat features in southeastern Oklahoma, USA at a range of spatial scales from local, in-stream variables to large-scale, regional characteristics to address the following questions: (1) How much variation in trichopteran community composition can be attributed to local, regional, and spatial variables? and (2) What environmental variables are most important in determining trichopteran community structure? We collected data on caddisfly community structure, local and regional environmental variables, and spatial location on the landscape from 25 sites in four rivers. We analyzed these data using canonical correspondence analysis (CCA) and variation partitioning. Our analysis explained approximately 60% of the variation in caddisfly community composition. We found that local and regional environmental variables were near equal in importance in governing caddisfly communities, with each accounting for approximately a quarter of the explained variation. Although pure spatial variables were less important, the amount of variation shared among spatial variables and local and regional variables was substantial, indicating that biogeographic history is also key to understanding caddisfly distributions. We also found a strong influence of human landuse (i.e., percent of land in agriculture, distance to roads) on caddisfly community composition. Our study indicated that communities are influenced by factors across scales, and that bioassessments should focus on not only local habitat conditions, but also incorporate larger-scale factors.     

Freshwater biodiversity at regional extent: determinants of macroinvertebrate taxonomic richness in headwater streams

We studied spatial variation of macroinvertebrate species richness in headwater streams at two spatial extents, within and across drainage systems, and assessed the relative importance of three groups of variables (local, landscape and regional) at each extent. We specifically asked whether the same variables proposed to control broad‐scale richness patterns of terrestrial organisms (temperature, topographic variability) are important determinants of species richness also in streams, or whether environmental factors effective at mainly local scales (in‐stream heterogeneity, potential productivity) constrain species richness in local communities. We used forward selection with two stopping criteria to identify the key environmental and spatial variables at each study extent. Eigenvector‐based spatial filtering was applied to evaluate spatial patterns in species richness, and variation partitioning was used to assess the amount of variation in richness attributable to purely environmental and spatial components. A prime regulator of richness variation at the bioregion extent was elevation range (increasing richness with higher topographic variability), whereas hydrological stability and temperature were unimportant. Water chemistry variables, particularly water color, exhibited strong spatially‐structured variation across drainage systems. Local environmental variables explained most of the variation in species richness at the drainage‐system extent, reflecting gradients in total phosphorus and water color (negative effect on richness). The importance of the pure spatial component was strongly region‐dependent, with a peak (60%) in one drainage system, suggesting the presence of unmeasured environmental factors. Our results emphasize the need for spatially‐explicit, regional studies to better understand geographical variation of freshwater biodiversity. Future studies need to relate species richness not only to local factors but also to broad‐scale climatic variables, recognizing the presence of spatially‐structured environmental variation.     

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.     

Spatial structure in lotic macroinvertebrate communities in England and Wales: relationship with physical, chemical and anthropogenic stress variables

We describe the relationship between macroinvertebrate community composition, the physicochemical environment and anthropogenic impacts, in running water sites across a range of water qualities in England and Wales. We have also investigated the degree of spatial structure present in both the macroinvertebrate community and the measured environment. Selected explanatory variables could account for 26% of the variation in lotic macroinvertebrate assemblage composition across England and Wales. The explanatory power of the CCA model was based predominantly on a combination of local scale variables (substrate, alkalinity, urban run-off) and regional scale variables (discharge category, northing). The physicochemical gradient associated with changes in stream type from headwaters to estuary dominated assemblage composition. The influence of pollution and habitat modification were of secondary importance. There was a substantial level of spatial structure to both the physicochemical (47% of its explanatory power spatially structured) and anthropogenic stress data (63% of its explanatory power spatially structured), which resulted in a high level of predictable spatial structuring in macroinvertebrate assemblages. Almost 40% of the variation in assemblage composition accounted for by the explanatory model exhibited spatial structure. Positive spatial autocorrelation in macroinvertebrate community composition extended to sites up to 150km apart. As a consequence, community composition could be described from northing and easting with 75% of the explanatory power of the eight physicochemical variables. Our study has confirmed the importance of the longitudinal gradient within catchments, as well as the geographical position of the catchment to macroinvertebrate communities. We have also demonstrated how quantifying the spatial structure in the dataset can improve our understanding of the factors influencing macroinvertebrate community structure.     

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

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Impacts of space,local environment and habitat connectivity on macrophyte communities in conservation lakes

Aim  To assess the relative impacts of spatial, local environmental and habitat connectivity on the structure of aquatic macrophyte communities in lakes designated for their conservation value. Location  Selected lakes of conservation importance all over Scotland, representing a wide variety of lake habitat types and associated macrophyte communities. Methods  Local environmental variables and species occurrence were measured in the field. Spatial variables were generated using principal coordinates of neighbour matrices (PCNM) analysis. Connectivity between each lake and its neighbours was defined as either (i) all lakes within a radius of 5, 10, 25, 50, 75 or 100 km; (ii) all lakes in same river system; or (iii) all lakes in the same catchment and upstream of the lake. Using variance partitioning within canonical correspondence analysis, the relative impact of E = local environment, S = space and C = lake connectivity was compared on submerged (n = 119 lakes) and emergent (n = 96 lakes) macrophyte assemblages. Results  Local environmental conditions, such as total phosphorus, alkalinity/conductivity and the presence of invasive species, as well as spatial gradients were key drivers of observed variation in macrophyte communities; e.g., for submerged macrophytes, a combination of local to moderate factors relating to water chemistry and broad‐scale gradients reflecting elevation and climate are important. Spatially structured environmental variables explained a large portion of observed variation. Main conclusions  Our findings confirmed the need to manage local environmental pressures such as eutrophication, but suggested that the traditional catchment approach was insufficient. The spatial aggregation of environmental and connectivity factors indicated that a landscape scale approach should be used in lake management to augment the risk assessment to conservation species from the deterioration of suitable lake sites over broad biogeographic areas.     

Environmental rather than spatial factors structure bacterioplankton communities in shallow lakes along a > 6000 km latitudinal gradient in South America

Metacommunity studies on lake bacterioplankton indicate the importance of environmental factors in structuring communities. Yet most of these studies cover relatively small spatial scales. We assessed the relative importance of environmental and spatial factors in shaping bacterioplankton communities across a > 6000 km latitudinal range, studying 48 shallow lowland lakes in the tropical, tropicali (isothermal subzone of the tropics) and tundra climate regions of South America using denaturing gradient gel electrophoresis. Bacterioplankton community composition (BCC) differed significantly across regions. Although a large fraction of the variation in BCC remained unexplained, the results supported a consistent significant contribution of local environmental variables and to a lesser extent spatial variables, irrespective of spatial scale. Upon correction for space, mainly biotic environmental factors significantly explained the variation in BCC. The abundance of pelagic cladocerans remained particularly significant, suggesting grazer effects on bacterioplankton communities in the studied lakes. These results confirm that bacterioplankton communities are predominantly structured by environmental factors, even over a large‐scale latitudinal gradient (6026 km), and stress the importance of including biotic variables in studies that aim to understand patterns in BCC.     

The roles of environment and space in shaping stream diatom communities

Diatoms are widely used in stream quality assessment due to their response to the local environment. Diatoms are also influenced by many large-scale processes and so the diatom communities of boreal streams incorporate a strong spatial component at a regional level. What is not properly known yet is whether the variation in diatom communities between regions is larger than the variation in measured environmental variables. We studied the roles of environment and space in accounting for variability in stream diatom communities across four regions in Finland. According to canonical correspondence analysis, geographical coordinates, nutrient concentrations (total N and P), and water conductivity were the most important factors affecting variation in diatom community composition. Of physical factors, depth and current velocity were also significant. According to Mantel tests, both environmental and geographical distances were related to dissimilarity in diatom community composition. Analysis of Similarities indicated that the regional differences in diatom community composition were larger than the regional differences in environmental variables. We also found many indicator species confined to certain regions. Our results suggest that the four study regions differ in their diatom species composition more than in their environmental features and that diatoms are structured not only by the local environment but also by large-scale processes, possibly related to history, climate and dispersal. These results imply that, while diatom species composition reflects well the environmental differences between regions, future bioassessments would benefit from regional stratification. Otherwise, relationships with environmental variables may be masked by trans-regional differences in species pools caused by the large-scale processes.     

Plant genetics shapes inquiline community structure across spatial scales

Recent research in community genetics has examined the effects of intraspecific genetic variation on species diversity in local communities. However, communities can be structured by a combination of both local and regional processes and to date, few community genetics studies have examined whether the effects of instraspecific genetic variation are consistent across levels of diversity. In this study, we ask whether host-plant genetic variation structures communities of arthropod inquilines within distinct habitat patches – rosette leaf galls on tall goldenrod ( Solidago altissima ). We found that genetic variation determined inquiline diversity at both local and regional spatial scales, but that trophic-level responses varied independently of one another. This result suggests that herbivores and predators likely respond to heritable plant traits at different spatial scales. Together, our results show that incorporating spatial scale is essential for predicting the effects of genetically variable traits on different trophic levels and levels of diversity within the communities that depend on host plants.     

Environmental and spatial correlates of community composition,richness and status of boreal lake macrophytes

We assessed the relative roles of natural covariates, human disturbance (water quality and catchment land use) together with geography in driving variation in aquatic macrophyte community composition, richness and status among 101 lakes in southern and central Finland. In addition to all species together, we studied different growth forms (i.e. emergent and submerged macrophytes and aquatic bryophytes) separately. Partial redundancy analysis (taxonomic composition) and partial least-squares regression (species richness and status index) were employed to display the share of variability in macrophyte assemblages that was attributable to the environmental factors (both natural and human-affected) and the spatial filters generated through principal coordinates of neighbor matrices (PCNM).Macrophyte community composition, richness and status were explained by natural covariates, together with joint effects of human disturbance variables and space. The contributions of pure fractions of human disturbance and space were mostly modest, albeit variable among macrophyte groups and status indices. Alkalinity, historical distributions, colour, dynamic ratio and lake area were most important natural covariates for macrophytes. Of those variables influenced by human, macrophytes were mostly explained by conductivity, total phosphorus, turbidity and chlorophyll-a.Our results demonstrate, as expected, that macrophytes are dominantly affected by local environmental variables, whereas dispersal-related processes seem not to be important at regional extent. Response of macrophyte growth forms to environment and space, however, varied significantly. Community composition and richness of emergent macrophytes showed congruent response to natural covariates and human disturbance. Aquatic bryophytes, which are rarely studied along lake macrophytes, responded stronger than other growth forms to human disturbance. Contrary to our expectations, ecological indices were not affected by dispersal-related processes, but were mainly explained by natural covariates. This study is the first to investigate spatial patterns in aquatic macrophytes derived bioassessment. Geographical structuring of environmental variables and regional extent negatively affected indices, suggesting that ecological status assessment needs further development.     

Ecological relationships between stream communities and spatial scale: implications for designing catchment-level monitoring programmes

1. Stream communities are structured by factors acting over multiple spatial and temporal scales. Identifying what factors are driving spatial patterns in stream communities is a central aim of ecology.
2. Here we used two large European data sets of fish, invertebrates, macrophytes, benthic diatoms and environmental data in two stream groups (lowland and mountain) to determine the importance of variables at different spatial scales (geographical, regional, local) on community structure.
3. Both geographical position and ecoregion were selected first in canonical correspondence analysis (CCA), clearly showing the broad spatial gradients covered in the data set. Secondary predictors (after accounting for spatial and/or ecoregion effects) were similar between stream groups and among the four organism groups. In particular, conductivity and N concentration were strong predictors reflecting catchment land use.
4. Using partial CCA, we assessed the individual importance of the three spatial scales on the community structure of the four organism groups in the two stream groups. The majority of among-site variability (22–29%) was accounted for by local scale variables (e.g. water chemistry and substratum type), with regional and spatial variables accounting 11–13% and 5–6%, respectively. Our findings indicate that the four organism groups are responding similarly to the different levels of spatial scale, implying much redundancy which should be consider when implementing studies of bioassessment.     

Local environment and connectivity are the main drivers of diatom species composition and trait variation in a set of tropical reservoirs

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Relationships between stream macroinvertebrates and environmental variables at multiple spatial scales

1. Aquatic communities are structured by multiple forces, and identifying the driving factors over multispatial scales is an important research issue. The East Asian monsoon region is globally one of the richest environments in terms of biodiversity, and is undergoing rapid human development, yet the river ecosystems in this region have not been well studied. We applied a hierarchical framework to incorporate regional and local environmental effects on stream macroinvertebrate communities in this region. The knowledge gained is expected to improve the understanding of the importance of spatial scale on regional and local diversity in the East Asian monsoon region. 2. A national data set of benthic macroinvertebrates and environmental variables (geographical, land‐use, hydrological, substratum and physicochemical elements) in Korean rivers was used to determine the habitat preferences of macroinvertebrates. 3. Latitude, proportion of forest coverage, riffle habitat, silt substratum and temperature were the most important determinants for the ordinations of macroinvertebrate communities in each category evaluated by canonical correspondence analysis (CCA). The optimal habitats for stream macroinvertebrates are not the same for all species, and overall community metrics and abundance of sensitive species tended to be lower in open agricultural and urban streams than in forested streams. The sensitivity of mayflies and stoneflies to anthropogenic disturbances implicated them as good indicators to assess the effects of urban and agricultural activities. 4. A partial CCA was used to evaluate the relative importance of macrohabitat and microhabitat variables on community composition at three spatial scales (whole country, the large Han River basin and two small sub‐basins in the lowlands and highlands). The majority of community variation (17–22% for each environmental element) was explained by macrohabitat variables at the regional spatial scale. In contrast, large proportions (15–18%) were explained by microhabitat variables at the local spatial scale. 5. Our findings indicate that the relative importance of habitat scales should be determined by geographical size and that comprehensive understanding of multispatial scale patterns can be important for implementing sound biodiversity conservation programmes.     

A model of geographical, environmental and regional variation in vegetation composition of pyrogenic grasslands of Florida

Aim To develop a landscape‐level model that partitions variance in plant community composition among local environmental, regional environmental, and purely spatial predictive variables for pyrogenic grasslands (prairies, savannas and woodlands) throughout northern and central Florida. Location North and central Florida, USA. Methods We measured plant species composition and cover in 271 plots throughout the study region. A variation‐partitioning model was used to quantify components of variation in species composition associated with the main and interaction effects of soil and topographic variables, climate variables and spatial coordinates. Partial correlations of environmental variables with community variation were identified using direct gradient analysis (redundancy analysis and partial redundancy analysis) and Monte Carlo tests of significance. Results Community composition was most strongly related to edaphic variables at local scales in association with topographic gradients, although geographically structured edaphic, climatic and pure spatial effects were also evident. Edaphic variables explained the largest portion of total variation explained (TVE) as a main effect (48%) compared with the main effects of climate (9%) and pure spatial factors (9%). The remaining TVE was explained by the interaction effect of climate and spatial factors (13%) and the three‐way interaction (22%). Correlation analyses revealed that the primary compositional gradient was related to soil fertility and topographic position corresponding to soil moisture. A second gradient represented distinct geographical separation between the Florida panhandle and peninsular regions, concurrent with differences in soil characteristics. Gradients in composition corresponded to species richness, which was lower in the Florida peninsula. Main conclusions Environmental variables have the strongest influence on the species composition of Florida pyrogenic grasslands at both local and regional scales. However, the limited distributions of many plant taxa suggest historical constraints on species distributions from one physiographical region to the other (Florida panhandle and peninsula), although this pattern is partially confounded by regionally spatially structured environmental variables. Our model provides insight into the relative importance of local‐ and regional‐scale environmental effects as well as possible historical constraints on floristic variation in pine‐dominated pyrogenic grasslands of the south‐eastern USA.     

Ecological relationships between phytoplankton communities and different spatial scales in European reservoirs: implications at catchment level monitoring programmes

Phytoplankton communities are structured by factors acting over temporal and spatial scales. Identifying which factors are driving spatial patterns in aquatic communities is the central aim of ecology. In this study, data sets of phytoplankton communities and environmental data of two Portuguese reservoirs types (lowland “riverine reservoirs” and higher altitude “artificial lake reservoirs”) were used to determine the importance of environmental variables at different spatial (geographical, regional and local) and time scales (seasons, years) on the community structure. In all the data sets, the multivariate ordination technique Canonical Correspondence Analysis (CCA) showed that regional and local scales explained the majority (9–18% and 13–19%, respectively) of the taxa variance. However, for “riverine reservoirs”, time variables were more important, explaining 27% of the variability in phytoplankton assemblages. Variance partitioning was used to assess the individual importance of the three spatial scales and time for the community structure of the two reservoir types. The majority of among-site variability (5.9–21.4%) was accounted for by time variables, with local, regional, and geographical scale variables accounting for 3.3–5.6%, 3.7–4.5% and 2.6–2.9%, respectively. The effects of different spatial scales on phytoplankton communities were clearly interrelated; thus, implying that phytoplankton assemblages are capable of detecting stress from catchment to site scales. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. Padisak     

Environmental,spatial and phylogenetic determinants of fish life‐history traits and functional composition of Australian rivers

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Similar regional effects among local habitats on the structure of tropical reef fish and coral communities

Aim We examined data on corals and reef fishes to determine how particular local habitat types contribute to variation in community structure across regions covering gradients in species richness and how consistent this was over time. Location Great Barrier Reef (GBR), Australia. Methods We compared large‐scale (1300 km), long‐term (11 years) data on fishes and corals that were collected annually at fixed sites in three habitats (inshore, mid‐shelf and outer‐shelf reefs) and six regions (latitudinal sectors) along a gradient of regional species richness in both communities. We used canonical approaches to partition variation in community structure (sites × species abundance data matrices) into components associated with habitat, region and time and Procrustes analyses to assess the degree of concordance between coral and fish community structure. Results Remarkably similar patterns emerged for both fish and coral communities occupying the same sites. Reefs that had similar coral communities also had similar fish communities. The fraction of the community data that could be explained by regional effects, independent of pure habitat effects, was similar in both fish (33%) and coral (36.9%) communities. Pure habitat effects were slightly greater in the fish (31.3%) than in the coral (20.1%) community. Time explained relatively little variation (fish = 7.9%, corals = 9.6%) compared with these two spatial factors. Conclusions Our results indicate either that fish and coral communities were structured in similar ways by processes associated with region, habitat and time, or that the variation in fish community structure tracked variation associated with the coral communities at these sites and thereby reflects an indirect link between the environment and the structure of fish communities mediated by corals. Irrespective of the causes of such commonality, we demonstrate that community structure, not just species richness, can be related to both habitat differences and regional setting simultaneously.     

Metacommunity ecology meets biogeography: effects of geographical region,spatial dynamics and environmental filtering on community structure in aquatic organisms

Metacommunity patterns and underlying processes in aquatic organisms have typically been studied within a drainage basin. We examined variation in the composition of six freshwater organismal groups across various drainage basins in Finland. We first modelled spatial structures within each drainage basin using Moran eigenvector maps. Second, we partitioned variation in community structure among three groups of predictors using constrained ordination: (1) local environmental variables, (2) spatial variables, and (3) dummy variable drainage basin identity. Third, we examined turnover and nestedness components of multiple-site beta diversity, and tested the best fit patterns of our datasets using the “elements of metacommunity structure” analysis. Our results showed that basin identity and local environmental variables were significant predictors of community structure, whereas within-basin spatial effects were typically negligible. In half of the organismal groups (diatoms, bryophytes, zooplankton), basin identity was a slightly better predictor of community structure than local environmental variables, whereas the opposite was true for the remaining three organismal groups (insects, macrophytes, fish). Both pure basin and local environmental fractions were, however, significant after accounting for the effects of the other predictor variable sets. All organismal groups exhibited high levels of beta diversity, which was mostly attributable to the turnover component. Our results showed consistent Clementsian-type metacommunity structures, suggesting that subgroups of species responded similarly to environmental factors or drainage basin limits. We conclude that aquatic communities across large scales are mostly determined by environmental and basin effects, which leads to high beta diversity and prevalence of Clementsian community types.