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 comparative analysis reveals weak relationships between ecological factors and beta diversity of stream insect metacommunities at two spatial levels

The hypotheses that beta diversity should increase with decreasing latitude and increase with spatial extent of a region have rarely been tested based on a comparative analysis of multiple datasets, and no such study has focused on stream insects. We first assessed how well variability in beta diversity of stream insect metacommunities is predicted by insect group, latitude, spatial extent, altitudinal range, and dataset properties across multiple drainage basins throughout the world. Second, we assessed the relative roles of environmental and spatial factors in driving variation in assemblage composition within each drainage basin. Our analyses were based on a dataset of 95 stream insect metacommunities from 31 drainage basins distributed around the world. We used dissimilarity‐based indices to quantify beta diversity for each metacommunity and, subsequently, regressed beta diversity on insect group, latitude, spatial extent, altitudinal range, and dataset properties (e.g., number of sites and percentage of presences). Within each metacommunity, we used a combination of spatial eigenfunction analyses and partial redundancy analysis to partition variation in assemblage structure into environmental, shared, spatial, and unexplained fractions. We found that dataset properties were more important predictors of beta diversity than ecological and geographical factors across multiple drainage basins. In the within‐basin analyses, environmental and spatial variables were generally poor predictors of variation in assemblage composition. Our results revealed deviation from general biodiversity patterns because beta diversity did not show the expected decreasing trend with latitude. Our results also call for reconsideration of just how predictable stream assemblages are along ecological gradients, with implications for environmental assessment and conservation decisions. Our findings may also be applicable to other dynamic systems where predictability is low.     

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.     

Role of environmental and spatial processes in structuring anuran communities across a tropical rain forest

This study aimed to evaluate if anuran species distributions in riparian and non‐riparian areas are influenced by environmental factors (i.e. niche) and/or by spatial factors (i.e. dispersal). The environmental variables analysed were altitude, distance from the stream and leaf litter depth. Spatial factors were represented by the eigenvectors extracted from geographical coordinates by eigenfunction analysis. The study was conducted in 24 km² of terra‐firme forest in Central Amazonia, Manaus – Amazonas, Brazil. Between November 2008 and May 2009, three samples were taken from 41 plots, 21 plots being placed at non‐riparian areas and another 20 placed in riparian areas. We submitted the assemblage dataset to a partial redundancy analysis to evaluate the contributions of environmental and spatial variables (selected with a forward selection procedure). In addition, we tested if communities differ from riparian and non‐riparian areas using a db‐MANOVA. Species richness and species composition differed between riparian and non‐riparian plots. Some species were restricted to riparian areas. Altitude was the only significant variable (P = 0.005) explaining 21% of the total variance. When analysing the data from all plots using the partial redundancy analysis, 27% of the variance was explained by spatial and environmental variables. The environmental variables explained exclusively 4% of the variance in assemblage composition, and 13% was explained by environmental variables that were also structured in space (i.e. the shared fraction), while 10% was explained exclusively by spatial variables. In conclusion, our results showed differences between the assemblages of riparian and non‐riparian areas which can be explained by the distribution of anuran species along environmental gradients altitude and distance to streams, with little evidence of dispersal limitation.     

Control of stream insect assemblages: roles of spatial configuration and local environmental factors

Abstract 1. Current views in ecology emphasise that community structure is the sum of multiple processes, with imprints of both regional and local drivers. However, the degree to which stream insect assemblages are structured by spatial configuration (complying with the dispersal‐based neutral hypothesis) and local environmental features (complying with the niche‐based species sorting hypothesis) has not been rigorously examined based on surveys in multiple years. 2. Stream sites in a boreal drainage system were surveyed during three consecutive years and the relative contribution of spatial configuration and local environmental variables to aquatic insect assemblage structure (characterised by both abundance and presence–absence data) was assessed. Separate analyses were conducted for mayflies (Ephemeroptera), stoneflies (Plecoptera), caddisflies (Trichoptera), and non‐biting midges (Diptera: Chironomidae) in each year. 3. There were no relationships between the spatial location and local environmental features of streams in Mantel tests, facilitating exploration of their independent effects on assemblage structure. The study found virtually no effects of spatial location on stream insect assemblages across the study drainage system, as evidenced by Mantel tests and canonical correspondence analyses (CCA). The environmental variables were also rather weakly associated with assemblage structure, with the total amount of explained variation ranging from 9.8% to 31.7% in the CCAs. There were no appreciable differences in the amount of environment‐related explained variation in assemblage structure between mayflies, stoneflies, caddisflies, and midges, but some between‐year differences were noticeable in most insect groups. The environmental variables that were significantly related to assemblage structure exhibited some between‐group and between‐year variability. In general, patterns shown by abundance and presence–absence data were highly similar. 4. It appears that stream insect assemblages comply with the niche‐based species sorting hypothesis in the context of metacommunity ecology. In contrast, the absence of spatial structuring suggests that stream insect assemblages do not comply with the neutral hypothesis, being not strongly dispersal limited at the within‐drainage basin scale.     

The relationship between species richness and biomass changes from boreal to subtropical forests in China

Diversity‐manipulation experiments suggest a positive effect of biodiversity on ecosystem properties (EPs), but variable relationships between species richness and EPs have been reported in natural communities. An explanation for this discrepancy is that observed richness–EPs relationships in natural communities change with environment and species functional identities. But how the relationships change along broad‐scale climatic gradients has rarely been examined. In this paper, we sampled 848 plots of 20 × 30 m² from boreal to tropical forests across China. We examined plot biomass with respect to environmental factors, tree species richness and functional group identity (FGI, i.e. evergreen vs deciduous, and coniferous vs broadleaf). Variation partitioning was used to evaluate the relative effects of the three classes of factors. We found that, most of the ‘effects’ (percentage of variation explained) of richness and FGI on forest biomass were shared with environmental factors, but species richness and FGI still revealed significant effects in addition to environment for plots across China. Richness and FGI explained biomass mainly through their shared effects instead of independent effects, suggesting that the positive biodiversity effect is closely associated with a sampling effect. The relative effects of richness, FGI and environment varied latitudinally: the independent effects of environment and richness decreased from boreal to subtropical forests, whereas the total effect of FGI increased. We also found that the slope of richness–biomass relationship decreased monotonically from boreal to subtropical forests, possibly because of decreasing complementarity and increasing competition with increasing productivity. Our results suggest that while species richness does have significant effects on forest biomass it is less important than environmental gradients and other biotic factors in shaping large‐scale biomass patterns. We suggest that understanding how and why the diversity–EPs relationships change along climatic gradient would be helpful for a better understanding of real biodiversity effects in natural communities.     

The Influence of Environmental,Biotic and Spatial Factors on Diatom Metacommunity Structure in Swedish Headwater Streams

Stream assemblages are structured by a combination of local (environmental filtering and biotic interactions) and regional factors (e.g., dispersal related processes). The relative importance of environmental and spatial (i.e., regional) factors structuring stream assemblages has been frequently assessed in previous large-scale studies, but biotic predictors (potentially reflecting local biotic interactions) have rarely been included. Diatoms may be useful for studying the effect of trophic interactions on community structure since: (1) a majority of experimental studies shows significant grazing effects on diatom species composition, and (2) assemblages can be divided into guilds that have different susceptibility to grazing. We used a dataset from boreal headwater streams in south-central Sweden (covering a spatial extent of ∼14000 km²), which included information about diatom taxonomic composition, abundance of invertebrate grazers (biotic factor), environmental (physicochemical) and spatial factors (obtained through spatial eigenfunction analyses). We assessed the relative importance of environmental, biotic, and spatial factors structuring diatom assemblages, and performed separate analyses on different diatom guilds. Our results showed that the diatom assemblages were mainly structured by environmental factors. However, unique spatial and biological gradients, specific to different guilds and unrelated to each other, were also evident. We conclude that biological predictors, in combination with environmental and spatial variables, can reveal a more complete picture of the local vs. regional control of species assemblages in lotic environments. Biotic factors should therefore not be overlooked in applied research since they can capture additional local control and therefore increase accuracy and performance of predictive models. The inclusion of biotic predictors did, however, not significantly influence the unique fraction explained by spatial factors, which suggests low bias in previous assessments of unique regional control of stream assemblages.     

The influence of niche and neutral processes on a neotropical anuran metacommunity

One of the most important questions in ecology is the relative importance of local conditions (niche processes) and dispersal ability (neutral processes) in driving metacommunity structure. Although many studies have been conducted in recent years, there is still much debate. We evaluated the processes (niche and neutral) responsible for variation in anuran composition in 28 lentic water bodies in southeastern Brazil. Because anurans depend heavily on environmental conditions, we hypothesized that environmental variables (niche processes) are the most important drivers of community composition. Additionally, as anurans have limited dispersal abilities, and the study region presents harsh conditions (high forest fragmentation, low rainfall and long dry season), we expected a lower, but significant, spatial signature in metacommunity structure, due to neutral dynamics. We used a partial redundancy analysis with variation partitioning to evaluate the relative influence of environmental and spatial variables as drivers of metacommunity structure. Additionally, we used a recently developed spatial autocorrelation analysis to test if neutral dynamics can be attributed to the pure spatial component. This analysis is based on predictions that species abundances are independent but similarly spatially structured, with correlograms similar in shape. Therefore, under neutral dynamics there is no expectation of a correlation between the pairwise distance of spatial correlograms and the pairwise correlation of species abundances predicted by the pure spatial component. We found that the environmental component explained 21.5%, the spatial component 10.2%, and the shared component 6.4% of the metacommunity structure. We found no correlation between correlograms and correlation of abundances predicted by the pure spatial component (Mantel test = ?0.109, P = 0.961). In our study, niche‐based processes are the dominant process that explained community composition. However, neutral processes are important because spatial variation can be attributed to pure neutral dynamics rather than to missing spatially structured environmental factors.     

Hierarchical Dynamics of Ecological Communities: Do Scales of Space and Time Match?

Theory posits that community dynamics organize at distinct hierarchical scales of space and time, and that the spatial and temporal patterns at each scale are commensurate. Here we use time series modeling to investigate fluctuation frequencies of species groups within invertebrate metacommunities in 26 boreal lakes over a 20-year period, and variance partitioning analysis to study whether species groups with different fluctuation patterns show spatial signals that are commensurate with the scale-specific fluctuation patterns identified. We identified two groups of invertebrates representing hierarchically organized temporal dynamics: one species group showed temporal variability at decadal scales (slow patterns of change), whilst another group showed fluctuations at 3 to 5-year intervals (faster change). This pattern was consistently found across all lakes studied. A spatial signal was evident in the slow but not faster-changing species groups. As expected, the spatial signal for the slow-changing group coincided with broad-scale spatial patterns that could be explained with historical biogeography (ecoregion delineation, and dispersal limitation assessed through a dispersal trait analysis). In addition to spatial factors, the slow-changing groups correlated with environmental variables, supporting the conjecture that boreal lakes are undergoing environmental change. Taken together our results suggest that regionally distinct sets of taxa, separated by biogeographical boundaries, responded similarly to broad-scale environmental change. Not only does our approach allow testing theory about hierarchically structured space-time patterns; more generally, it allows assessing the relative role of the ability of communities to track environmental change and dispersal constraints limiting community structure and biodiversity at macroecological scales.     

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.     

Spatial autocorrelation analysis allows disentangling the balance between neutral and niche processes in metacommunities

One of the most popular approaches for investigating the roles of niche and neutral processes driving metacommunity patterns consists of partitioning variation in species data into environmental and spatial components. The logic is that the distance decay of similarity in communities is expected under neutral models. However, because environmental variation is often spatially structured, the decay could also be attributed to environmental factors that are missing from the analysis. Here, we use a spatial autocorrelation analysis protocol, previously developed to detect isolation‐by‐distance in allele frequencies, to evaluate patterns of species abundances under neutral dynamics. We show that this protocol can be linked with variation partitioning analyses. Moreover, in an attempt to test the neutral model, we derive three predictions to be applied both to original species abundances and to abundances predicted by a pure spatial model species abundances will be uncorrelated; Moran's I correlograms will reveal similar short‐distance autocorrelation patterns; an increasing degree of non‐neutrality will tend to generate patterns of correlation among abundances within groups of species with similar correlograms (i.e. within species with neutral and non‐neutral dynamics). We illustrate our protocol by analyzing spatial patterns in abundance of 28 terrestrially breeding anuran species from Central Amazonia. We recommend that researchers should investigate spatial autocorrelation patterns of abundances predicted by pure spatial models to identify similar patterns of spatial autocorrelation at short distances and lack of correlation between species abundances. Therefore, the hypothesis that spatial patterns in abundances are primarily due to pure neutral dynamics (rather than to missing spatiallystructured environmental factors) can be confirmed after taking environmental variables into account.     

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.     

Spatial, environmental and human influences on the distribution of otter (Lutra lutra) in the Spanish provinces

In a previous survey of otters (Lutra lutra L. 1758) in Spain, different causes were invoked to explain the frequency of the species in each province. To find common causes of the distribution of the otter in Spain, we recorded a number of spatial, environmental and human variables in each Spanish province. We then performed a stepwise linear multiple regression of the proportion of positive sites of otter in the Spanish provinces separately on each of the three groups of variables. Geographic longitude, January air humidity, soil permeability and highway density were the variables selected. A linear regression of the proportion of otter presence on these variables explained 62.4% of the variance. We then used the selected variables in a partial regression analysis to specify which proportions of the variation are explained exclusively by spatial, environmental and human factors, and which proportions are attributable to interactions between these components. Pure environmental effects accounted for only 5.5% of the variation, while pure spatial and pure human effects explained 18% and 9.7%, respectively. Shared variation among the components totalled 29.2%, of which 10.9% was explained by the interaction between environmental and spatial factors. Human factors explained globally less variance than spatial and environmental ones, but the pure human influence was higher than the pure environmental one. We concluded that most of the variation in the proportion of occurrences of otter in Spanish provinces is spatially structured, and that environmental factors have more influence on otter presence than human ones; however, the human influence on otter distribution is less structured in space, and thus can be more disruptive. This effect of large infrastructures on wild populations must be taken into account when planning large‐scale conservation policies.     

Modelling spatial and environmental effects at the edge of the distribution: the red-backed shrike Lanius collurio in Northern Portugal

This study examined the interplay of spatial and environmental effects shaping the range margin of the red‐backed shrike (Lanius collurio) in northern Portugal. The occurrence of shrikes in 10 × 10 km UTM squares was related to three sets of explanatory variables, reflecting environmental effects (climate and habitat), large‐scale spatial trends, and neighbourhood influences (considering an autologistic term); spatial variables were used as surrogates for historical and demographic factors. Multiple logistic regression models were built for each set, and then variation partitioning based on partial regressions isolated the unique and shared components of explained variation. The environmental model revealed a dominant influence of climate effects, with the occurrence of shrikes increasing with frost and thermal amplitude, declining with insolation, and responding unimodally to rainfall. There was a weaker influence of habitat conditions, though shrikes were more likely with increasing cover by annual crops and pastures, and decreasing forest cover. Only a relatively small proportion of explained variation was due to a ‘pure’ environmental component (10.4%), as most variation explained by environmental factors appeared spatially structured (51.9%). The unique contributions of spatial variables to the overall model were also small, though the neighbourhood effects appeared relatively stronger than large‐scale trends. Taken together, results suggested that the south‐western range margin of the red‐backed shrike was largely determined by spatially structured environmental factors. Nevertheless, there were also ‘pure’ environmental factors determining some isolate occurrences irrespective of any spatial structure, and ‘pure’ spatial factors that appeared to favour the occupation of squares surrounding the core distribution areas irrespective of environmental conditions. These results add to the growing evidence that both environmental and spatial factors need to be considered in predictive modelling of species range margins.     

Does dispersal ability affect the relative importance of environmental control and spatial structuring of littoral macroinvertebrate communities?

Both spatial processes and environmental control may structure metacommunities, but their relative importance may be contingent on the dispersal ability of organisms. I examined the roles of spatial and environmental factors for the structuring of littoral macroinvertebrate communities across a set of lakes in a boreal drainage basin. I hypothesized that dispersal ability would affect the relative importance of spatial processes and environmental control, and thus the biological data were divided into four groups of species differing in dispersal ability. In general, the group of the strongest aerial dispersers showed greatest relative pure environmental control and least pure spatial structuring of community structure and species richness, while spatial processes seemed to be more important for the other three dispersal ability groups. However, these results were contingent on the indirect measure of spatial processes, with the spatial variables and connectivity variables providing slightly different insights into the spatial processes and environmental control of metacommunity structuring. It appears, however, that dispersal ability has effects on the spatial processes and environmental control important in metacommunity organization, with strong dispersers being more under environmental control and less affected by spatial processes compared to weak dispersers.     

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.     

Hydrological connectivity structures concordant plant and animal assemblages according to niche rather than dispersal processes

1. Understanding the processes that structure community assembly across landscapes is fundamental to ecology and for predicting and managing the consequences of anthropogenically induced changes to ecosystems. 2. We assessed the community similarity of fish, macroinvertebrate and vegetation communities against geographic distances ranging from 4 to 480 km (i.e. distance–decay relationships) to determine the balance between local environmental factors and regional dispersal processes, and thus whether species‐sorting (niche processes) or dispersal limitation (neutral processes) was more important in structuring these assemblages in Australia’s wet‐dry tropics. We investigated whether the balance between niche and dispersal processes depended on the degree of hydrological connectivity, predicting that dispersal processes would be more important at connected sites, and also whether there was spatial concordance among these three assemblage types. 3. There was significant but weak spatial concordance among the study communities, suggesting limited potential for surrogacy among them. Distance–decay in community similarity was not observed for any study assemblage at perennial sites, suggesting dispersal was not limiting and assemblages were structured more strongly by local niche processes at these connected sites. At intermittent sites, weak distance–decay relationships for each assemblage type were confounded by significant relationships with environmental dissimilarity, suggesting that dispersal limitation contributed, albeit weakly, to niche processes in structuring our three study assemblages at disconnected sites. 4. Two environmental factors, flow regime and channel width, explained significant proportions of variation in all three assemblages, potentially contributing to the observed spatial concordance between them and representing local environmental gradients along which these communities re‐assemble after the wet season, according to niche rather than dispersal processes.     

Determinants of beta diversity of spiders in coastal dunes along a gradient of mediterraneity

Aim The Mediterranean Basin is recognized for its high levels of species richness, rarity and endemicity. Our main aim was to evaluate the relative effects of environmental and spatial variables and their scale‐specific importance on beta diversity patterns along a gradient of mediterraneity, using spiders as a model group. Location This study was carried out in 18 coastal dune sites along the Portuguese Atlantic coast. This area encompasses 445 km and comprises two distinct biogeographic regions, Eurosiberian (northern coast) and Mediterranean (centre and south). Methods A forward selection procedure was carried out to select environmental and spatial variables responsible for determining beta diversity patterns. Variation partitioning and principal coordinates of neighbour matrices (PCNM) were used to estimate the contribution of pure environmental and pure spatial effects and their shared influence on beta diversity patterns and to estimate the relative importance of environmental structured variation and pure spatial variation at multiple spatial scales. Results Climate, ground vegetation dune cover and area were selected by a forward selection procedure. The same procedure identified three PCNM variables, all corresponding to large and medium spatial scales. Variation partitioning revealed that 46.1% of the variation of beta diversity patterns was explained by a combination of environmental and PCNM variables. Most of this variation (42.5%) corresponded to spatial variation (environmental spatially structured and pure spatial). Climate and vegetation structure influences were predominant at the PCNM1 and PCNM3 scales, while area was more important at the intermediate PCNM2 scale. Main conclusions Our study revealed that beta diversity of spiders was primarily controlled by a broad‐scale gradient of mediterraneity. The relative importance of environmental variables on the spider assemblage composition varied with spatial scale. This study highlights the need of considering the scale‐specific influence of niche and neutral processes on beta diversity patterns.     

Multivariate analysis of a fine-scale breeding bird atlas using a geographical information system and partial canonical correspondence analysis: environmental and spatial effects

Aim To assess the relative roles of environment and space in driving bird species distribution and to identify relevant drivers of bird assemblage composition, in the case of a fine‐scale bird atlas data set. Location The study was carried out in southern Belgium using grid cells of 1 × 1 km, based on the distribution maps of the Oiseaux nicheurs de Famenne: Atlas de Lesse et Lomme which contains abundance for 103 bird species. Methods Species found in < 10% or > 90% of the atlas cells were omitted from the bird data set for the analysis. Each cell was characterized by 59 landscape metrics, quantifying its composition and spatial patterns, using a Geographical Information System. Partial canonical correspondence analysis was used to partition the variance of bird species matrix into independent components: (a) ‘pure’ environmental variation, (b) spatially‐structured environmental variation, (c) ‘pure’ spatial variation and (d) unexplained, non‐spatial variation. Results The variance partitioning method shows that the selected landscape metrics explain 27.5% of the variation, whilst ‘pure’ spatial and spatially‐structured environmental variables explain only a weak percentage of the variation in the bird species matrix (2.5% and 4%, respectively). Avian community composition is primarily related to the degree of urbanization and the amount and composition of forested and open areas. These variables explain more than half of the variation for three species and over one‐third of the variation for 12 species. Main conclusions The results seem to indicate that the majority of explained variation in species assemblages is attributable to local environmental factors. At such a fine spatial resolution, however, the method does not seem to be appropriated for detecting and extracting the spatial variation of assemblages. Consequently, the large amount of unexplained variation is probably because of missing spatial structures and ‘noise’ in species abundance data. Furthermore, it is possible that other relevant environmental factors, that were not taken into account in this study and which may operate at different spatial scales, can drive bird assemblage structure. As a large proportion of ecological variation can be shared by environment and space, the applied partitioning method was found to be useful when analysing multispecific atlas data, but it needs improvement to factor out all‐scale spatial components of this variation (the source of ‘false correlation’) and to bring out the ‘pure’ environmental variation for ecological interpretation.