Unraveling the effects of water–sediment conditions and spatial patterns on Unionida assemblages in seasonally connected floodplain lakes

Floodplain lakes are good metacommunity systems to study the environmental and spatial processes structuring local assemblages. They are more connected during high-water periods and are more isolated during low-water periods. We evaluated the effects of lake spatial patterning and water and sediment conditions on Unionida species assemblages. Moran Eigenvector Maps were used to generate spatial variables representing spatial patterns at different scales. We sampled 35 lakes from the Pantanal floodplain, Brazil. To understand the effects of environmental and spatial variables, we performed Redundancy Analyses and variation partitioning to separate environmental and spatial pattern effects. Environmental variables explained almost twice the variation in the Pantanal mussel assemblages than did spatial variables. Unionida species presence was driven mainly by variations in sediment coarse sand and silt contents. The weak spatial patterns observed may be related to increased connectivity between lakes during floods, which facilitates mussel host fish dispersal. Mussel abundances were driven mainly by organic matter availability, but varied between species. Changes in lake connectivity can affect the regional sediment dynamics and affect mussel assemblages.

     

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.     

Environmental factors exert predominant effects on testate amoeba metacommunities during droughts in floodplains

Metacommunities have been evaluated as models for the relative importance of environmental and spatial processes in assembling ecological communities. Here, we tested the hypothesis that different hydrological periods (drought and flooding) influence the environmental associations of planktonic testate amoeba metacommunities. We predicted that environmental factors would exert the strongest effects on species dispersal under drought, but that they would be less significant during flooding. Testate amoebae were sampled during drought and flooding, from 72 lakes in four Brazilian floodplains (Amazonian, Araguaia, Pantanal and Paraná). Partial redundancy analysis indicated that only environmental factors were significant; they were significant in all floodplain lakes during the drought. Only the Paraná floodplain had significant results for environmental factors during both hydrological periods. Spatial factors did not contribute significantly to any of the metacommunities. The depth, pH and variables related to environmental productivity were identified as major predictors in the assembly of testate amoeba communities. Our results highlight that different hydrological periods vary in their relative importance in environmental and spatial processes. The species‐sorting model was predominant during drought, while stochastic processes prevailed during flooding in all but the Paraná floodplain. In the Paraná floodplain, the construction of dams could potentially alter the effects of environmental and spatial factors on the dispersal of planktonic testate amoebae. The pH and the environmental productivity factors were largely responsible for species selection and the structuring of the planktonic testate amoebae metacommunities in Brazilian floodplains.     

Correlates of Zooplankton Beta Diversity in Tropical Lake Systems

The changes in species composition between habitat patches (beta diversity) are likely related to a number of factors, including environmental heterogeneity, connectivity, disturbance and productivity. Here, we used data from aquatic environments in five Brazilian regions over two years and two seasons (rainy and dry seasons or high and low water level periods in floodplain lakes) in each year to test hypotheses underlying zooplankton beta diversity variation. The regions present different levels of hydrological connectivity, where three regions present lakes that are permanent and connected with the main river, while the water bodies of the other two regions consist of permanent lakes and temporary ponds, with no hydrological connections between them. We tested for relationships between zooplankton beta diversity and environmental heterogeneity, spatial extent, hydrological connectivity, seasonality, disturbance and productivity. Negative relationships were detected between zooplankton beta diversity and both hydrological connectivity and disturbance (periodic dry-outs). Hydrological connectivity is likely to affect beta diversity by facilitating dispersal between habitats. In addition, the harsh environmental filter imposed by disturbance selected for only a small portion of the species from the regional pool that were able to cope with periodic dry-outs (e.g., those with a high production of resting eggs). In summary, this study suggests that faunal exchange and disturbance play important roles in structuring local zooplankton communities.     

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.     

Disentangling the role of connectivity,environmental filtering,and spatial structure on metacommunity dynamics

Dispersal is a key process in metacommunity dynamics, allowing the maintenance of diversity in complex community networks. Geographic distance is usually used as a surrogate for connectivity implying that communities that are closely located are considered more prone to exchange individuals than distant communities. However, in some natural systems, organisms may be subjected to directional dispersal (air or water flows, particular landscape configuration), possibly leading close communities to be isolated from each other and distant communities to be connected. Using geographic distance as a proxy for realised connectivity may then yield misleading results regarding the role of dispersal in structuring communities in such systems. Here, we quantified the relative importance of flow connectivity, geographic distance, and environmental gradients to explain polychaete metacommunity structure along the coasts of the Gulf of Lions (northwest Mediterranean Sea). Flow connectivity was estimated by Lagrangian particle dispersal simulations. Our results revealed that this metacommunity is strongly structured by the environment at large spatial scales, and that both flow connectivity and geographic distance play an important role within homogeneous environments at smaller spatial scales. We thus strongly advocate for a wider use of connectivity measures, in addition to geographic distance, to study spatial patterns of biological diversity (e.g. distance decay) and to infer the processes behind these patterns at different spatial scales. Synthesis Everything is connected, but connections are seldom accurately quantified. Biological communities are often studied separately, using observations, experiments and models to unravel local dynamics of organisms interacting with each other. However, regional processes such as dispersal through ocean and air circulation, likely to connect distant communities and influence their local dynamics, are not always accounted for, or, at best, used as an homogeneous and distance‐related factor. Ocean models have being extensively developed and validated during the past decades with the increasing availability of accurate meteorological data. Using such model outputs, precise quantifi cation of exchange rates of organisms between communities was performed in a marine Mediterranean coastal area. Jointly with local environmental and biological data, these results were used to quantify the effects of realistic connectivity on local and regional polychaete community structure, and revealed that the environmental gradient, geographic distance, and connectivity were responsible for community structure at different spatial scales.     

Spatial and environmental effects on a rock‐pool metacommunity depend on landscape setting and dispersal mode

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Ecological uniqueness of macroinvertebrate communities in high-latitude streams is a consequence of deterministic environmental filtering processes

Variation in biological communities is a consequence of stochastic and deterministic factors. Examining the relative importance of these factors helps to understand variation in the whole biodiversity in a region. We examined the roles of stochastic and deterministic factors in structuring macroinvertebrate communities in high-latitude streams across two seasons. We predicted that if communities are the result of deterministic environmental filtering processes, the communities should show strong association with environmental variables, as taxa would be selected according to stream environmental conditions. However, if communities are driven by stochastic factors, they should show strong association with spatial variables, as the distribution of taxa in communities would be driven by spatially related dispersal factors. We studied these predictions by calculating the degree of uniqueness of the streams in terms of their taxonomic and functional community compositions and by modelling the resulting index values using spatial and environmental variables. Our results supported the first prediction where the communities are more influenced by the environmental filtering processes, although indications of the effect of spatial processes in structuring the communities were present especially in autumn. High-latitude stream communities also seem to be sensitive to environmental changes, as even small changes in environment were enough to affect the ecological uniqueness of the streams. These findings highlight the vulnerability of northern streams in the face of the climate change. To maintain biodiversity in high-latitude catchments, it would be important to protect varying habitat conditions, which are the main forces affecting the ecological uniqueness of the streams.     

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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The role of environmental and spatial processes in structuring native and non‐native fish communities across thousands of lakes

Quantifying the role of spatial patterns is an important goal in ecology to further understand patterns of community composition. We quantified the relative role of environmental conditions and regional spatial patterns that could be produced by environmental filtering and dispersal limitation on fish community composition for thousands of lakes. A database was assembled on fish community composition, lake morphology, water quality, climatic conditions, and hydrological connectivity for 9885 lakes in Ontario, Canada. We utilized a variation partitioning approach in conjunction with Moran's Eigenvector Maps (MEM) and Asymmetric Eigenvector Maps (AEM) to model spatial patterns that could be produced by human‐mediated and natural modes of dispersal. Across 9885 lakes and 100 fish species, environmental factors and spatial structure explained approximately 19% of the variation in fish community composition. Examining the proportional role of spatial structure and environmental conditions revealed that as much as 90% of the explained variation in native species assemblage composition is governed by environmental conditions. Conversely on average, 67% of the explained variation in non‐native assemblage composition can be related to human‐mediated dispersal. This study highlights the importance of including spatial structure and environmental conditions when explaining patterns of community composition to better discriminate between the ecological processes that underlie biogeographical patterns of communities composed of native and non‐native fish species.     

Metacommunity organisation,spatial extent and dispersal in aquatic systems: patterns,processes and prospects

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No evidence for environmental and spatial processes in structuring phytoplankton communities

The relative importance of local and regional processes in shaping natural communities within a metacommunity context has been a focus of intense debate in recent years. Floodplain lakes provide a good system for testing this theoretical approach, as they undergo seasonal variations in physical, chemical and biological factors, as well as in their degree of connectivity. Here, we investigated how local phytoplankton communities in lakes of a tropical river-floodplain system (Araguaia River floodplain – Central Brazil) were affected by environmental and spatial (dispersal) predictors in two rainy and two dry seasons (two consecutive years). Partial redundancy analysis indicated that during the periods analyzed the effects of neither predictor were significant. Although we cannot exclude the possibility that these tropical phytoplankton communities could be regulated by stochastic events, we suggested that further studies will have greater explanatory power if they include other variables related to biotic interactions (e.g., abundance of grazers) and fine-scale environmental variation.     

Variance partitioning of deconstructed periphyton communities: does the use of biological traits matter?

The use of species traits offers a promising approach to the understanding of the main processes underlying metacommunity patterns. We analyzed samples of periphytic algae in 30 environments of the Upper Paraná River floodplain in southeastern Brazil, to test the hypotheses that variation in species composition of algal groups with low dispersal abilities would be mainly explained by spatial variables; on the other hand, algal groups with higher dispersal abilities would be better explained by environmental variables. The variation in community structure was mainly correlated with environmental variables. This result is in line with a growing body of evidence indicating a predominant role of species-sorting processes. The more-refined prediction that the spatial variables would gradually become more important across a gradient of adherence or size was, however, not supported by our analyses. Also, the large unexplained variation suggested that these periphytic communities were assembled by idiosyncratic events, or that other variables that are often neglected in studies of aquatic metacommunities needed to be included.     

The relative importance of local conditions and regional processes in structuring aquatic plant communities

1. The structure of biological communities reflects the influence of both local environmental conditions and processes such as dispersal that create patterns in species’ distribution across a region. 2. We extend explicit tests of the relative importance of local environmental conditions and regional spatial processes to aquatic plants, a group traditionally thought to be little limited by dispersal. We used partial canonical correspondence analysis and partial Mantel tests to analyse data from 98 lakes and ponds across Connecticut (northeastern United States). 3. We found that aquatic plant community structure reflects the influence of local conditions (pH, conductivity, water clarity, lake area, maximum depth) as well as regional processes. 4. Only 27% of variation in a presence/absence matrix was explained by environmental conditions and spatial processes such as dispersal. Of the total explained, 45% was related to environmental conditions and 40% to spatial processes. 5. Jaccard similarity declined with Euclidean distance between lakes, even after accounting for the increasing difference in environmental conditions, suggesting that dispersal limitation may influence community composition in the region. 6. The distribution of distances among lakes where species occurred was associated with dispersal‐related functional traits, providing additional evidence that dispersal ability varies among species in ways that affect community composition. 7. Although environmental and spatial variables explained a significant amount of variation in community structure, a substantial amount of stochasticity also affects these communities, probably associated with unpredictable colonisation and persistence of the plants.     

The role of watercourse versus overland dispersal and niche effects on ostracod distribution in Mediterranean streams (eastern Iberian Peninsula)

The processes behind the heterogeneous distribution of species involve a combination of environmental and spatial effects. In the spatial context, stream networks constitute appropriate systems to compare the relative importance of two dispersal modes in aquatic organisms: overland and watercourse dispersal. In the present study, we analyzed the distribution of ostracod species in a river network in the eastern Iberian Peninsula, with variation partitioning between environmental and spatial factors, using Moran and Asymmetric Eigenvector Maps (MEMs, AEMs) as spatial variables. Our aims were to determine the relative importance of environmental and spatial control and to compare the importance of overland and watercourse dispersal for species distribution of passively-dispersing aquatic organisms. Our results suggest that watercourse was the most important dispersal mode, favoring mass-effects. The role of species sorting was significant and related to temperature, stream width and water quality, measured as a biotic index (IBMWP). These results stress the major importance of connectivity, besides niche-related factors, in structuring riverine communities of passively-dispersing aquatic organisms.     

Metacommunity Structure of Stream Insects across Three Hierarchical Spatial scales

A major challenge in community ecology is to understand the underlying factors driving metacommunity (i.e., a set of local communities connected through species dispersal) dynamics. However, little is known about the effects of varying spatial scale on the relative importance of environmental and spatial (i.e., dispersal related) factors in shaping metacommunities and on the relevance of different dispersal pathways. Using a hierarchy of insect metacommunities at three spatial scales (a small, within‐stream scale, intermediate, among‐stream scale, and large, among‐sub‐basin scale), we assessed whether the relative importance of environmental and spatial factors shaping metacommunity structure varies predictably across spatial scales, and tested how the importance of different dispersal routes vary across spatial scales. We also studied if different dispersal ability groups differ in the balance between environmental and spatial control. Variation partitioning showed that environmental factors relative to spatial factors were more important for community composition at the within‐stream scale. In contrast, spatial factors (i.e., eigenvectors from Moran's eigenvector maps) relative to environmental factors were more important at the among‐sub‐basin scale. These results indicate that environmental filtering is likely to be more important at the smallest scale with highest connectivity, while dispersal limitation seems to be more important at the largest scale with lowest connectivity. Community variation at the among‐stream and among‐sub‐basin scales were strongly explained by geographical and topographical distances, indicating that overland pathways might be the main dispersal route at the larger scales among more isolated sites. The relative effect of environmental and spatial factors on insect communities varied between low and high dispersal ability groups; this variation was inconsistent among three hierarchical scales. In sum, our study indicates that spatial scale, connectivity, and dispersal ability jointly shape stream metacommunities.     

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.     

Phytoplankton community response to hydrological variations in oxbow lakes with different levels of connection to a tropical river

Changes to the structure of the phytoplankton community and to the physical and chemical variables of the water were investigated in oxbow lakes with different levels of connection to a tropical river and subject to annual hydrological pulse variations. The selected lentic environments are located at the mouth region of the main tributary in a reservoir built for water storage and electric power generation. The temporal variation of phytoplankton in the studied lentic environments can be attributed mainly to the hydrological level of the river. A similar variation pattern of the ecological attributes was observed in the structure of the phytoplankton community in the connected lakes and Paranapanema River, evidencing the high degree of association that the lacustrine systems maintain with the river. The highest values of richness and diversity for connected environments were observed at the end of the emptying period and in the drought. However, considering the isolated lake, the highest values of these attributes were recorded during the flooding period.     

Macrozoobenthos patterns along environmental gradients and hydrological connectivity of oxbow lakes

Hydrological connectivity and the frequency and intensity of floods are the key factors determining the structure of macroinvertebrates inhabiting wetland ecosystems in river valleys. In 2007, water and macroinvertebrate samples were collected on four occasions in the middle course of the S?upia River and in five oxbow lakes (Northern Poland) to determine the hydrological relations in a regulated lowland river environment marked by a moderate climate. The water bodies selected for the study featured different types of connections with the main river valley: two of them were completely cut off from the valley, one was connected via a single branch, one featured a forced-flow connection through drainage pipes, and one was connected by a system of drainage channels. Macroinvertebrates, mostly Chironomidae larvae, were predominant in the eutrophic waters of the river. The prevalent macroinvertebrates found in the eutrophicated oxbow lakes isolated from the river were Chironomidae larvae and Crustacea (mainly Asellus aquaticus). In unobstructed oxbow lakes, the main component of benthic fauna was Crustacea, while Ephemeroptera were found mostly in the water body connected to the river via a drainage channel. A canonical correspondence analysis (CCA) showed that hydrological connectivity was the main factor responsible for the structure of invertebrate populations, followed by the physical and chemical parameters of the local environment. A non-conformance analysis revealed that hydrological connectivity enhanced invertebrate abundance and biological diversity, while the overall abundance was marked by unimodal distribution. The developed general model indicates that in the group of measured environmental variables, nitrite concentrations were highly correlated with Shannon diversity and invertebrate composition, while sulphate levels were closely associated with invertebrate abundance in the waters of the analyzed ecosystems.