Beta diversity of diatom species and ecological guilds : Response to environmental and spatial mechanisms along the stream watercourse

1. Understanding the mechanisms that drive beta diversity (i.e. beta-diversity), an important aspect of regional biodiversity, remains a priority for ecological research. beta-diversity and its components can provide insights into the processes generating regional biodiversity patterns. We tested whether environmental filtering or dispersal related processes predominated along the stream watercourse by analysing the responses of taxonomic and functional diatom beta-diversity to environmental and spatial factors. 2. We examined the variation in total beta-diversity and its components (turnover and nestedness) in benthic diatom species and ecological guilds (motile, planktonic, low-and high profile) with respect to watercourse position (up-, mid-and downstream) in 2,182 sites throughout France. We tested the effects of pure environmental and pure spatial factors on beta-diversity with partial Mantel tests. Environmental factors included eight physicochemical variables, while geographical distances between sites were used as spatial factors. We also correlated a and c-diversity, and the degree of nestedness (NODF metric) with environmental variables. 3. Total beta-diversity and its turnover component displayed higher values upstream than mid-and downstream. The nestedness component exhibited low values, even when NODF values increased from up-to downstream. Pure environmental factors were highly significant for explaining total beta-diversity and turnover regardless of watercourse position, but pure spatial factors were mostly significant mid-and downstream, with geographical distance being positively correlated with beta-diversity. Across sites, nutrient enrichment decreased turnover but increased the degree of nestedness. Motile and low profile diatoms comprised the most abundant guilds, but their beta-diversity patterns varied in an opposite way. The lowest guild beta-diversity was observed upstream for low profile species, and downstream for motile species. 4. In conclusion, environmental filtering seemed to play a major role in structuring metacommunities irrespective of site watercourse position. Filtering promoted strong turnover patterns, especially in disconnected upstream sites. The greater role of spatial factors mid-and downstream was consistent with mass effects rather than neutral processes because these sites had lower total beta-diversity than upstream sites. Motile species were most strongly affected by mass effects processes, whereas low profile species were primarily influenced by environmental conditions. Collectively, these findings suggest that partitioning of total beta-diversity into its components and the use of diatom ecological guilds provide a useful framework for assessing the mechanisms underlying metacommunity patterns along the stream watercourse.Peer reviewe

Environmental filtering and taxonomic relatedness underlie the species richness-evenness relationship

We examined the relationship between species richness (S) and evenness (J) within a novel, community assembly framework. We hypothesized that environmental stress leads to filtering (increasing the proportional abundance of tolerant species) and taxonomic dispersion (decreasing the number of species within genera and families). Environmental filtering would cause a decline in S by eliminating some stress-sensitive species and a reduction of J by allowing only tolerant species to maintain large populations. Taxonomic relatedness may influence both S and J by controlling the nature of interspecific interactions-positive under taxonomic dispersion versus negative under taxonomic clustering. Therefore, the S-J relationship may be a product of environmental filtering and taxonomic relatedness. We tested this framework with redundancy analyses and structural equation models using continental stream diatom and fish data. We confirmed that (i) environmental stress, defined by watershed forest cover, slope, and temperature, caused filtering (lower sensitive: tolerant species abundance ratios) and taxonomic dispersion (elevated genus: species richness and family: species richness ratios); (ii) S and J, which declined with filtering and taxonomic dispersion, exhibited a positive relationship; and (iii) the role of filtering on J was pronounced only under stressful conditions, while taxonomic dispersion remained an important predictor of J across stressful and favorable environments.Peer reviewe

Biogeographical Patterns of Species Richness and Abundance Distribution in Stream Diatoms Are Driven by Climate and Water Chemistry

In this intercontinental study of stream diatoms, we asked three important but still unresolved ecological questions: (1) What factors drive the biogeography of species richness and species abundance distribution (SAD)? (2) Are climate-related hypotheses, which have dominated the research on the latitudinal and altitudinal diversity gradients, adequate in explaining spatial biotic variability? and (3) Is the SAD response to the environment independent of richness? We tested a number of climatic theories and hypotheses (i.e., the species-energy theory, the metabolic theory, the energy variability hypothesis, and the climatic tolerance hypothesis) but found no support for any of these concepts, as the relationships of richness with explanatory variables were nonexistent, weak, or unexpected. Instead, we demonstrated that diatom richness and SAD evenness generally increased with temperature seasonality and at mid- to high total phosphorus concentrations. The spatial patterns of diatom richness and the SADmainly longitudinal in the United States but latitudinal in Finlandwere defined primarily by the covariance of climate and water chemistry with space. The SAD was not entirely controlled by richness, emphasizing its utility for ecological research. Thus, we found support for the operation of both climate and water chemistry mechanisms in structuring diatom communities, which underscores their complex response to the environment and the necessity for novel predictive frameworks.Peer reviewe

Dispersal-niche continuum index : a new quantitative metric for assessing the relative importance of dispersal versus niche processes in community assembly

Patterns in community composition are scale-dependent and generally difficult to distinguish. Therefore, quantifying the main assembly processes in various systems and across different datasets has remained challenging. Building on the PER-SIMPER method, we propose a new metric, the dispersal-niche continuum index (DNCI), which estimates whether dispersal or niche processes dominate community assembly and facilitates the comparisons of processes among datasets. The DNCI was tested for robustness using simulations and applied to observational datasets comprising organismal groups with different trophic level and dispersal potential. Based on the robustness tests, the DNCI discriminated the respective contribution of niche and dispersal processes in pairwise comparisons of site groups with less than 40% and 30% differences in their taxa and site numbers, respectively. In the observational datasets, the DNCI suggested that dispersal rather than niche assembly was the dominant assembly process which, however, varied in intensity among organismal groups and study contexts, including spatial scale and ecosystem types. The proposed DNCI measures the relative strength of community assembly processes in a way that is simple, easily quantifiable and comparable across datasets. We discuss the strengths and weaknesses of the DNCI and provide perspectives for future research.Peer reviewe

Stream diatom community assembly processes in islands and continents: a global perspective

[EN] Understanding the roles of deterministic and stochastic processes in community assembly is essential for gaining insights into the biogeographical patterns of biodiversity. However, the way community assembly processes operate is still not fully understood, especially in oceanic islands. In this study, we examine the importance of assembly processes in shaping diatom communities in islands and continents, while also investigating the influence of climate and local water chemistry variables on species distributions. Location Global. Taxon Stream benthic diatoms. Methods We used diatom datasets from five continents and 19 islands and applied beta diversity analyses with a null model approach and hierarchical joint species distribution modelling. To facilitate comparisons with continents, we used continental area equivalents (CAEs), which represent continental subsets with comparable areas and the same number of study sites as their corresponding islands counterparts. Results We found that homogeneous selection (i.e., communities being more similar than the random expectation) was the dominant assembly process within islands whereas stochastic processes tended to be more important within continents. In addition, assembly processes were influenced by study scale and island isolation. Climatic variables showed a greater influence on species distribution than local factors. However, in islands, local environmental variables had a greater impact on the distributions of unique taxa as opposed to non-unique taxa. Main Conclusions We observed that the assembly processes of diatom communities were complex and influenced by a combination of deterministic and stochastic forces, which varied across spatial scales. In islands, there was no universal pattern of assembly processes, given that their influence depends on abiotic conditions such as area, isolation, and environmental heterogeneity. In addition, the sensitivity of species occurring uniquely in islands to local environmental variables suggests that they are perhaps less vulnerable to climatic changes but may be more influenced by changes in local physicochemistrySIFor financial support, the authors thank the Academy of Finland (grant nr. 346812 to JS); the Institut Francais de Finlande; the Embassy of France to Finland; the French Ministry of Education and Higher Education; Finnish Society of Sciences and Letters. J.J. Wang was further supported by the National Natural Science Foundation of China (91851117, 41871048), CAS Key Research Program of Frontier Sciences (QYZDB-SSW-DQC043), and The National Key Research and Development Program of China (2019YFA0607100

Data from: Abundance inequality in freshwater communities has an ecological origin

The hollow-shaped species abundance distribution (SAD) and its allied rank abundance distribution (RAD)—showing that abundance is unevenly distributed among species—are some of the most studied patterns in ecology. To explain the nature of abundance inequality, I developed a novel framework identifying environmental favorability, which controls the balance between reproduction and immigration, as the ultimate source and species stress tolerance as a proximate factor. Thus, under harsh conditions, only a few tolerant species can reproduce, while some sensitive species can be present in low numbers due to chance immigration. This would lead to high abundance inequality between the two groups of species. Under benign conditions, both groups can reproduce and give rise to higher abundance equality. To test these ideas, I examined the variability in the parameters of a Poisson lognormal fit of the SAD and a square root fit of the RAD in diatom and fish communities across US streams. Indeed, as environmental favorability increased, more sensitive forms were able to establish large populations, diminishing the abundance disparity between locally common and rare species. Finally, it was demonstrated that in diatoms, the RAD belonged to the same family of relationships as those of population density with body size and regional distribution

Framework for community functioning: synthesis of stress gradient and resource partitioning concepts

To understand how communities function and generate abundance, I develop a framework integrating elements from the stress gradient and resource partitioning concepts. The framework suggests that guild abundance depends on environmental and spatial factors but also on inter-guild interactions (competitor or facilitator richness), which can alter the fundamental niche of constituent species in negative (competition) or positive direction (facilitation). Consequently, the environmental and spatial mechanisms driving guild abundance would differ across guilds and interaction modes. Using continental data on stream diatoms and physico-chemistry, the roles of these mechanisms were tested under three interaction modes—shared preference, distinct preference, and facilitative, whereby pairs of guilds exhibited, respectively, a dominance-tolerance tradeoff along a eutrophication gradient, specialization along a pH gradient, or a donor-recipient relationship along a nitrogen gradient. Representative of the shared preference mode were the motile (dominant) and low profile (tolerant) guilds, of the distinct preference mode—the acidophilous and alkaliphilous (low profile) guilds, and of the facilitative mode—nitrogen fixers (donors) and motile species (recipients). In each mode, the influences of environment, space (latitude and longitude), and competitor or facilitator richness on guild density were assessed by variance partitioning. Pure environment constrained most strongly the density of the dominant, the acidophilous, and the recipient guild in the shared preference, distinct preference, and facilitative mode, respectively, while spatial effects were important only for the low profile guild. Higher competitor richness was associated with lower density of the tolerant guild in the shared preference mode, both guilds in the distinct preference mode, and the donor guild in the facilitative mode. Conversely, recipient density in the facilitative mode increased with donor richness in stressful nitrogen-poor environments. Thus, diatom guild abundance patterns were determined primarily by biotic and/or environmental impacts and, with the exception of the low profile guild, were insensitive to spatial effects. This framework identifies major sources of variability in diatom guild abundance with implications for the understanding of biodiversity-ecosystem functioning

Fish data

The file “Fish data” contains 3 Excel spreadsheets, i.e. “Raw data,” “Community data,” and “Species data.” The Raw data spreadsheet shows the abundance, measured as number of individuals, and the abundance rank for all species (BioDataTaxonName) across 761 fish samples, listed by their sample identification. The fish samples were collected from 399 distinct stream localities, listed by their station identification. The Community data spreadsheet contains relevant information for the fish samples, including latitude, longitude, date of collection, species richness, evenness, total abundance (number of individuals), statistics of the Poisson lognormal fit to the sample species abundance distribution (parameters mu and sigma, gof (goodness of fit), and p (proportion of species revealed by the sample)), statistics of the square root fit to the sample rank-abundance distribution (intercept a0, slope a, and their standard errors and 95% confidence intervals), ln Tolerant:Intolerant ratio, and abundance and richness of the three fish tolerance groups (tolerant, intolerant, and moderate). Environmental data were available for a subset of 732 samples. The Species data spreadsheet lists the 460 fish species (BioDataTaxonName) encountered in this study with tolerance group designation whenever known and occurrence across 1005 stream localities

Diatom data

The file “Diatom data” contains 3 Excel spreadsheets, i.e. “Raw data,” “Community data,” and “Species data.” The Raw data spreadsheet shows the population density, measured in cells/cm^2, and the population density rank for all species (BioDataTaxonName) across 3213 diatom samples, listed by their sample identification. The diatom samples were collected from 1580 distinct stream localities, listed by their station identification. The Community data spreadsheet contains relevant information for the diatom samples, including latitude, longitude, date of collection, species richness, evenness, total density (cells/cm^2), statistics of the Poisson lognormal fit to the sample species abundance distribution (parameters mu and sigma, gof (goodness of fit), and p (proportion of species revealed by the sample)), statistics of the square root fit to the sample rank-abundance distribution (intercept a0, slope a, and their standard errors and 95% confidence intervals), ln Tolerant:Sensitive ratio, and density and richness of the three diatom guilds (low profile, motile, and high profile). Environmental data were available for a subset of 2949 samples. The Species data spreadsheet lists the 1777 diatom species (BioDataTaxonName) encountered in this study with guild designation whenever possible and occurrence across 1582 stream localities