The structure of bacteria-fungi bipartite networks along elevational gradients in contrasting climates.

Climate change is altering species distribution and modifying interactions in microbial communities. Understanding microbial community structure and their interactions is crucial to interpreting ecosystem responses to climate change. Here, we examined the assemblages of stream bacteria and fungi, and the associations between the two groups along elevational gradients in two regions with contrasting precipitation and temperature, that is the Galong and Qilian mountains of the Tibetan Plateau. In the wetter and warmer region, the species richness significantly increased and decreased with elevation for bacteria and fungi, respectively, while were nonsignificant in the drier and colder region. Their bipartite network structure was also different by showing significant increases in connectance and nestedness towards higher elevations only in the wetter and warmer region. In addition, these correlation network structure generally exhibited similar positive association with species richness in the wetter and warmer region and the drier and colder region. In the wetter and warmer region, climatic change along elevation was more important in determining connectance and nestedness, whereas microbial species richness exerted a stronger influence on network structure and robustness in the drier and colder region. These findings indicate substantial forthcoming changes in microbial diversity and network structure in warming climates, especially in wetter and warmer regions on Earth, advancing the understanding of microbial bipartite interactions' response to climate change.

Embracing mountain microbiome and ecosystem functions under global change.

Mountains are pivotal to maintaining habitat heterogeneity, global biodiversity, ecosystem functions and services to humans. They have provided classic model natural systems for plant and animal diversity gradient studies for over 250 years. In the recent decade, the exploration of microorganisms on mountainsides has also achieved substantial progress. Here, we review the literature on microbial diversity across taxonomic groups and ecosystem types on global mountains. Microbial community shows climatic zonation with orderly successions along elevational gradients, which are largely consistent with traditional climatic hypotheses. However, elevational patterns are complicated for species richness without general rules in terrestrial and aquatic environments and are driven mainly by deterministic processes caused by abiotic and biotic factors. We see a major shift from documenting patterns of biodiversity towards identifying the mechanisms that shape microbial biogeographical patterns and how these patterns vary under global change by the inclusion of novel ecological theories, frameworks and approaches. We thus propose key questions and cutting-edge perspectives to advance future research in mountain microbial biogeography by focusing on biodiversity hypotheses, incorporating meta-ecosystem framework and novel key drivers, adapting recently developed approaches in trait-based ecology and manipulative field experiments, disentangling biodiversity-ecosystem functioning relationships and finally modelling and predicting their global change responses.

Microbial and Environmental Processes Shape the Link between Organic Matter Functional Traits and Composition.

Dissolved organic matter (DOM) is a large and complex mixture of molecules that fuels microbial metabolism and regulates biogeochemical cycles. Individual DOM molecules have unique functional traits, but how their assemblages vary deterministically under global change remains poorly understood. Here, we examine DOM and associated bacteria in 300 aquatic microcosms deployed on mountainsides that span contrasting temperatures and nutrient gradients. Based on molecular trait dimensions of reactivity and activity, we partition the DOM composition into labile-active, recalcitrant-active, recalcitrant-inactive, and labile-inactive fractions and quantify the relative influences of deterministic and stochastic processes governing the assembly of each. At both subtropical and subarctic study sites, the assembly of labile or recalcitrant molecules in active fractions is primarily governed by deterministic processes, while stochastic processes are more important for the assembly of molecules within inactive fractions. Surprisingly, the importance of deterministic selection increases with global change gradients for recalcitrant molecules in both active and inactive fractions, and this trend is paralleled by changes in the deterministic assembly of microbial communities and environmental filtering, respectively. Together, our results highlight the shift in focus from potential reactivity to realized activity and indicate that active and inactive fractions of DOM assemblages are structured by contrasting processes, and their recalcitrant components are consistently sensitive to global change. Our study partitions the DOM molecular composition across functional traits and links DOM with microbes via a shared ecological framework of assembly processes. This integrated approach opens new avenues to understand the assembly and turnover of organic carbon in a changing world.

Anthropogenic land‒use impacts on the size structure of macroinvertebrate assemblages are jointly modulated by local conditions and spatial processes.

Body size descriptors and associated resemblance measurements may provide useful tools for forecasting ecological responses to increasing anthropogenic land‒use disturbances. Yet, the influences of agriculture and urbanisation on the size structure of biotic assemblages have seldom been investigated in running waters. Using a comprehensive dataset on stream macroinvertebrates from 21 river basins across Western Finland, we assessed whether the structure of assemblages via changes in taxonomic composition and body size distributions responded predictably to anthropogenic land‒use impacts. Specifically, we applied a combination of resemblance measurements based on cumulative abundance profiles and spatially constrained null models to understand faunal impairment by agricultural and urban development, and the most likely mechanisms underlying the observed shifts in assemblage size structure. Anthropogenically impacted stream sites showed less variation in assemblage composition and size distributions compared with least‒disturbed sites, with strong declines in internal variation also occurring for the transition from near‒pristine to moderately impacted landscapes. These results were consistent whether based on species‒level or genus‒level data. Variation in assemblage size structure seemed to be more predictable than taxonomic composition, supporting the notion that resemblance measurements based on body size distributions can represent an improvement to more traditional approaches based on taxonomic identities alone. In addition, we showed that macroinvertebrate assemblages resulted from effects of land‒use degradation mediated through local conditions and spurious spatial structures in the distribution of anthropogenic activities across the landscape. Overall, our findings suggest that existing water policies and agri‒environment schemes should be guided not only by understanding the individual effects of agricultural and urban development on taxonomic composition at a given stream site. Rather, we should also acknowledge the size structure of stream assemblages and whether concomitant changes in local conditions and the non‒random distribution of human infrastructures are likely to mitigate or accelerate these effects.

Does trait-based joint species distribution modelling reveal the signature of competition in stream macroinvertebrate communities?

The occupancy and abundance of species are jointly driven by local factors, such as environmental characteristics and biotic interactions, and regional-scale factors, such as dispersal and climate. Recently, it has been shown that biotic interactions shape species occupancies and abundances beyond local extents. However, for small ectothermic animals, particularly for those occurring in freshwater environments, the importance of biotic interactions remains understudied. Species-to-species associations from joint species distribution models (i.e. species associations while controlling for environmental characteristics) are increasingly used to draw hypotheses of which species possibly show biotic interactions. We studied whether species-to-species associations from joint species distribution models show signs of competition using a hypothesis testing framework in stream macroinvertebrate communities at regional extent. We sampled aquatic macroinvertebrates from 105 stream sites in western Finland encompassing a latitudinal gradient of c. 500 kilometres. We hypothesized that if competition drives these associations (H1) functionally, similar species are mostly negatively associated, whereas functionally dissimilar species show random associations. We further hypothesized that the relationship between functional dissimilarity and the strength of association is more pronounced (H2) for abundances rather than occupancies, (H3) at small grain (i.e. stream site) rather than at large grain (i.e. river basin), and (H4) among species having weak dispersal ability than among species with high dispersal ability. Stream macroinvertebrates showed both negative and positive species-to-species associations while controlling for habitat characteristics. However, the negative associations were mostly at large grain (river basin) rather than at small grain (stream site), in occupancy rather than abundance, and not related to species functional dissimilarity or to their dispersal ability. Thus, all our hypotheses considering possible competition (H1-H4) were rejected. Competition does not appear to be a major driving force of stream macroinvertebrate communities at the spatial grain sizes considered. The observed positive associations in occupancy at small grain (stream site) may be attributed to species' similar microhabitat preferences, whereas at large grain (river basin), they may stem from metacommunity dynamics. Our results highlight that species traits were necessary to interpret whether or not species-to-species associations from joint species distribution models resulted from biotic interactions.

Biodiversity Loss Threatens the Current Functional Similarity of Beta Diversity in Benthic Diatom Communities.

The global biodiversity loss has increased the need to understand the effects of decreasing diversity, but our knowledge on how species loss will affect the functioning of communities and ecosystems is still very limited. Here, the levels of taxonomic and functional beta diversity and the effect of species loss on functional beta diversity were investigated in an estuary that provides a naturally steep environmental gradient. The study was conducted using diatoms that are among the most important microorganisms in all aquatic ecosystems and globally account for 40% of marine primary production. Along the estuary, the taxonomic beta diversity of diatom communities was high (Bray-Curtis taxonomic similarity 0.044) and strongly controlled by the environment, particularly wind exposure, salinity, and temperature. In contrast, the functional beta diversity was low (Bray-Curtis functional similarity 0.658) and much less controlled by the environment. Thus, the diatom communities stayed functionally almost similar despite large changes in species composition and environment. This may indicate that, through high taxonomic diversity and redundancy in functions, microorganisms provide an insurance effect against environmental change. However, when studying the effect of decreasing species richness on functional similarity of communities, simulated species loss to 45% of the current species richness decreased functional similarity significantly. This suggests that decreasing species richness may increase variability and reduce the stability and resilience of communities. These results highlight the importance of high taxonomic biodiversity for the stable functioning of benthic communities.

A Metacommunity Approach to Improve Biological Assessments in Highly Dynamic Freshwater Ecosystems.

Rapid shifts in biotic communities due to environmental variability challenge the detection of anthropogenic impacts by current biomonitoring programs. Metacommunity ecology has the potential to inform such programs, because it combines dispersal processes with niche-based approaches and recognizes variability in community composition. Using intermittent rivers-prevalent and highly dynamic ecosystems that sometimes dry-we develop a conceptual model to illustrate how dispersal limitation and flow intermittence influence the performance of biological indices. We produce a methodological framework integrating physical- and organismal-based dispersal measurements into predictive modeling, to inform development of dynamic ecological quality assessments. Such metacommunity-based approaches could be extended to other ecosystems and are required to underpin our capacity to monitor and protect ecosystems threatened under future environmental changes.

Elevational patterns and hierarchical determinants of biodiversity across microbial taxonomic scales.

Microbial biogeography is gaining increasing attention due to recent molecular methodological advance. However, the diversity patterns and their environmental determinants across taxonomic scales are still poorly studied. By sampling along an extensive elevational gradient in subarctic ponds of Finland and Norway, we examined the diversity patterns of aquatic bacteria and fungi from whole community to individual taxa across taxonomic coverage and taxonomic resolutions. We further quantified cross-phylum congruence in multiple biodiversity metrics and evaluated the relative importance of climate, catchment and local pond variables as the hierarchical drivers of biodiversity across taxonomic scales. Bacterial community showed significantly decreasing elevational patterns in species richness and evenness, and U-shaped patterns in local contribution to beta diversity (LCBD). Conversely, no significant species richness and evenness patterns were found for fungal community. Elevational patterns in species richness and LCBD, but not in evenness, were congruent across bacterial phyla. When narrowing down the taxonomic scope towards higher resolutions, bacterial diversity showed weaker and more complex elevational patterns. Taxonomic downscaling also indicated a notable change in the relative importance of biodiversity determinants with stronger local environmental filtering, but decreased importance of climatic variables. This suggested that niche conservatism of temperature preference was phylogenetically deeper than that of water chemistry variables. Our results provide novel perspectives for microbial biogeography and highlight the importance of taxonomic scale dependency and hierarchical drivers when modelling biodiversity and species distribution responses to future climatic scenarios.

Are drivers of microbial diatom distributions context dependent in human-impacted and pristine environments?

Species occurrences are influenced by numerous factors whose effects may be context dependent. Thus, the magnitude of the effects and their relative importance to species distributions may vary among ecosystems due to anthropogenic stressors. To investigate context dependency in factors governing microbial bioindicators, we developed species distribution models (SDMs) for epilithic stream diatom species in human-impacted and pristine sites separately. We performed SDMs using boosted regression trees for 110 stream diatom species, which were common to both data sets, in 164 human-impacted and 164 pristine sites in Finland (covering ~1,000 km, 60° to 68° N). For each species and site group, two sets of models were conducted: climate model, comprising three climatic variables, and full model, comprising the climatic and six local environmental variables. No significant difference in model performance was found between the site groups. However, climatic variables had greater importance compared with local environmental variables in pristine sites, whereas local environmental variables had greater importance in human-impacted sites as hypothesized. Water balance and conductivity were the key variables in human-impacted sites. The relative importance of climatic and local environmental variables varied among individual species, but also between the site groups. We found a clear context dependency among the variables influencing stream diatom distributions as the most important factors varied both among species and between the site groups. In human-impacted streams, species distributions were mainly governed by water chemistry, whereas in pristine streams by climate. We suggest that climatic models may be suitable in pristine ecosystems, whereas the full models comprising both climatic and local environmental variables should be used in human-impacted ecosystems.

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 SAD-mainly longitudinal in the United States but latitudinal in Finland-were 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.

Thermal barriers constrain microbial elevational range size via climate variability.

Range size is invariably limited and understanding range size variation is an important objective in ecology. However, microbial range size across geographical gradients remains understudied, especially on mountainsides. Here, the patterns of range size of stream microbes (i.e., bacteria and diatoms) and macroorganisms (i.e., macroinvertebrates) along elevational gradients in Asia and Europe were examined. In bacteria, elevational range size showed non-significant phylogenetic signals. In all taxa, there was a positive relationship between niche breadth and species elevational range size, driven by local environmental and climatic variables. No taxa followed the elevational Rapoport's rule. Climate variability explained the most variation in microbial mean elevational range size, whereas local environmental variables were more important for macroinvertebrates. Seasonal and annual climate variation showed negative effects, while daily climate variation had positive effects on community mean elevational range size for all taxa. The negative correlation between range size and species richness suggests that understanding the drivers of range is key for revealing the processes underlying diversity. The results advance the understanding of microbial species thermal barriers by revealing the importance of seasonal and diurnal climate variation, and highlight that aquatic and terrestrial biota may differ in their response to short- and long-term climate variability.

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.

Eutrophication increases the similarity of cyanobacterial community features in lakes and reservoirs.

Eutrophication of inland waters is a mostly anthropogenic phenomenon impacting aquatic biodiversity worldwide, and might change biotic community structure and ecosystem functions. However, little is known about the patterns of cyanobacterial community variations and changes both on alpha and beta diversity levels in response to eutrophication. Here, we investigated cyanobacterial communities sampled at 140 sites from 59 lakes and reservoirs along a strong eutrophication gradient in eastern China through using CPC-IGS and 16S rRNA gene amplicon sequencing. We found that taxonomic diversity increased, but phylogenetic diversity decreased significantly along the eutrophication gradient. Both niche width and niche overlap of cyanobacteria significantly decreased from low- to high-nutrient waterbodies. Cyanobacterial community distance-decay relationship became weaker from mesotrophic to hypereutrophic waterbodies, while ecological uniqueness (i.e., local contributions to beta diversity) tended to increase in high-nutrient waterbodies. Latitude and longitude were more important in shaping cyanobacterial community structure than other environmental variables. These findings suggest that eutrophication affects alpha and beta diversity of cyanobacterial communities, leading to increasingly similar community structures in lakes and reservoirs with a higher level of eutrophication. Our work highlights how cyanobacterial communities respond to anthropogenic eutrophication and calls for an urgent need to develop conservation and management strategies to control lake eutrophication and protect freshwater biodiversity.

Stochastic species distributions are driven by organism size.

The strengths of environmental drivers and biotic interactions are expected to show large variability across organism groups. We tested two ideas related to the degree of ecological determinism vs. stochasticity using a large data set comprising bacterio-, phyto-, and zooplankton. We expected that (1) there are predictable, size-driven differences in the degree to which planktonic taxa respond to different drivers such as water chemistry, biotic interactions, and climatic variables; and (2) species distribution models show lowest predictive performance for the smallest taxa due to the stochastic distributions of microbes. Generalized linear models (GLMs), generalized additive models (GAMs), and generalized boosted methods (GBMs) were constructed for 84 species to model their occurrence as a function of eight predictors. Predictive performance was measured as the area under the curve (AUC) of the receiver-operating characteristic plot and true skill statistic (TSS) using independent model evaluation data. We found that the model performances were typically remarkably low for all planktonic groups. The proportion of satisfactory models (AUC > 0.7) was lowest for bacteria (11.1% of the models), followed by phyto- (24.2%) and zooplankton (38.1%). The occurrences of taxa within all planktonic groups were related to climatic variables to a certain degree, but bacteria showed the strongest associations with the climatic variables. Moreover, zooplankton occurrences were more related to biotic variables than the occurrences of smaller taxa, while phytoplankton occurrences were more related to water chemistry. We conclude that the occurrences of planktonic taxa are highly unpredictable and that stochasticity in occurrences is negatively related to the organism size perhaps due to efficient dispersal and fast population dynamics among the smallest taxa.

Seasonal variation of phytoplankton community assembly processes in Tibetan Plateau floodplain.

Uncovering the mechanisms underlying phytoplankton community assembly remains a major challenge in freshwater ecology. The roles of environmental filtering and spatial processes in shaping phytoplankton metacommunity in Tibetan floodplain ecosystems under various hydrological conditions are still unclear. Here, multivariate statistics and a null model approach were used to compare the spatiotemporal patterns and assembly processes of phytoplankton communities in the river-oxbow lake system of Tibetan Plateau floodplain between non-flood and flood periods. The results showed that phytoplankton communities had significant seasonal and habitat variations, with the seasonal variations being more remarkable. Phytoplankton density, biomass, and alpha diversity were distinctly lower in the flood than non-flood period. The habitat differences (rivers vs. oxbow lakes) in phytoplankton community were less pronounced during the flood than non-flood period, most likely due to the increased hydrological connectivity. There was a significant distance-decay relationship only in lotic phytoplankton communities, and such relationship was stronger in the non-flood than flood period. Variation partitioning and PER-SIMPER analysis showed that the relative role of environmental filtering and spatial processes affecting phytoplankton assemblages varied across hydrological periods, with environmental filtering dominating in the non-flood period and spatial processes in the flood period. These results suggest that the flow regime plays a key role in balancing environmental and spatial factors in shaping phytoplankton communities. This study contributes to a deeper understanding of ecological phenomena in highland floodplains and provides a theoretical basis for floodplain ecosystem maintenance and ecological health management.

Local eukaryotic and bacterial stream community assembly is shaped by regional land use effects.

With anticipated expansion of agricultural areas for food production and increasing intensity of pressures stemming from land-use, it is critical to better understand how species respond to land-use change. This is particularly true for microbial communities which provide key ecosystem functions and display fastest responses to environmental change. However, regional land-use effects on local environmental conditions are often neglected, and, hence, underestimated when investigating community responses. Here we show that the effects stemming from agricultural and forested land use are strongest reflected in water conductivity, pH and phosphorus concentration, shaping microbial communities and their assembly processes. Using a joint species distribution modelling framework with community data based on metabarcoding, we quantify the contribution of land-use types in determining local environmental variables and uncover the impact of both, land-use, and local environment, on microbial stream communities. We found that community assembly is closely linked to land-use type but that the local environment strongly mediates the effects of land-use, resulting in systematic variation of taxon responses to environmental conditions, depending on their domain (bacteria vs. eukaryote) and trophic mode (autotrophy vs. heterotrophy). Given that regional land-use type strongly shapes local environments, it is paramount to consider its key role in shaping local stream communities.

On the shape and origins of the freshwater species-area relationship.

The species-area relationship (SAR) has over a 150-year-long history in ecology, but how its shape and origins vary across scales and organisms remains incompletely understood. This is the first subcontinental freshwater study to examine both these properties of the SAR in a spatially explicit way across major organismal groups (diatoms, insects, and fish) that differ in body size and dispersal capacity. First, to describe the SAR shape, we evaluated the fit of three commonly used models, logarithmic, power, and Michaelis-Menten. Second, we proposed a hierarchical framework to explain the variability in the SAR shape, captured by the parameters of the SAR model. According to this framework, scale and species group were the top predictors of the SAR shape, climatic factors (heterogeneity and median conditions) represented the second predictor level, and metacommunity properties (intraspecific spatial aggregation, γ-diversity, and species abundance distribution) the third predictor level. We calculated the SAR as a sample-based rarefaction curve using 60 streams within landscape windows (scales) in the United States, ranging from 160,000 to 6,760,000 km2 . First, we found that all models provided good fits (R2 ≥ 0.93), but the frequency of the best-fitting model was strongly dependent on organism, scale, and metacommunity properties. The Michaelis-Menten model was most common in fish, at the largest scales, and at the highest levels of intraspecific spatial aggregation. The power model was most frequent in diatoms and insects, at smaller scales, and in metacommunities with the lowest evenness. The logarithmic model fit best exclusively at the smallest scales and in species-poor metacommunities, primarily fish. Second, we tested our framework with the parameters of the most broadly used SAR model, the log-log form of the power model, using a structural equation model. This model supported our framework and revealed that the SAR slope was best predicted by scale- and organism-dependent metacommunity properties, particularly spatial aggregation, whereas the intercept responded most strongly to species group and γ-diversity. Future research should investigate from the perspective of our framework how shifts in metacommunity properties due to climate change may alter the SAR.

European river typologies fail to capture diatom, fish, and macrophyte community composition.

Typology systems are frequently used in applied and fundamental ecology and are relevant for environmental monitoring and conservation. They aggregate ecosystems into discrete types based on biotic and abiotic variables, assuming that ecosystems of the same type are more alike than ecosystems of different types with regard to a specific property of interest. We evaluated whether this assumption is met by the Broad River Types (BRT), a recently proposed European river typology system, that classifies river segments based on abiotic variables, when it is used to group biological communities. We compiled data on the community composition of diatoms, fishes, and aquatic macrophytes throughout Europe and evaluated whether the composition is more similar in site groups with the same river type than in site groups of different river types using analysis of similarities, classification strength, typical species analysis, and the area under zeta diversity decline curves. We compared the performance of the BRT with those of four region-based typology systems, namely, Illies Freshwater Ecoregions, the Biogeographic Regions, the Freshwater Ecoregions of the World, and the Environmental Zones, as well as spatial autocorrelation (SA) classifications. All typology systems received low scores from most evaluation methods, relative to predefined thresholds and the SA classifications. The BRT often scored lowest of all typology systems. Within each typology system, community composition overlapped considerably between site groups defined by the types of the systems. The overlap tended to be the lowest for fishes and between Illies Freshwater Ecoregions. In conclusion, we found that existing broad-scale river typology systems fail to delineate site groups with distinct and compositionally homogeneous communities of diatoms, fishes, and macrophytes. A way to improve the fit between typology systems and biological communities might be to combine segment-based and region-based typology systems to simultaneously account for local environmental variation and historical distribution patterns, thus potentially improving the utility of broad-scale typology systems for freshwater biota.

Is catchment productivity a useful predictor of taxa richness in lake plankton communities?

The influence of catchment variables on lake organisms is understudied. The terrestrial zone in the vicinity of lakes is, however, probably highly important for biota due to the effects on water chemistry and to various processes operating across ecosystem boundaries. We examined the relative importance of lake and catchment variables, as well as large-scale geographical factors, on the taxa richness of phyto- and zooplankton in 100 small lakes in Finland. In variation partitioning, the variability of phytoplankton richness was most strongly related to the effects of lake variables, the joint effects of lake and catchment variables, and the joint effects of all three groups of variables. Zooplankton richness, in turn, was most strongly related to the effects of lake and catchment variables and the joint effect of lake and catchment variables. The exact results of the variation partitioning depended on the catchment sizes considered in the regression models. Among lake variables, planktonic richness was strongly related to variables indicating productivity. Among catchment variables, the normalized difference vegetation index (NDVI), indicating catchment productivity, showed a relatively strong association with planktonic richness. These results provide evidence that catchment variables such as the NDVI may be efficient predictors of planktonic richness in small lakes. It is possible that individual lakes embedded in a highly productive landscape have higher taxa richness than solitary, potentially productive lakes because of the high influx of dispersing propagules from the regional pool. We also suggest that catchment variables may respond to environmental changes at different scales than the lake variables, and explicit consideration of catchment productivity would therefore be useful when planning research and monitoring programs for freshwater organisms.

The relationship between species richness and evenness: a meta-analysis of studies across aquatic ecosystems.

Biological diversity comprises both species richness, i.e., the number of species in a community, and evenness, measuring how similar species are in their abundances. The relationship between species richness and evenness (RRE) across communities remains, however, a controversial issue in ecology because no consistent pattern has been reported. We conducted a systematic meta-review of RRE in aquatic ecosystems along regional to continental gradients and across trophic groups, differing in body size by 13 orders of magnitude. Hypotheses that RRE responded to latitudinal and scale variability across trophic groups were tested by regression analyses. Significant correlations of species richness and evenness only existed in 71 out of 229 datasets. Among the RRE, 89 were negative and 140 were positive. RRE did not vary with latitude but showed a positive response to scale. In a meta-analysis with ecosystem type as a single explaining variable, RRE did not vary among ecosystem types, i.e. between marine and freshwater. Finally, autotrophs had more positive RRE than heterotrophs. The weak RRE in many aquatic datasets suggests that richness and evenness often reflect independent components of biodiversity, highlighting that richness alone may be an incomplete surrogate for biodiversity. Our results further elucidate that RRE is driven by organismal and environmental properties, both of which must be considered to gain a deeper understanding of large-scale patterns of biodiversity.