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.

Perceived multiple stressor effects depend on sample size and stressor gradient length.

Multiple stressors are continuously deteriorating surface waters worldwide, posing many challenges for their conservation and restoration. Combined effect types of multiple stressors range from single-stressor dominance to complex interactions. Identifying prevalent combined effect types is critical for environmental management, as it helps to prioritise key stressors for mitigation. However, it remains unclear whether observed single and combined stressor effects reflect true ecological processes unbiased by sample size and length of stressor gradients. Therefore, we examined the role of sample size and stressor gradient lengths in 158 paired-stressor response cases with over 120,000 samples from rivers, lakes, transitional and marine ecosystems around the world. For each case, we split the overall stressor gradient into two partial gradients (lower and upper) and investigated associated changes in single and combined stressor effects. Sample size influenced the identified combined effect types, and stressor interactions were less likely for cases with fewer samples. After splitting gradients, 40 % of cases showed a change in combined effect type, 30 % no change, and 31 % showed a loss in stressor effects. These findings suggest that identified combined effect types may often be statistical artefacts rather than representing ecological processes. In 58 % of cases, we observed changes in stressor effect directions after the gradient split, suggesting unimodal stressor effects. In general, such non-linear responses were more pronounced for organisms at higher trophic levels. We conclude that observed multiple stressor effects are not solely determined by ecological processes, but also strongly depend on sampling design. Observed effects are likely to change when sample size and/or gradient length are modified. Our study highlights the need for improved monitoring programmes with sufficient sample size and stressor gradient coverage. Our findings emphasize the importance of adaptive management, as stress reduction measures or further ecosystem degradation may change multiple stressor-effect relationships, which will then require associated changes in management strategies.

Lakes in the era of global change: moving beyond single-lake thinking in maintaining biodiversity and ecosystem services.

The Anthropocene presents formidable threats to freshwater ecosystems. Lakes are especially vulnerable and important at the same time. They cover only a small area worldwide but harbour high levels of biodiversity and contribute disproportionately to ecosystem services. Lakes differ with respect to their general type (e.g. land-locked, drainage, floodplain and large lakes) and position in the landscape (e.g. highland versus lowland lakes), which contribute to the dynamics of these systems. Lakes should be generally viewed as 'meta-systems', whereby biodiversity is strongly affected by species dispersal, and ecosystem dynamics are contributed by the flow of matter and substances among locations in a broader waterscape context. Lake connectivity in the waterscape and position in the landscape determine the degree to which a lake is prone to invasion by non-native species and accumulation of harmful substances. Highly connected lakes low in the landscape accumulate nutrients and pollutants originating from ecosystems higher in the landscape. The monitoring and restoration of lake biodiversity and ecosystem services should consider the fact that a high degree of dynamism is present at local, regional and global scales. However, local and regional monitoring may be plagued by the unpredictability of ecological phenomena, hindering adaptive management of lakes. Although monitoring data are increasingly becoming available to study responses of lakes to global change, we still lack suitable integration of models for entire waterscapes. Research across disciplinary boundaries is needed to address the challenges that lakes face in the Anthropocene because they may play an increasingly important role in harbouring unique aquatic biota as well as providing ecosystem goods and services in the future.

Assessing mining impacts: The influence of background geochemical conditions on diatom and macroinvertebrate communities in subarctic streams.

Mining has changed landscapes locally in northern Fennoscandia and there is an increasing pressure for exploitation of the remaining mineral deposits of the region. Mineral deposits, even if unmined, can strongly influence stream water chemistry, stream biological communities and the ability of organisms to tolerate stressors. Using data sampled from six mining areas with three active (gold and chrome), two closed (gold) and one planned mine (phosphate), we examined how mineral deposits and mining influence water chemistry and diatom and macroinvertebrate communities in subarctic streams in Finnish Lapland. We supplemented the data by additional samples compiled from databases and further assessed how variation in background geological conditions influences bioassessments of the impacts arising from mining. We found that water specific conductivity was elevated in our study streams draining through catchments with a high mineral potential. Mining effects were mainly seen as increased concentration of nitrogen. Influence of mineral deposits was detected in composition of diatom and macroinvertebrate communities, but communities in streams in areas with a high mineral potential were as diverse as those in streams in areas with a low mineral potential. Mining impacts were better detected for diatoms using a reference condition based on sites with a high than low mineral potential, while for macroinvertebrates, the responses were generally less evident, likely because of only minor effects of mining on water chemistry. Community composition and frequencies of occurrence of macroinvertebrate taxa were, however, highly similar between mine-influenced streams and reference streams with a high potential for minerals indicating that the communities are strongly structured by the natural influence of mineral deposits. Incorporating geochemistry into the reference condition would likely improve bioassessments of both taxonomic groups. Replicated monitoring in potentially impacted sites and reference sites would be the most efficient framework for detecting environmental impacts in streams draining through mineral-rich catchments.

Climate mediates continental scale patterns of stream microbial functional diversity.

BACKGROUND: Understanding the large-scale patterns of microbial functional diversity is essential for anticipating climate change impacts on ecosystems worldwide. However, studies of functional biogeography remain scarce for microorganisms, especially in freshwater ecosystems. Here we study 15,289 functional genes of stream biofilm microbes along three elevational gradients in Norway, Spain and China. RESULTS: We find that alpha diversity declines towards high elevations and assemblage composition shows increasing turnover with greater elevational distances. These elevational patterns are highly consistent across mountains, kingdoms and functional categories and exhibit the strongest trends in China due to its largest environmental gradients. Across mountains, functional gene assemblages differ in alpha diversity and composition between the mountains in Europe and Asia. Climate, such as mean temperature of the warmest quarter or mean precipitation of the coldest quarter, is the best predictor of alpha diversity and assemblage composition at both mountain and continental scales, with local non-climatic predictors gaining more importance at mountain scale. Under future climate, we project substantial variations in alpha diversity and assemblage composition across the Eurasian river network, primarily occurring in northern and central regions, respectively. CONCLUSIONS: We conclude that climate controls microbial functional gene diversity in streams at large spatial scales; therefore, the underlying ecosystem processes are highly sensitive to climate variations, especially at high latitudes. This biogeographical framework for microbial functional diversity serves as a baseline to anticipate ecosystem responses and biogeochemical feedback to ongoing climate change. Video Abstract.

Relationships between nutrient, chlorophyll a and Secchi depth in lakes of the Chinese Eastern Plains ecoregion: Implications for eutrophication management.

Eutrophication and associated algal blooms are principal environmental challenges confronting lakes in China, particularly in the Eastern Plains ecoregion. The empirical relationships between nutrient and chlorophyll a (Chla) level and Secchi depth (SD) are widely used as a theoretical basis for lake eutrophication management. However, these relationships are largely influenced by hydromorphological conditions and biogeochemical processes. Thus, there is a need to establish a type-specific understanding of these interactions. In the current study, lakes in the Chinese Eastern Plains ecoregion were subdivided into four lake types according to water retention time (LRT), water depth, and water area. Regression analyses indicated that the impacts of nutrient (total nitrogen, TN; total phosphorus, TP) concentrations on summer Chla were significantly reduced in lakes with high inorganic suspended solids (ISS) (P<0.05). Meanwhile, the decrease in SD in these lakes were found to relate mainly to non-algal turbidity. In lakes characterized by both short LRT and high ISS content, the Chla exhibited limited response to nutrients. In contrast, in lakes with low ISS content and long LRT, the observed slopes of both Chla=f(TP) and SD=f(Chla) were significantly steeper (P < 0.05). The factors limiting summer algal growth and the development of type-specific nutrient criteria (TN and TP) of all four investigated lake types in the Eastern Plains ecoregion are discussed in the context of specific nutrients. Based on these results, we establish type-specific eutrophication assessment equations of TN, TP, Chla, and SD in our study lakes. Our results may provide essential information for achieving the cost-effective eutrophication management of lakes both in the Eastern Plains ecoregion and elsewhere with similar climatic and hydromorphological conditions. Moreover, we believe that the subdivision of lakes to allow type-specific eutrophication management framework may prove valuable for other ecoregions where the interpretation of empirical nutrient-Chla and SD relationships suffer from similar serious limitations.

Ecological processes underlying community assembly of aquatic bacteria and macroinvertebrates under contrasting climates on the Tibetan Plateau.

Understanding the role of climatic variation on biodiversity is of chief importance due to the ongoing biodiversity loss and climate change. Freshwaters, one of the most threatened ecosystems in the world, offer a valuable context to study biodiversity patterns of distinct organism groups in relation to climatic variation. In the Tibetan Plateau biodiversity hotspot - Hengduan Mountain region, we studied the effects of climate and local physico-chemical factors on stream microorganisms (i.e. bacteria) and macroorganisms (i.e. macroinvertebrates) in two parallel catchments with contrasting precipitation and temperature, that is, the Nujiang and Lancang Rivers. Diversities and community structures were better explained by climatic and local environmental variables in the drier and colder catchment and at higher elevations, than in the warmer and wetter conditions and at lower elevations. This suggests that communities may be more strongly assembled by deterministic processes in the former, comparatively harsher conditions, compared to the latter, more benign conditions. Macroinvertebrates were more strongly affected by climatic and local environmental factors compared to bacteria, but the diversities and community structures of the two groups showed spatially similar responses to overall abiotic variation, being especially evident with their community structures' responses to climate. Furthermore, bacterial and macroinvertebrate diversities were positively correlated in the drier and colder catchment, implying that these biologically and ecologically distinct organism groups are likely to be driven by similar processes in areas with such climatic conditions. We conclude that changes in climatic and local environmental conditions may affect the diversity of macroorganisms more strongly than that of microorganisms, at least in subtropical mountainous stream ecosystems studied here, but simultaneous responses of both groups to environmental changes can also be expected.

Relative roles of spatial processes, natural factors and anthropogenic stressors in structuring a lake macroinvertebrate metacommunity.

Studies of aquatic metacommunities have so far been focused almost entirely on relatively isolated systems, such as a set of streams, lakes or ponds. Here, we aimed to quantify the relative importance of spatial processes, natural factors and anthropogenic stressors in structuring of a macroinvertebrate metacommunity within a large, highly-connected shallow lake system. The roles of different drivers were evaluated for the entire metacommunity, 10 trait-based deconstructed metacommunities and four common species by incorporating extensive sampling and a large number of abiotic explanatory variables. Contrary to our expectations, we found that variation in community structure among sites was mostly correlated to spatial and wind-wave variables rather than anthropogenic disturbance factors even though the lake presented strong environmental gradients associated with long-term human pressures. In addition, the relative importance of the three groups of drivers varied slightly among the deconstructed trait matrices (i.e. based on dispersal ability, feeding mode and degree of occurrence). Importantly, the distributions of the most common species showed significant and strong spatial autocorrelation, indicating the prominent role of high dispersal rate for their distributions. These findings suggest that the influences of high dispersal rates and natural disturbance may even override the roles of anthropogenic stressors in metacommunity organization in highly-connected aquatic systems. Hence, we strongly encourage that spatial processes and natural drivers are taken into account in the development of bioassessment approaches in highly-connected aquatic systems.

Bacterial metacommunity organization in a highly connected aquatic system.

The spatial structure and underlying assembly mechanisms of bacterial communities have been studied widely across aquatic systems, focusing primarily on isolated sites, such as different lakes, ponds and streams. Here, our main aim was to determine the underlying mechanisms for bacterial biofilm assembly within a large, highly connected lake system in Northern Finland using associative methods based on taxonomic and phylogenetic alpha- and beta-diversity and a large number of abiotic and biotic variables. Furthermore, null model approaches were used to quantify the relative importance of different community assembly processes. We found that spatial variation in bacterial communities within the lake was structured by different assembly processes, including stochasticity, species sorting and potentially even dispersal limitation. Species sorting by abiotic environmental conditions explained more of the taxonomic and particularly phylogenetic turnover in community composition compared with that by biotic variables. Finally, we observed clear differences in alpha diversity (species richness and phylogenetic diversity), which were to a stronger extent determined by abiotic compared with biotic factors, but also by dispersal effects. In summary, our study shows that the biodiversity of bacterial biofilm communities within a lake ecosystem is driven by within-habitat gradients in abiotic conditions and by stochastic and deterministic dispersal processes.

Freshwater diatoms as environmental indicators: evaluating the effects of eutrophication using species morphology and biological indices.

Anthropogenic eutrophication is a major form of perturbation in freshwaters, and several approaches aim to recognise its effects on lake ecosystems. We compared the responses of diatom species morphology, diversity indices and diatom indices to total phosphorus, total nitrogen and distance from a point stressor causing eutrophication in a large lake. We specifically examined the degree to which extent nutrients and distance to the stressor affect variation in the values of various biological indices and diatom valve size. In addition, special attention was given to the adequate repetition of diatom valve width measurements in the context of environmental assessment. Our results showed that diatom valve width was a better indicator of nutrient concentrations than any of the diatom and diversity indices examined. However, the results varied between the two study transects, suggesting that the diatom-based variables not only respond to nutrients but also to other environmental factors (e.g. shoreline morphology). We also found that when using the method based on diatom morphology, one should measure more valves than has been originally suggested to provide a more reliable picture of response to eutrophication. We argue that diatom morphology could be considered as an additional environmental assessment tool, because it may complement the information provided by the traditional diatom indices. Diatom valve width may also be more sensitive to early phases of the eutrophication process and its effects on freshwater ecosystems than various diatom indices that were developed in regional contexts with wide ranges in nutrient levels.