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.

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.

Excess of nitrogen reduces temporal variability of stream diatom assemblages.

Nutrient enrichment degrades water quality and threatens aquatic biota. However, our knowledge on (dis)similarities in temporal patterns of biota among sites of varying level of nutrient stress is limited. We addressed this gap by assessing temporal (among seasons) variation in algal biomass, species diversity and composition of diatom assemblages in three streams that differ in nutrient stress, but are otherwise similar and share the same regional species pool. We monitored three riffle sections in each stream bi-weekly from May to October in 2014. Temporal variation in water chemistry and other environmental variables was mainly synchronous among riffles within streams and often also among streams, indicating shared environmental forcing through time. We found significant differences in diatom assemblage composition among streams and, albeit less so, also among riffles within streams. Diatom assemblages in the two nutrient-enriched streams were more similar to each other than to those in the nutrient-poor stream. Taxa richness did not differ consistently among the streams, and did not vary synchronously at any spatial scale. Temporal variation in diatom assemblage composition decreased with increasing DIN:TotP ratio, likely via a negative effect on sensitive taxa while maintaining favorable conditions for certain tolerant taxa, irrespective of season. This relationship weakened but remained significant even after controlling for stochastic effects, suggesting deterministic mechanisms between nutrient levels and diatom assemblage stability. After controlling for stochastic effects temporal variability was best explained by DIN suggesting that excess of nitrogen reduces temporal variability(intra-annual beta diversity) of diatom assemblages. The high temporal variation, and especially the lack of temporal synchrony at the within streams scale, suggests that single sampling at a single site may be insufficient to reliably assess and monitor a complete stream water body. Our results also showed that measures including species identity outperform traditional diversity metrics in detecting nutrient stress in streams.

Correlates of different facets and components of beta diversity in stream organisms.

Recently, community ecology has emphasized the multi-facetted aspects of biological diversity by linking species traits and the environment. Here, we explored environmental correlates of taxonomically-based and traits-based compositional distances using a comprehensive data set of diatom and macroinvertebrate communities. We also explored the responses of different beta diversity components (i.e., overall beta diversity, turnover, and nestedness) of beta diversity facets (i.e., taxonomically and traits-based beta diversity) to environmental distances. Partial Mantel tests were used to test the relationships between beta diversity and environmental distance (while controlling for spatial distances). Taxonomically-based beta diversity varied much more than traits-based beta diversity, indicating strong functional convergence. We found that taxonomically-based beta diversity was largely driven by the turnover component. However, the nestedness component contributed more to overall traits-based beta diversity than the turnover component. Taxonomically-based beta diversity was significantly correlated with environmental distances for both diatoms and macroinvertebrates. Thus, we found support for the role of environmental filtering as a driver of community dissimilarities of rather different biological groups. However, the strength of these relationships between beta diversity and environmental distances varied depending on the biological group, facet, component, and the way which the environmental variables were selected to calculate the explanatory (distance) matrix. Our results indicated that both taxonomically and traits-based approaches are still needed to better understand patterns and mechanisms affecting the organization of biological communities in streams. This is because different facets of biological communities may be driven by different mechanisms.

Local geology determines responses of stream producers and fungal decomposers to nutrient enrichment: A field experiment.

We examined how short-term (19 days) nutrient enrichment influences stream fungal and diatom communities, and rates of leaf decomposition and algal biomass accrual. We conducted a field experiment using slow-releasing nutrient pellets to increase nitrate (NO3-N) and phosphate (PO4-P) concentrations in a riffle section of six naturally acidic (naturally low pH due to catchment geology) and six circumneutral streams. Nutrient enrichment increased microbial decomposition rate on average by 14%, but the effect was significant only in naturally acidic streams. Nutrient enrichment also decreased richness and increased compositional variability of fungal communities in naturally acidic streams. Algal biomass increased in both stream types, but algal growth was overall very low. Diatom richness increased in response to nutrient addition by, but only in circumneutral streams. Our results suggest that primary producers and decomposers are differentially affected by nutrient enrichment and that their responses to excess nutrients are context dependent, with a potentially stronger response of detrital processes and fungal communities in naturally acidic streams than in less selective environments.

Elevated nutrient concentrations in headwaters affected by drained peatland.

Nutrient export from drained peatland has significant impacts on aquatic environments in Nordic catchments. Spatial information on variations in nutrient concentrations across different landscapes and land uses is needed to design measures for achieving the good ecological status of the EU Water Framework Directive. In this study we determined background concentrations in natural peatland-dominated streams and examined effects of peatland use practices on water quality in headwater catchments. We quantified sources for different elements by joint analysis of water chemistry, isotopes, and hydrology for 62 small catchments in North Ostrobothnia, Finland. Concentrations of nutrients and suspended solids were relatively high in catchments containing drained peatland. In particular, dissolved nitrogen and phosphorus concentrations were elevated during baseflow conditions when flow likely originated from deeper soil layers. Total concentrations of nutrients, suspended solids, and loss on ignition also showed elevated values, and changes in the ratio of dissolved and particulate forms, especially the C/N ratio, were observed. Past drainage had a stronger effect on organic matter and nutrients concentrations than local hydroclimate conditions. These results strongly indicate that current water protection methods are not sufficient to capture all seasonal variations in nutrient and suspended solid loads from drained peatland. Thus, more effort and actions are needed for water protection in such areas.

Predicting occupancy and abundance by niche position, niche breadth and body size in stream organisms.

The regional occupancy and local abundance of species are thought to be strongly correlated to their body size, niche breadth and niche position. The strength of the relationships among these variables can also differ between different organismal groups. Here, we analyzed data on stream diatoms and insects from a high-latitude drainage basin to investigate these relationships. To generate measures of niche position and niche breadth for each species, we used sets of local environmental and catchment variables separately, applying the outlying mean index analysis. Beta regression and negative binomial generalized linear models were run to predict regional occupancy and mean local abundance, respectively. We found a positive occupancy-abundance relationship in both diatoms and insects, and that niche-based variables were the main predictors of variation in regional occupancy and local abundance. This finding was mainly due to local environmental niche position, whereas the effects of niche breadth on regional occupancy and local abundance were less important. We also found a relationship between body size and local abundance or regional occupancy of diatoms. Our results thus add to current macroecological research by emphasizing the strong importance of niche position rather than niche breadth and body size for regional occupancy and local abundance in rarely studied organisms (e.g., diatoms and insects) and ecosystems (i.e., wilderness streams).

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.

Redundancy in the ecological assessment of lakes: Are phytoplankton, macrophytes and phytobenthos all necessary?

Although the Water Framework Directive specifies that macrophytes and phytobenthos should be used for the ecological assessment of lakes and rivers, practice varies widely throughout the EU. Most countries have separate methods for macrophytes and phytobenthos in rivers; however, the situation is very different for lakes. Here, 16 countries do not have dedicated phytobenthos methods, some include filamentous algae within macrophyte survey methods whilst others use diatoms as proxies for phytobenthos. The most widely-cited justification for not having a dedicated phytobenthos method is redundancy, i.e. that macrophyte and phytoplankton assessments alone are sufficient to detect nutrient impacts. Evidence from those European Union Member States that have dedicated phytobenthos methods supports this for high level overviews of lake condition and classification; however, there are a number of situations where phytobenthos may contribute valuable information for the management of lakes.

Disentangling the responses of boreal stream assemblages to low stressor levels of diffuse pollution and altered channel morphology.

Non-point diffuse pollution from land use and alteration of hydromorphology are among the most detrimental stressors to stream ecosystems. We explored the independent and interactive effects of morphological channel alteration (channelization for water transport of timber) and diffuse pollution on species richness and community structure of four organism groups in boreal streams: diatoms, macrophytes, macroinvertebrates, and fish. Furthermore, the effect of these stressors on stream condition was evaluated by Ecological Quality Ratios (EQR) from the national Water Framework Directive (WFD) assessment system. We grouped 91 study sites into four groups that were impacted by either diffuse pollution or hydromorphological alteration, by both stressors, or by neither one. Macroinvertebrate richness was reduced by diffuse pollution, whereas other biological groups were unaltered. Hydromorphological modification had no effect on taxon richness of any of the assemblages. Community structure of all groups was significantly affected by diffuse pollution but not by hydromorphology. Similarly, EQRs indicated negative response by diatoms, macroinvertebrates and fish to diffuse pollution, but not to hydromorphological alteration. Agricultural diffuse pollution thus affected species identities and abundances rather than taxonomic richness. Our results suggest that channelization of boreal streams for timber transport has not altered hydromorphological conditions sufficiently to have a strong impact on stream biota, whereas even moderate nutrient enrichment may be ecologically harmful. Controlling diffuse pollution and associated land use stressors should be prioritized over restoration of in-stream habitat structure to improve the ecological condition of boreal streams.

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.