Bacterial rarity in a subarctic stream network: Biodiversity patterns, assembly mechanisms and types of rarity.

Dendritic stream networks are an intriguing subject for exploring the spatial and temporal variability of the rare and common bacterial biosphere, yet very few such studies have been conducted. We sampled riverine bacterioplankton at 13 sites in a subarctic riverine network across 3 years, with five sampling times each year. Ordinations showed a consistent pattern of downstream shift for both rare and abundant subcommunities. We also detected a temporal signal, with seasonal community shifts reflecting changes in water temperature and groundwater contribution, and an inter-annual pattern where the year 2018 differed from other years. Phylogenetic turnover of the rare subcommunity indicated homogeneous selection, whereas the abundant subcommunity was mainly stochastically structured. Transiently rare taxa were the dominant type of rarity with the highest proportion at the headwater regions. The bacterioplankton community was characterized by a small group of core taxa that occurred at most sites with little temporal variation, a very large number of permanently or transiently rare taxa, and taxa shifting through time between the rare and abundant biosphere. While this basic structure could have been detected with less extensive temporal replication, a comprehensive understanding of the rare biosphere in riverine bacterioplankton can only be achieved via inter-annual, spatially replicated sampling that covers the whole stream network.

Multi-decadal improvements in the ecological quality of European rivers are not consistently reflected in biodiversity metrics.

Humans impact terrestrial, marine and freshwater ecosystems, yet many broad-scale studies have found no systematic, negative biodiversity changes (for example, decreasing abundance or taxon richness). Here we show that mixed biodiversity responses may arise because community metrics show variable responses to anthropogenic impacts across broad spatial scales. We first quantified temporal trends in anthropogenic impacts for 1,365 riverine invertebrate communities from 23 European countries, based on similarity to least-impacted reference communities. Reference comparisons provide necessary, but often missing, baselines for evaluating whether communities are negatively impacted or have improved (less or more similar, respectively). We then determined whether changing impacts were consistently reflected in metrics of community abundance, taxon richness, evenness and composition. Invertebrate communities improved, that is, became more similar to reference conditions, from 1992 until the 2010s, after which improvements plateaued. Improvements were generally reflected by higher taxon richness, providing evidence that certain community metrics can broadly indicate anthropogenic impacts. However, richness responses were highly variable among sites, and we found no consistent responses in community abundance, evenness or composition. These findings suggest that, without sufficient data and careful metric selection, many common community metrics cannot reliably reflect anthropogenic impacts, helping explain the prevalence of mixed biodiversity trends.

Understanding temporal variability across trophic levels and spatial scales in freshwater ecosystems.

A tenet of ecology is that temporal variability in ecological structure and processes tends to decrease with increasing spatial scales (from locales to regions) and levels of biological organization (from populations to communities). However, patterns in temporal variability across trophic levels and the mechanisms that produce them remain poorly understood. Here we analyzed the abundance time series of spatially structured communities (i.e., metacommunities) spanning basal resources to top predators from 355 freshwater sites across three continents. Specifically, we used a hierarchical partitioning method to disentangle the propagation of temporal variability in abundance across spatial scales and trophic levels. We then used structural equation modeling to determine if the strength and direction of relationships between temporal variability, synchrony, biodiversity, and environmental and spatial settings depended on trophic level and spatial scale. We found that temporal variability in abundance decreased from producers to tertiary consumers but did so mainly at the local scale. Species population synchrony within sites increased with trophic level, whereas synchrony among communities decreased. At the local scale, temporal variability in precipitation and species diversity were associated with population variability (linear partial coefficient, ß = 0.23) and population synchrony (ß = -0.39) similarly across trophic levels, respectively. At the regional scale, community synchrony was not related to climatic or spatial predictors, but the strength of relationships between metacommunity variability and community synchrony decreased systematically from top predators (ß = 0.73) to secondary consumers (ß = 0.54), to primary consumers (ß = 0.30) to producers (ß = 0). Our results suggest that mobile predators may often stabilize metacommunities by buffering variability that originates at the base of food webs. This finding illustrates that the trophic structure of metacommunities, which integrates variation in organismal body size and its correlates, should be considered when investigating ecological stability in natural systems. More broadly, our work advances the notion that temporal stability is an emergent property of ecosystems that may be threatened in complex ways by biodiversity loss and habitat fragmentation.

Bacterial communities in a subarctic stream network: Spatial and seasonal patterns of benthic biofilm and bacterioplankton.

Water-column bacterial communities are assembled by different mechanisms at different stream network positions, with headwater communities being controlled by mass effects (advection of bacteria from terrestrial soils) while downstream communities are mainly driven by environmental sorting. Conversely, benthic biofilms are colonized largely by the same set of taxa across the entire network. However, direct comparisons of biofilm and bacterioplankton communities along whole stream networks are rare. We used 16S rRNA gene amplicon sequencing to explore the spatiotemporal variability of benthic biofilm (2 weeks old vs. mature biofilm) and water-column communities at different network positions of a subarctic stream from early summer to late autumn. Amplicon sequence variant (ASV) richness of mature biofilm was about 2.5 times higher than that of early biofilm, yet the pattern of seasonality was the same, with the highest richness in midsummer. Biofilm bacterial richness was unrelated to network position whereas bacterioplankton diversity was negatively related to water residence time and distance from the source. This pattern of decreasing diversity along the network was strongest around midsummer and diminished greatly as water level increased towards autumn. Biofilm communities were phylogenetically clustered at all network positions while bacterioplankton assemblages were phylogenetically clustered only at the most downstream site. Both early and mature biofilm communities already differed significantly between upstream (1st order) and midstream (2nd order) sections. Network position was also related to variation in bacterioplankton communities, with upstream sites harbouring substantially more unique taxa (44% of all upstream taxa) than midstream (20%) or downstream (8%) sites. Some of the taxa that were dominant in downstream sections were already present in the upmost headwaters, and even in riparian soils, where they were very rare (relative abundance <0.01%). These patterns in species diversity and taxonomic and phylogenetic community composition of the riverine bacterial metacommunity were particularly strong for water-column communities, whereas both early and mature biofilm exhibited weaker spatial patterns. Our study demonstrated the benefits of studying bacterioplankton and biofilm communities simultaneously to allow testing of ecological hypotheses about biodiversity patterns in freshwater bacteria.

Evaluating the biological validity of European river typology systems with least disturbed benthic macroinvertebrate communities.

Humans have severely altered freshwater ecosystems globally, causing a loss of biodiversity. Regulatory frameworks, like the Water Framework Directive, have been developed to support actions that halt and reverse this loss. These frameworks use typology systems that summarize freshwater ecosystems into environmentally delineated types. Within types, ecosystems that are minimally impacted by human activities, i.e., in reference conditions, are expected to be similar concerning physical, chemical, and biological characteristics. This assumption is critical when water quality assessments rely on comparisons to type-specific reference conditions. Lyche Solheim et al. (2019) developed a pan-European river typology system, the Broad River Types, that unifies the national Water Framework Directive typology systems and is gaining traction within the research community. However, it is unknown how similar biological communities are within these individual Broad River Types. We used analysis of similarities and classification strength analysis to examine if the Broad River Types delineate distinct macroinvertebrate communities across Europe and whether they outperform two ecoregional approaches: the European Biogeographical Regions and Illies' Freshwater Ecoregions. We determined indicator and typical taxa for the types of all three typology systems and evaluated their distinctiveness. All three typology systems captured more variation in macroinvertebrate communities than random combinations of sites. The results were similar among typology systems, but the Broad River Types always performed worse than either the Biogeographic Regions or Illies' Freshwater Ecoregions. Despite reaching statistical significance, the statistics of analysis of similarity and classification strength were low in all tests indicating substantial overlap among the macroinvertebrate communities of different types. We conclude that the Broad River Types do not represent an improvement upon existing freshwater typologies when used to delineate macroinvertebrate communities and we propose future avenues for advancement: regionally constrained types, better recognition of intermittent rivers, and consideration of biotic communities.

Dark matters: Contrasting responses of stream biofilm to browning and loss of riparian shading.

Concentrations of terrestrial-derived dissolved organic carbon (DOC) in freshwater ecosystems have increased consistently, causing freshwater browning. The mechanisms behind browning are complex, but in forestry-intensive regions browning is accelerated by land drainage. Forestry actions in streamside riparian forests alter canopy shading, which together with browning is expected to exert a complex and largely unpredictable control over key ecosystem functions. We conducted a stream mesocosm experiment with three levels of browning (ambient vs. moderate vs. high, with 2.7 and 5.5-fold increase, respectively, in absorbance) crossed with two levels of riparian shading (70% light reduction vs. open canopy) to explore the individual and combined effects of browning and loss of shading on the quantity (algal biomass) and nutritional quality (polyunsaturated fatty acid and sterol content) of the periphytic biofilm. We also conducted a field survey of differently colored (4.7 to 26.2 mg DOC L-1 ) streams to provide a 'reality check' for our experimental findings. Browning reduced greatly the algal biomass, suppressed the availability of essential polyunsaturated fatty acids, especially eicosapentaenoic acid (EPA), and sterols, but increased the availability of terrestrial-derived long-chain saturated fatty acids (LSAFA). In contrast, loss of shading increased primary productivity, which resulted in elevated sterol and EPA contents of the biofilm. The field survey largely repeated the same pattern: biofilm nutritional quality decreased significantly with increasing DOC, as indicated particularly by a decrease of the ω-3:ω-6 ratio and increase in LSAFA content. Algal biomass, in contrast, was mainly controlled by dissolved inorganic nitrogen (DIN) concentration, while DOC concentration was of minor importance. The ongoing browning process is inducing a dramatic reduction in the nutritional quality of the stream biofilm. Such degradation of the major high-quality food source available for stream consumers may reduce the trophic transfer efficiency in stream ecosystems, potentially extending across the stream-forest ecotone.

Bacterial communities at a groundwater-surface water ecotone: gradual change or abrupt transition points along a contamination gradient?

Microbial communities contribute greatly to groundwater quality, but the impacts of land-use practices on bacteria in groundwaters and groundwater-dependent ecosystems remain poorly known. With 16S rRNA gene amplicon sequencing, we assessed bacterial community composition at the groundwater-surface water ecotone of boreal springs impacted by urbanization and agriculture, using spring water nitrate-N as a surrogate of contamination. We also measured the rate of a key ecosystem process, organic matter decomposition. We documented a recurrent pattern across all major bacterial phyla where diversity started to decrease at unexpectedly low nitrate-N concentrations (100-300 µg L-1 ). At 400 NO3 - -N µg L-1 , 25 bacterial exact sequence variants showed a negative response, resulting in a distinct threshold in bacterial community composition. Chthonomonas, Acetobacterales and Hyphomicrobium were the most sensitive taxa, while only three taxa (Duganella, Undibacterium and Thermoanaerobaculaceae) were enriched due to increased contamination. Decomposition rate responded unimodally to increasing nitrate-N concentration, with a peak rate at ~400 NO3 - -N µg L-1 , parallelly with a major shift in bacterial community composition. Our results emphasize the utility of bacterial communities in the assessment of groundwater-dependent ecosystems. They also call for a careful reconsideration of threshold nitrate values for defining groundwater ecosystem health and protecting their microbial biodiversity.

Stochastic processes and ecological connectivity drive stream invertebrate community responses to short-term drought.

Community responses to and recovery from disturbances depend on local (e.g. presence of refuges) and regional (connectivity to recolonization sources) factors. Droughts are becoming more frequent in boreal regions, and are likely to constitute a severe disturbance for boreal stream communities where organisms largely lack adaptations to such hydrological extremes. We conducted an experiment in 24 semi-natural stream flumes to assess the effects of local and regional factors on the responses of benthic invertebrate communities to a short-term drought. We manipulated flow (drought vs. constant-flow), spatial arrangement of leaf litter patches (aggregated vs. evenly distributed) and colonization from regional species pool (enhanced vs. ambient connectivity) to test the combined effects of disturbance, resource arrangement and connectivity on the structural and functional responses of benthic invertebrate communities. We found that a drought as short as 1 week reduced invertebrate taxonomic richness and abundance, mainly through stochastic extinctions. Such changes in richness were not reflected in functional diversity. This suggests that communities were characterized by a high degree of functional redundancy, which allowed maintenance of functional diversity despite species losses. Feeding groups responded differently to drought, with organic matter decomposers responding more than scrapers and predators. Three weeks were insufficient for complete invertebrate community recovery from drought. However, recovery was greater in channels subjected to enhanced connectivity, which increased taxonomic diversity and abundance of certain taxa. Spatial configuration of resources explained the least variation in our response variables, having a significant effect only on invertebrate abundance and evenness (both sampling occasions) and taxonomic richness (end of recovery period). Even a short drought, if occurring late in the season, may not allow communities to recover before the onset of winter, thus having a potentially long-lasting effect on stream communities. For boreal headwaters, extreme dewatering poses a novel disturbance regime that may trigger substantial and potentially irreversible changes. An improved understanding of such changes is needed to underpin adaptive management strategies in these increasingly fragmented and disturbed ecosystems.

Multiple stressors in small streams in the forestry context of Fennoscandia: The effects in time and space.

In this paper we describe how forest management practices in Fennoscandian countries, namely Sweden and Finland, expose streams to multiple stressors over space and time. In this region, forestry includes several different management actions and we explore how these may successively disturb the same location over 60-100 year long rotation periods. Of these actions, final harvest and associated road construction, soil scarification, and/or ditch network maintenance are the most obvious sources of stressors to aquatic ecosystems. Yet, more subtle actions such as planting, thinning of competing saplings and trees, and removing logging residues also represent disturbances around waterways in these landscapes. We review literature about how these different forestry practices may introduce a combination of physicochemical stressors, including hydrological change, increased sediment transport, altered thermal and light regimes, and water quality deterioration. We further elaborate on how the single stressors may combine and interact and we consequently hypothesise how these interactions may affect aquatic communities and processes. Because production forestry is practiced on a large area in both countries, the various stressors appear multiple times during the rotation cycles and potentially affect the majority of the stream network length within most catchments. We concluded that forestry practices have traditionally not been the focus of multiple stressor studies and should be investigated further in both observational and experimental fashion. Stressors accumulate across time and space in forestry dominated landscapes, and may interact in unpredictable ways, limiting our current understanding of what forested stream networks are exposed to and how we can design and apply best management practices.

Habitat patchiness, ecological connectivity and the uneven recovery of boreal stream ecosystems from an experimental drought.

Ongoing climate change is increasing the occurrence and intensity of drought episodes worldwide, including in boreal regions not previously regarded as drought prone, and where the impacts of drought remain poorly understood. Ecological connectivity is one factor that might influence community structure and ecosystem functioning post-drought, by facilitating the recovery of sensitive species via dispersal at both local (e.g. a nearby habitat patch) and regional (from other systems within the same region) scales. In an outdoor mesocosm experiment, we investigated how impacts of drought on boreal stream ecosystems are altered by the spatial arrangement of local habitat patches within stream channels, and variation in ecological connectivity with a regional species pool. We measured basal ecosystem processes underlying carbon and nutrient cycling: (a) algal biomass accrual; (b) microbial respiration; and (c) decomposition of organic matter, and sampled communities of aquatic fungi and benthic invertebrates. An 8-day drought event had strong impacts on both community structure and ecosystem functioning, including algal accrual, leaf decomposition and microbial respiration, with many of these impacts persisting even after water levels had been restored for 3.5 weeks. Enhanced connectivity with the regional species pool and increased aggregation of habitat patches also affected multiple response variables, especially those associated with microbes, and in some cases reduced the effects of drought to a small extent. This indicates that spatial processes might play a role in the resilience of communities and ecosystem functioning, given enough time. These effects were however insufficient to facilitate significant recovery in algal growth before seasonal dieback began in autumn. The limited resilience of ecosystem functioning in our experiment suggests that even short-term droughts can have extended consequences for stream ecosystems in the world's vast boreal region, and especially on the ecosystem processes and services mediated by algal biofilms.

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.

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.

Combined effects of local habitat, anthropogenic stress, and dispersal on stream ecosystems: a mesocosm experiment.

The effects of anthropogenic stressors on community structure and ecosystem functioning can be strongly influenced by local habitat structure and dispersal from source communities. Catchment land uses increase the input of fine sediments into stream channels, clogging the interstitial spaces of benthic habitats. Aquatic macrophytes enhance habitat heterogeneity and mediate important ecosystem functions, being thus a key component of habitat structure in many streams. Therefore, the recovery of macrophytes following in-stream habitat modification may be prerequisite for successful stream restoration. Restoration success is also affected by dispersal of organisms from the source community, with potentially the strongest responses in relatively isolated headwater sites that receive a limited amount of dispersing individuals. We used a factorial design in a set of stream mesocosms to study the independent and combined effects of an anthropogenic stressor (sand sedimentation), local habitat (macrophytes, i.e., moss transplants), and enhanced dispersal (two levels: high vs. low) on organic matter retention, algal accrual rate, leaf decomposition, and macroinvertebrate community structure. Overall, all responses were simple additive effects with no interactions between treatments. Sand reduced algal accumulation, total invertebrate density, and density of a few individual taxa. Mosses reduced algal accrual rate and algae-grazing invertebrates, but enhanced organic matter retention and the number of detritus and filter feeders. Mosses also reduced macroinvertebrate diversity by increasing the dominance by a few taxa. Mosses reduced leaf mass loss, possibly because the organic matter retained by mosses provided an additional food source for leaf-shredding invertebrates and thus reduced shredder aggregation into leaf packs. The effect of mosses on macroinvertebrate communities and ecosystem functioning was distinct irrespective of the level of dispersal, suggesting strong environmental control of community structure. The strong environmental control of macroinvertebrate community composition even under enhanced dispersal suggests that re-establishing key habitat features, such as natural stream vegetation, could aid ecosystem recovery in boreal streams.

Do latitudinal gradients exist in New Zealand stream invertebrate metacommunities?

That biodiversity declines with latitude is well known, but whether a metacommunity process is behind this gradient has received limited attention. We tested the hypothesis that dispersal limitation is progressively replaced by mass effects with increasing latitude, along with a series of related hypotheses. We explored these hypotheses by examining metacommunity structure in stream invertebrate metacommunities spanning the length of New Zealand's two largest islands (∼1,300 km), further disentangling the role of dispersal by deconstructing assemblages into strong and weak dispersers. Given the highly dynamic nature of New Zealand streams, our alternative hypothesis was that these systems are so unpredictable (at different stages of post-flood succession) that metacommunity structure is highly context dependent from region to region. We rejected our primary hypotheses, pinning this lack of fit on the strong unpredictability of New Zealand's dynamic stream ecosystems and fauna that has evolved to cope with these conditions. While local community structure turned over along this latitudinal gradient, metacommunity structure was highly context dependent and dispersal traits did not elucidate patterns. Moreover, the emergent metacommunity types exhibited no trends, nor did the important environmental variables. These results provide a cautionary tale for examining singular metacommunities. The considerable level of unexplained contingency suggests that any inferences drawn from one-off snapshot sampling may be misleading and further points to the need for more studies on temporal dynamics of metacommunity processes.

Groundwater contamination and land drainage induce divergent responses in boreal spring ecosystems.

Degradation of freshwater ecosystems has engendered legislative mandates for the protection and management of surface waters while groundwater-dependent ecosystems (GDEs) have received much less attention. This is so despite biodiversity and functioning of GDEs are currently threatened by several anthropogenic stressors, particularly intensified land use and groundwater contamination. We assessed the impacts of land drainage (increased input of dissolved organic carbon, DOC, from peatland drainage) and impaired groundwater chemical quality (NO3--N enrichment from agricultural or urban land use) on biodiversity and ecosystem functioning in 20 southern Finnish cold-water springs using several taxonomic and functional measures. Groundwater contamination decreased macroinvertebrate and bacterial diversity and altered their community composition. Changes in macroinvertebrate and bacterial communities along the gradient of water-quality impairment were caused by the replacement of native with new taxa rather than by mere disappearance of some of the original taxa. Also species richness of habitat specialist (but not headwater generalist) bryophytes decreased due to impaired groundwater quality. Periphyton accrual rate showed a subsidy-stress response to elevated nitrate concentrations, with peak values at around 2500 µg L-1, while drainage-induced spring water brownification (increased DOC) reduced both periphyton accrual and leaf decomposition rates already at very low concentrations. Our results highlight the underutilized potential of ecosystem-level functional measures in GDE bioassessment as they seem to respond to the first signs of spring ecosystem impairment, at least for the anthropogenic stressors studied by us.

Identifying taxonomic and functional surrogates for spring biodiversity conservation.

Surrogate approaches are widely used to estimate overall taxonomic diversity for conservation planning. Surrogate taxa are frequently selected based on rarity or charisma, whereas selection through statistical modeling has been applied rarely. We used boosted-regression-tree models (BRT) fitted to biological data from 165 springs to identify bryophyte and invertebrate surrogates for taxonomic and functional diversity of boreal springs. We focused on these 2 groups because they are well known and abundant in most boreal springs. The best indicators of taxonomic versus functional diversity differed. The bryophyte Bryum weigelii and the chironomid larva Paratrichocladius skirwithensis best indicated taxonomic diversity, whereas the isopod Asellus aquaticus and the chironomid Macropelopia spp. were the best surrogates of functional diversity. In a scoring algorithm for priority-site selection, taxonomic surrogates performed only slightly better than random selection for all spring-dwelling taxa, but they were very effective in representing spring specialists, providing a distinct improvement over random solutions. However, the surrogates for taxonomic diversity represented functional diversity poorly and vice versa. When combined with cross-taxon complementarity analyses, surrogate selection based on statistical modeling provides a promising approach for identifying groundwater-dependent ecosystems of special conservation value, a key requirement of the EU Water Framework Directive.

Thermal and hydrologic responses to climate change predict marked alterations in boreal stream invertebrate assemblages.

Air temperature at the northernmost latitudes is predicted to increase steeply and precipitation to become more variable by the end of the 21st century, resulting in altered thermal and hydrological regimes. We applied five climate scenarios to predict the future (2070-2100) benthic macroinvertebrate assemblages at 239 near-pristine sites across Finland (ca. 1200 km latitudinal span). We used a multitaxon distribution model with air temperature and modeled daily flow as predictors. As expected, projected air temperature increased the most in northernmost Finland. Predicted taxonomic richness also increased the most in northern Finland, congruent with the predicted northwards shift of many species' distributions. Compositional changes were predicted to be high even without changes in richness, suggesting that species replacement may be the main mechanism causing climate-induced changes in macroinvertebrate assemblages. Northern streams were predicted to lose much of the seasonality of their flow regimes, causing potentially marked changes in stream benthic assemblages. Sites with the highest loss of seasonality were predicted to support future assemblages that deviate most in compositional similarity from the present-day assemblages. Macroinvertebrate assemblages were also predicted to change more in headwaters than in larger streams, as headwaters were particularly sensitive to changes in flow patterns. Our results emphasize the importance of focusing protection and mitigation on headwater streams with high-flow seasonality because of their vulnerability to climate change.

Disentangling the effects of low pH and metal mixture toxicity on macroinvertebrate diversity.

One of the primary goals of biological assessment of streams is to identify which of a suite of chemical stressors is limiting their ecological potential. Elevated metal concentrations in streams are often associated with low pH, yet the effects of these two potentially limiting factors of freshwater biodiversity are rarely considered to interact beyond the effects of pH on metal speciation. Using a dataset from two continents, a biogeochemical model of the toxicity of metal mixtures (Al, Cd, Cu, Pb, Zn) and quantile regression, we addressed the relative importance of both pH and metals as limiting factors for macroinvertebrate communities. Current environmental quality standards for metals proved to be protective of stream macroinvertebrate communities and were used as a starting point to assess metal mixture toxicity. A model of metal mixture toxicity accounting for metal interactions was a better predictor of macroinvertebrate responses than a model considering individual metal toxicity. We showed that the direct limiting effect of pH on richness was of the same magnitude as that of chronic metal toxicity, independent of its influence on the availability and toxicity of metals. By accounting for the direct effect of pH on macroinvertebrate communities, we were able to determine that acidic streams supported less diverse communities than neutral streams even when metals were below no-effect thresholds. Through a multivariate quantile model, we untangled the limiting effect of both pH and metals and predicted the maximum diversity that could be expected at other sites as a function of these variables. This model can be used to identify which of the two stressors is more limiting to the ecological potential of running waters.

Effects of changing climate on European stream invertebrate communities: A long-term data analysis.

Long-term observations on riverine benthic invertebrate communities enable assessments of the potential impacts of global change on stream ecosystems. Besides increasing average temperatures, many studies predict greater temperature extremes and intense precipitation events as a consequence of climate change. In this study we examined long-term observation data (10-32years) of 26 streams and rivers from four ecoregions in the European Long-Term Ecological Research (LTER) network, to investigate invertebrate community responses to changing climatic conditions. We used functional trait and multi-taxonomic analyses and combined examinations of general long-term changes in communities with detailed analyses of the impact of different climatic drivers (i.e., various temperature and precipitation variables) by focusing on the response of communities to climatic conditions of the previous year. Taxa and ecoregions differed substantially in their response to climate change conditions. We did not observe any trend of changes in total taxonomic richness or overall abundance over time or with increasing temperatures, which reflects a compensatory turnover in the composition of communities; sensitive Plecoptera decreased in response to warmer years and Ephemeroptera increased in northern regions. Invasive species increased with an increasing number of extreme days which also caused an apparent upstream community movement. The observed changes in functional feeding group diversity indicate that climate change may be associated with changes in trophic interactions within aquatic food webs. These findings highlight the vulnerability of riverine ecosystems to climate change and emphasize the need to further explore the interactive effects of climate change variables with other local stressors to develop appropriate conservation measures.

Sediments and flow have mainly independent effects on multitrophic stream communities and ecosystem functions.

Stream ecosystems are affected by multiple abiotic stressors, and species responses to simultaneous stressors may differ from those predicted based on single-stressor responses. Using 12 semi-natural stream channels, we examined the individual and interactive effects of flow level (low or high flow) and addition of fine sediments (grain size <2 mm) on key ecosystem processes (leaf breakdown, algal biomass accrual) and benthic macroinvertebrate and fungal communities. Both stressors had mostly independent effects on biological responses, with sand addition being the more influential of the two. Sand addition decreased algal biomass and microbe-mediated leaf breakdown significantly, whereas invertebrate shredder-mediated breakdown only responded to flow level. Macroinvertebrate community composition responded significantly to both stressors. Fungal biomass decreased and shredder abundance increased when sand was added; thus, organisms at different trophic levels can exhibit highly variable responses to the same stressor. Terrestrial endophytic fungi were abundant in low-flow flumes where leaf mass loss was also highest, indicating that terrestrial endophytes may contribute importantly to leaf decomposition in the aquatic environment. Leaf breakdown rates depended on the identity and abundance of the dominant decomposer species, suggesting that the effects of anthropogenic activities on ecosystem processes may be driven by changes in the abundance of a few key species. The few observed interactive effects were all antagonistic (i.e., less than the sum of the individual effects); for example, increased flow stimulated algal biomass accumulation but this effect was largely cancelled by sand. While our finding that sand and stream flow did not have strong synergistic effects can be considered reassuring for management, future experiments should manipulate these and other human stressors in experiments that run for much longer periods, thus focusing on the long-term impacts of multiple simultaneously operating stressors.