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.

From meta-system theory to the sustainable management of rivers in the Anthropocene.

Regional-scale ecological processes, such as the spatial flows of material, energy, and organisms, are fundamental for maintaining biodiversity and ecosystem functioning in river networks. Yet these processes remain largely overlooked in most river management practices and underlying policies. Here, we propose adoption of a meta-system approach, where regional processes acting at different levels of ecological organization - populations, communities, and ecosystems - are integrated into conventional river conservation, restoration, and biomonitoring. We also describe a series of measurements and indicators that could be assimilated into the implementation of relevant biodiversity and environmental policies. Finally, we highlight the need for alternative management strategies that can guide practitioners toward applying recent advances in ecology to preserve and restore river ecosystems and the ecosystem services they provide, in the context of increasing alteration of river network connectivity worldwide.

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.

Gaps in current Baltic Sea environmental monitoring - Science versus management perspectives.

Legislations and commitments regulate Baltic Sea status assessments and monitoring. These assessments suffer from monitoring gaps that need prioritization. We used three sources of information; scientific articles, project reports and a stakeholder survey to identify gaps in relation to requirements set by the HELCOM's Baltic Sea Action Plan, the Marine Strategy Framework Directive and the Water Framework Directive. The most frequently mentioned gap was that key requirements are not sufficiently monitored in space and time. Biodiversity monitoring was the category containing most gaps. However, whereas more than half of the gaps in reports related to biodiversity, scientific articles pointed out many gaps in the monitoring of pollution and water quality. An important finding was that the three sources differed notably with respect to which gaps were mentioned most often. Thus, conclusions about gap prioritization for management should be drawn after carefully considering the different viewpoints of scientists and stakeholders.

Benthic macroinvertebrates in lake ecological assessment: A review of methods, intercalibration and practical recommendations.

Legislation in Europe has been adopted to determine and improve the ecological integrity of inland and coastal waters. Assessment is based on four biotic groups, including benthic macroinvertebrate communities. For lakes, benthic invertebrates have been recognized as one of the most difficult organism groups to use in ecological assessment, and hitherto their use in ecological assessment has been limited. In this study, we review and intercalibrate 13 benthic invertebrate-based tools across Europe. These assessment tools address different human impacts: acidification (3 methods), eutrophication (3 methods), morphological alterations (2 methods), and a combination of the last two (5 methods). For intercalibration, the methods were grouped into four intercalibration groups, according to the habitat sampled and putative pressure. Boundaries of the 'good ecological status' were compared and harmonized using direct or indirect comparison approaches. To enable indirect comparison of the methods, three common pressure indices and two common biological multimetric indices were developed for larger geographical areas. Additionally, we identified the best-performing methods based on their responsiveness to different human impacts. Based on these experiences, we provide practical recommendations for the development and harmonization of benthic invertebrate assessment methods in lakes and similar habitats.

A comparative analysis reveals weak relationships between ecological factors and beta diversity of stream insect metacommunities at two spatial levels.

The hypotheses that beta diversity should increase with decreasing latitude and increase with spatial extent of a region have rarely been tested based on a comparative analysis of multiple datasets, and no such study has focused on stream insects. We first assessed how well variability in beta diversity of stream insect metacommunities is predicted by insect group, latitude, spatial extent, altitudinal range, and dataset properties across multiple drainage basins throughout the world. Second, we assessed the relative roles of environmental and spatial factors in driving variation in assemblage composition within each drainage basin. Our analyses were based on a dataset of 95 stream insect metacommunities from 31 drainage basins distributed around the world. We used dissimilarity-based indices to quantify beta diversity for each metacommunity and, subsequently, regressed beta diversity on insect group, latitude, spatial extent, altitudinal range, and dataset properties (e.g., number of sites and percentage of presences). Within each metacommunity, we used a combination of spatial eigenfunction analyses and partial redundancy analysis to partition variation in assemblage structure into environmental, shared, spatial, and unexplained fractions. We found that dataset properties were more important predictors of beta diversity than ecological and geographical factors across multiple drainage basins. In the within-basin analyses, environmental and spatial variables were generally poor predictors of variation in assemblage composition. Our results revealed deviation from general biodiversity patterns because beta diversity did not show the expected decreasing trend with latitude. Our results also call for reconsideration of just how predictable stream assemblages are along ecological gradients, with implications for environmental assessment and conservation decisions. Our findings may also be applicable to other dynamic systems where predictability is low.

The influence of environmental, biotic and spatial factors on diatom metacommunity structure in Swedish headwater streams.

Stream assemblages are structured by a combination of local (environmental filtering and biotic interactions) and regional factors (e.g., dispersal related processes). The relative importance of environmental and spatial (i.e., regional) factors structuring stream assemblages has been frequently assessed in previous large-scale studies, but biotic predictors (potentially reflecting local biotic interactions) have rarely been included. Diatoms may be useful for studying the effect of trophic interactions on community structure since: (1) a majority of experimental studies shows significant grazing effects on diatom species composition, and (2) assemblages can be divided into guilds that have different susceptibility to grazing. We used a dataset from boreal headwater streams in south-central Sweden (covering a spatial extent of ∼14000 km(2)), which included information about diatom taxonomic composition, abundance of invertebrate grazers (biotic factor), environmental (physicochemical) and spatial factors (obtained through spatial eigenfunction analyses). We assessed the relative importance of environmental, biotic, and spatial factors structuring diatom assemblages, and performed separate analyses on different diatom guilds. Our results showed that the diatom assemblages were mainly structured by environmental factors. However, unique spatial and biological gradients, specific to different guilds and unrelated to each other, were also evident. We conclude that biological predictors, in combination with environmental and spatial variables, can reveal a more complete picture of the local vs. regional control of species assemblages in lotic environments. Biotic factors should therefore not be overlooked in applied research since they can capture additional local control and therefore increase accuracy and performance of predictive models. The inclusion of biotic predictors did, however, not significantly influence the unique fraction explained by spatial factors, which suggests low bias in previous assessments of unique regional control of stream assemblages.

Metacommunity structure in a small boreal stream network.

Current ecological frameworks emphasize the relative importance of local and regional drivers for structuring species communities. However, most research has been carried out in systems with discrete habitat boundaries and a clear insular structure. Stream networks deviate from the insular structure and can serve as excellent model systems for studying hierarchical community dynamics over different temporal and spatial extents. We used benthic invertebrate data from streams in a small northern Swedish catchment to test whether metacommunity dynamics change between seasons, across spatial hierarchies (i.e. at the whole catchment scale vs. the scales of first-order and second/third-order sites within the catchment) and between stream-order groups. We assessed metacommunity structure as a function of three relevant dispersal dimensions (directional downstream processes, along-stream dispersal and overland dispersal). These dispersal dimensions were related to species groups with relevant dispersal traits (flying capacity, drift propensity) and dispersal capacities (weak vs. strong) to elucidate whether the observed spatial signals were due to dispersal limitation or mass effects. Results showed complex community organization that varied between seasons, with the scale of observation, and with stream order. The importance of spatial factors and specific dispersal dimensions was highly dependent on the time of sampling and the scale of observation. The importance of environmental factors was more consistent in our analyses, but their effect on species community structure peaked at first-order sites. Our analyses of species dispersal traits were not unequivocal, but indicated that both mass effects and dispersal limitation could simultaneously contribute to the spatial signal at the scale of the whole catchment through different dispersal pathways. We conclude that the study of hierarchically organized ecosystems uncovers complex patterns of metacommunity organization that may deviate substantially from those of systems with insular structure and discrete habitat boundaries. Moreover, we show that dispersal constraints imposed by the dendritic structure of stream networks and distinct dispersal mechanisms (e.g. dispersal limitation) may be evident also at very small spatial extents. Thus, even at this small scale, a landscape management approach that takes the dendritic nature of stream networks into account is needed to effectively conserve stream biodiversity.

Assessing the effects of hydromorphological degradation on macroinvertebrate indicators in rivers: examples, constraints, and outlook.

An extensive amount of literature on linkages between the in-stream physical environment and river benthic macroinvertebrates reports a number of relationships across multiple spatial scales. We analyzed data on different spatial scales to elucidate the linkages between different measurements of hydromorphological degradation and commonly used macroinvertebrate indices. A regression analysis of 1049 sites from 3 countries revealed that the strongest relationship between a biotic metric--average score per taxon--and physiochemical variables (R2 = 0.61) was obtained with a multiple regression model that included concentration of total phosphorus and percent arable land in the catchment, as well as hydromorphological quality variables. Analyses of 3 data sets from streams primarily affected by hydromorphological degradation showed an overall weak relationship (max R2 = 0.25) with the River Habitat Survey data of 28 Swedish streams, whereas moderate (R2 approximately 0.43) relationships with more detailed measurements of morphology were found in 2 Danish studies (39 and 6 streams, respectively). Although evidence exists in the literature on the importance of physical features for in-stream biota in general and macroinvertebrates specifically, we found only relatively weak relationships between various measures of hydromorphological stress and commonly used macroinvertebrate assessment tools. We attribute this to a combination of factors, including 1) the mixed nature of pressures acting on the majority of river reaches, 2) scaling issues (spatial and temporal) when relating habitat surveys to macroinvertebrate assessments, and 3) the scope of commonly used macroinvertebrate assessment systems (mainly focusing on water chemistry perturbation, such as eutrophication and acidification). The need is urgent to develop refined and updated biological assessment systems targeting hydromorphological stress for the use of the European Water Framework Directive (WFD) and national water-related policies.