Ecosystem-level effects of re-oligotrophication and N:P imbalances in rivers and estuaries on a global scale.

Trends and ecological consequences of phosphorus (P) decline and increasing nitrogen (N) to phosphorus (N:P) ratios in rivers and estuaries are reviewed and discussed. Results suggest that re-oligotrophication is a dominant trend in rivers and estuaries of high-income countries in the last two-three decades, while in low-income countries widespread eutrophication occurs. The decline in P is well documented in hundreds of rivers of United States and the European Union, but the biotic response of rivers and estuaries besides phytoplankton decline such as trends in phytoplankton composition, changes in primary production, ecosystem shifts, cascading effects, changes in ecosystem metabolism, etc., have not been sufficiently monitored and investigated, neither the effects of N:P imbalance. N:P imbalance has significant ecological effects that need to be further investigated. There is a growing number of cases in which phytoplankton biomass have been shown to decrease due to re-oligotrophication, but the potential regime shift from phytoplankton to macrophyte dominance described in shallow lakes has been documented only in a few rivers and estuaries yet. The main reasons why regime shifts are rarely described in rivers and estuaries are, from one hand the scarcity of data on macrophyte cover trends, and from the other hand physical factors such as peak flows or high turbidity that could prevent a general spread of submerged macrophytes as observed in shallow lakes. Moreover, re-oligotrophication effects on rivers may be different compared to lakes (e.g., lower dominance of macrophytes) or estuaries (e.g., limitation of primary production by N instead of P) or may be dependent on river/estuary type. We conclude that river and estuary re-oligotrophication effects are complex, diverse and still little known, and in some cases are equivalent to those described in shallow lakes, but the regime shift is more likely to occur in mid to high-order rivers and shallow estuaries.

Ecological relevance of non-perennial rivers for the conservation of terrestrial and aquatic communities.

River conservation efforts traditionally focus on perennial watercourses (i.e., those that do not dry) and their associated aquatic biodiversity. However, most of the global river network is not perennial and thus supports both aquatic and terrestrial biodiversity. We assessed the conservation value of nonperennial rivers and streams (NPRS) in one of Europe's driest regions based on aquatic (macroinvertebrates, diatoms) and terrestrial (riparian plants, birds, and carabid beetles) community data. We mapped the distribution of taxa at 90 locations and across wide environmental gradients. Using the systematic planning tool Marxan, we identified priority conservation sites under 2 scenarios: aquatic taxa alone or aquatic and terrestrial taxa together. We explored how environmental factors (runoff, flow intermittence, elevation, salinity, anthropogenic impact) influenced Marxan's site selection frequency. The NPRS were selected more frequently (over 13% on average) than perennial rivers when both aquatic and terrestrial taxa were considered, suggesting that NPRS have a high conservation value at the catchment scale. We detected an underrepresentation of terrestrial taxa (8.4-10.6% terrestrial vs. 0.5-1.1% aquatic taxa were unrepresented in most Marxan solutions) when priority sites were identified based exclusively on aquatic biodiversity, which points to a low surrogacy value of aquatic taxa for terrestrial taxa. Runoff explained site selection when focusing on aquatic taxa (all best-fitting models included runoff, r2  = 0.26-0.27), whereas elevation, salinity, and flow intermittence were more important when considering both groups. In both cases, site selection frequency declined as anthropogenic impact increased. Our results highlight the need to integrate terrestrial and aquatic communities when identifying priority areas for conservation in catchments with NPRS. This is key to overcoming drawbacks of traditional assessments based only on aquatic taxa and to ensure the conservation of NPRS, especially as NPRS become more prevalent worldwide due to climate change and increasing water demands.

Los esfuerzos de conservación fluvial se enfocan tradicionalmente en los cauces permanentes (aquellos que no se secan) y la biodiversidad acuática asociada. Sin embargo, la mayor parte de la red hidrográfica mundial no es permanente, por lo que sustenta biodiversidad tanto acuática como terrestre. Evaluamos el valor de conservación de los ríos y arroyos no permanentes (RANP) en una de las regiones más secas de Europa con datos de comunidades acuáticas (macroinvertebrados, diatomeas) y terrestres (escarabajos carábidos). Mapeamos la distribución de los taxones en 90 localidades que cubren gradientes ambientales amplios. Con la herramienta de planificación sistemática Marxan identificamos los sitios prioritarios de conservación bajo dos escenarios: considerando sólo los taxones acuáticos o los taxones acuáticos y terrestres juntos. Exploramos cómo los factores ambientales (escorrentía, intermitencia del caudal, altitud, salinidad, impacto antropogénico) influyeron sobre la frecuencia de selección de sitio de Marxan. Los RANP fueron seleccionados con mayor frecuencia (más del 13% en promedio) que los ríos permanentes cuando consideramos los taxones acuáticos y terrestres, lo que sugiere que los RANP tienen un valor elevado de conservación a escala de cuenca. Detectamos que los taxones terrestres estaban infrarrepresentados (8.4-10.6% taxones terrestres vs. 0.5-1.1% acuáticos no tuvieron representación en la mayoría de las soluciones de Marxan) cuando los sitios prioritarios para la conservación se identificaban exclusivamente con la biodiversidad acuática, lo que indica que los taxones acuáticos tienen un reducido valor indicador para los taxones terrestres. La escorrentía determinó la selección de sitios cuando se basó en los taxones acuáticos (los mejores modelos incluyeron la escorrentía, r2 = 0.26-0.27), mientras que la altitud, la salinidad y la intermitencia del caudal fueron más importantes cuando se consideraron ambos grupos. En ambos casos, la frecuencia de selección disminuyó conforme se incrementó el impacto antropogénico. Nuestros resultados resaltan la necesidad de integrar las comunidades terrestres y acuáticas a la identificación de las áreas prioritarias para la conservación de la biodiversidad en cuencas con RANP. Lo anterior es importante para superar las evaluaciones tradicionales basadas solamente en los taxones acuáticos y para garantizar la conservación de los RANP, especialmente ahora que estos son cada vez más frecuentes a nivel mundial debido al cambio climático y a la creciente demanda de agua.

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.

Agricultural impacts on streams near Nitrate Vulnerable Zones: A case study in the Ebro basin, Northern Spain.

Agricultural intensification during the last century has caused river degradation across Europe. From the wide range of stressors derived from agricultural activities that impact rivers, diffuse agricultural pollution has received most of the attention from managers and scientists. The aim of this study was to determine the main stressors exerted by intensive agriculture on streams around Nitrate Vulnerable Zones (NVZs), which are areas of land that drain into waters polluted by nitrates according to the European Nitrate Directive (91/676/EEC). The study area was located in the NW of La Rioja (Northern Spain), which has some of the highest nitrate concentrations within the Ebro basin. The relationships between 40 environmental variables and the taxonomic and functional characteristics of the macroinvertebrate assemblages (which are useful indicators of water quality) were analyzed in 11 stream reaches differentially affected by upstream agricultural activity. The streams affected by a greater percentage of agricultural land cover in the surrounding catchment had significantly higher nitrate concentrations than the remaining sites. However, hydromorphological alteration (i.e. channel simplification, riparian forest and habitat degradation), which is closely linked to agricultural practices, was the main factor affecting macroinvertebrate assemblages. We suggest that "good agricultural practices" should be implemented in streams affected by NVZs to reverse stream degradation, in concordance with the European Water Framework Directive (WFD).

Structural and functional responses of invertebrate communities to climate change and flow regulation in alpine catchments.

Understanding and predicting how biological communities respond to climate change is critical for assessing biodiversity vulnerability and guiding conservation efforts. Glacier- and snow-fed rivers are one of the most sensitive ecosystems to climate change, and can provide early warning of wider-scale changes. These rivers are frequently used for hydropower production but there is minimal understanding of how biological communities are influenced by climate change in a context of flow regulation. This study sheds light on this issue by disentangling structural (water temperature preference, taxonomic composition, alpha, beta and gamma diversities) and functional (functional traits, diversity, richness, evenness, dispersion and redundancy) effects of climate change in interaction with flow regulation in the Alps. For this, we compared environmental and aquatic invertebrate data collected in the 1970s and 2010s in regulated and unregulated alpine catchments. We hypothesized a replacement of cold-adapted species by warming-tolerant ones, high temporal and spatial turnover in taxa and trait composition, along with reduced taxonomic and functional diversities in consequence of climate change. We expected communities in regulated rivers to respond more drastically due to additive or synergistic effects between flow regulation and climate change. We found divergent structural but convergent functional responses between free-flowing and regulated catchments. Although cold-adapted taxa decreased in both of them, greater colonization and spread of thermophilic species was found in the free-flowing one, resulting in higher spatial and temporal turnover. Since the 1970s, taxonomic diversity increased in the free flowing but decreased in the regulated catchment due to biotic homogenization. Colonization by taxa with new functional strategies (i.e. multivoltine taxa with small body size, resistance forms, aerial dispersion and reproduction by clutches) increased functional diversity but decreased functional redundancy through time. These functional changes could jeopardize the ability of aquatic communities facing intensification of ongoing climate change or new anthropogenic disturbances.

Hydrological classification of natural flow regimes to support environmental flow assessments in intensively regulated Mediterranean rivers, Segura River Basin (Spain).

Hydrological classification constitutes the first step of a new holistic framework for developing regional environmental flow criteria: the "Ecological Limits of Hydrologic Alteration (ELOHA)". The aim of this study was to develop a classification for 390 stream sections of the Segura River Basin based on 73 hydrological indices that characterize their natural flow regimes. The hydrological indices were calculated with 25 years of natural monthly flows (1980/81-2005/06) derived from a rainfall-runoff model developed by the Spanish Ministry of Environment and Public Works. These indices included, at a monthly or annual basis, measures of duration of droughts and central tendency and dispersion of flow magnitude (average, low and high flow conditions). Principal Component Analysis (PCA) indicated high redundancy among most hydrological indices, as well as two gradients: flow magnitude for mainstream rivers and temporal variability for tributary streams. A classification with eight flow-regime classes was chosen as the most easily interpretable in the Segura River Basin, which was supported by ANOSIM analyses. These classes can be simplified in 4 broader groups, with different seasonal discharge pattern: large rivers, perennial stable streams, perennial seasonal streams and intermittent and ephemeral streams. They showed a high degree of spatial cohesion, following a gradient associated with climatic aridity from NW to SE, and were well defined in terms of the fundamental variables in Mediterranean streams: magnitude and temporal variability of flows. Therefore, this classification is a fundamental tool to support water management and planning in the Segura River Basin. Future research will allow us to study the flow alteration-ecological response relationship for each river type, and set the basis to design scientifically credible environmental flows following the ELOHA framework.