Drying niches of aquatic macroinvertebrates identify potential biomonitoring indicators in intermittent and ephemeral streams

Intermittent rivers and ephemeral streams (IRES) compose the majority of draining networks on Earth, supporting a unique fraction of biodiversity. Despite their high ecological value, IRES are increasingly threatened by global change and require appropriate biomonitoring and restoration tools. However, indices and indicators used in routine biomonitoring programs are often confounded by drying effects. This occurs because most pollutionsensitive taxa are lost over drying gradients, limiting the utility of current biomonitoring tools in IRES. To address this challenge, there is a need to evaluate which taxa should be used to calculate biomonitoring metrics and indicators over the different portions of the drying gradient. Here, using high-resolution drying data from 33 unpolluted streams, we explored the drying preferences of macroinvertebrates to identify their potential as biomonitoring indicators in IRES. To do this, we characterized macroinvertebrate drying niches and identified those with drying resistance and sensitivity preferences. Next, we evaluated the capacity of functional traits to predict drying specialization and affinity. Finally, to identify potential biomonitoring metrics and indicators for IRES, we evaluated how drying influenced the density, relative abundance and richness of taxa in drying sensitive and drying-resistant niches. Our results identified three macroinvertebrate groups with drying resistant niches (partly tolerant, generalist and specialist taxa) and one group of drying-sensitive taxa. We also found that functional traits had a limited capacity to represent differences in drying niches, with shredding trophic preferences and body size showing the strongest correlations. In addition, we observed that the density, relative abundance and richness of drying-resistant taxa were less influenced by drying intensity than those of drying-sensitive taxa. Finally, we found that some pollution-sensitive taxa with partial (e.g., Lepidostoma), moderate (e.g., Corduliidae) or high drying tolerance (e.g., Nemoura) can serve as potential indicators in IRES. Taken together, our results demonstrate that characterizing drying niches can be a useful strategy for developing biomonitoring tools in IRES and for highlighting the limitations of taxonomic and trait-based approaches

Multiple drying aspects shape dissolved organic matter composition in intermittent streams

Water availability is a fundamental driver of biogeochemical processing in highly dynamic ecosystems such as intermittent rivers and ephemeral streams (IRES), which are recognized as the most common fluvial ecosystem globally. Because of their global extent, IRES have a remarkable contribution to organic matter processing, which is expected to intensify as climate change and water extraction expand IRES extension. Nevertheless, the effect of the complexity of the drying process on river biogeochemistry remains unclear. This study investigated how drying aspects affect the dissolved organic carbon (DOC) concentration and composition in 35 streams along a wide flow-intermittence gradient in the NE Iberian Peninsula. To do that, four drying aspects: annual drying duration, annual frequency, duration of the last drying event, and time since the last drying event were characterized. Results showed that DOC concentration and the contribution of humic-like compounds were positively associated with intensifying drying conditions. In addition, protein-like compounds decreased over the drying gradient. More specifically, changes in DOC concentration were driven mainly by annual drying duration, whereas annual drying frequency and the duration of the last drying event jointly explained dissolved organic matter composition. These results suggest that the quantity and composition of dissolved organic matter in streams respond differently to the temporal aspects of the drying process. Our study can help to better anticipate changes in organic matter in the context of climate change

Remote sensing depicts riparian vegetation responses to water stress in a humid Atlantic region

Riparian areas in the Cantabrian Atlantic ecoregion (northwest Portugal) play a key role in soil formation and conservation, regulation of nutrient and water cycle, creation of landscape aesthetic value and the preservation of biodiversity. The maintenance of their ecological integrity is crucial given the ever increase in multiple anthropogenic (water demand and agriculture) and climatic pressures (droughts and extreme events). We developed a transferable remote sensing approach, taking advantage of the latest freely available technologies (Sentinel-2 and Copernicus Land products), to detect intra-annual and inter-annual changes in riparian vegetation productivity at the river basin scale related to water stress. This study has used the normalized difference vegetation index (NDVI) to investigate riparian vegetation productivity dynamics on three different vegetation types (coniferous, broadleaved and grassland) over the past 5 years (2015-2019). Our results indicated that inter-annual seasonality differed between drier (2017) and wetter (2016) years. We found that intra-annual dynamics of NDVI were influenced by the longitudinal river zonation. Our model ranked first (r2m = 0.73) showed that the productivity of riparian vegetation during the dry season was positively influenced by annual rainfall and by the type of riparian vegetation. The emergent long lags between climatic variation and riparian plant productivity provides opportunities to forecast early warnings of climatically-driven impacts. In addition, the different average productivity levels among vegetation types should be considered when assessing climatic impacts on riparian vegetation. Future applications of Sentinel 2 products could seek to distinguish riparian areas that are likely to be more vulnerable to changes in the annual water balance from those that are more resistant under longer-term changes in climate.Contrato-Programa UIDB/04050/2020. ERA4CS/0004/2016. CLIMALERT: Climate Alert Smart System for Sustainable Water and Agriculture, an ERA-NET initiated by JPI Climate (ERA4CS programme) co-funded by the EU commission (Grant Agreement 690462) and FCT (ERA4CS/0004/2016). This work was supported by the “Contrato-Programa” UIDB/04050/2020 funded by national funds through the FCT I.P. (GP

Diversity mediates the responses of invertebrate density to duration and frequency of rivers annual drying regime

Predicting the impacts of global change on highly dynamic ecosystems requires a better understanding of how communities respond to disturbance duration, frequency and timing. Intermittent rivers and ephemeral streams are dynamic ecosystems that are recognized as the most common fluvial ecosystem globally. The complexity of the drying process can give rise to different annual and antecedent hydrological conditions, but their effect on aquatic communities remains unclear. Here, using aquatic invertebrates from 33 streams across a flow-intermittence gradient, we assessed how annual (drying duration and frequency) and recent drying characteristics (duration of the last dry period and flowing duration since the last rewetting) affect the density and diversity metrics of communities and trophic groups while controlling for other key abiotic factors (dissolved oxygen and altitude). We characterized invertebrate communities using taxonomy and functional traits to capture biological features that increase vulnerability to drying. In addition, using structural equation modelling (SEM), we evaluated pathways by which drying characteristics directly impact invertebrate density and whether diversity indirectly mediates such relationships. We show that drying frequency drove reductions in diversity at the community level and within trophic groups, whereas both the drying duration and frequency had a negative influence on density metrics. Reductions in taxonomic richness were linked to increased annual drying duration, whereas functional diversity declined in response to annual drying frequency. Filterer, predator and shredder trophic groups exhibited the strongest negative responses to drying. Recent drying characteristics had a minor effect on density and diversity metrics. Our SEM results demonstrated that diversity mediates the negative impacts of annual drying duration and frequency on invertebrate density through reductions in their taxonomic richness and functional diversity. Our results underscore the importance of considering multiple drying characteristics together with the interdependence of density and diversity to better anticipate drying responses in freshwater ecosystems. Keywords: biodiversity-ecosystem functioning, drying, flow intermittence, functional traits, intermittent rivers and ephemeral streams, Mediterranean stream

Remote sensing indicators to assess riparian vegetation and river ecosystem health

Environmental managers need information to quickly detect which stressor combinations should be addressed to reverse river degradation across large study areas. The pivotal role of riparian vegetation in regulating thermal regimes and inputs of light, nutrients and organic matter has made it a major target of stressor-mitigation and conservation actions. However, due to the dendritic and extensive nature of river networks, field-based monitoring of local riparian conditions is expensive and time-consuming. Ongoing developments in remote sensing offer an unparalleled opportunity to address this challenge. Nonetheless, there is still a limited understanding of the capacity of remote sensing indicators to predict changes in local riparian and river conditions, urging for local calibration with in situ measurements. This study aims to evaluate the capacity of remote sensing to detect impacts on quality elements commonly used in river biomonitoring: riparian vegetation, abiotic river condition and macrophyte biomass. To this end, four remote sensing metrics were tested against field-based indicators in 50 stream locations from four river basins across the Northwest of Portugal: i) the lateral riparian continuity at reach scale (riparian forest buffer width), ii) the riparian vegetation density at reach scale (Normalized Difference Vegetation Index, NDVI100m), and iii) the land use intensification at both reach (LUI100m) and iv) segment (LUI500m) scales. We found that the combination of remote sensing variables (riparian forest buffer width and the land use intensification index) correlated with riparian vegetation quality and dissolved inorganic nitrogen concentrations. We also found that the riparian vegetation density was able to predict changes in vascular plant biomass except for bryophytes. Our study provides new insights on the capacity of satellite-based indicators to assess riparian and river health, illustrating their utility for land and water managers, to identify and monitor, at a reduced cost and time, po- This work was supported by the River2Ocean project (NORTE-01-0145-FEDER-000068), co-financed by the European Regional Development Fund (ERDF), through Programa Operacional Regional do Norte (NORTE 2020).The work was also supported by the "Contrato-Programa" UIDB/04050/2020 funded by national funds through the FCT I.P., the Centre of Molecular and Environmental Biology (CBMA). CG-C was supported by a "Juan de la Cierva -Incoporacion" contract (MINECO, IJC2018-036642-I). CCS was supported by the "Financiamento Programatico" UIDP/04050/2020 funded by national funds through the FCT I.P

Use of trait concepts and terminology in freshwater ecology: Historic, current, and future perspectives

1.Trait-based approaches have received increasing interest among freshwater scientists given their capacity to predict community structure and biodiversity effects on ecosystem functioning. However, the inconsistent development and use of trait concepts and terms across freshwater scientific disciplines may have limited realisation of the potential of traits. 2.Here, we reviewed trait definitions and terms use to provide recommendations for their consistent application in freshwater science. To do so, we first reviewed literature to identify established trait definitions, historical and current use of trait terms and challenges restricting the application of trait-based approaches in freshwater science. Next, we surveyed 414 freshwater researchers from 54 countries to assess variability in the current use of trait terminology in relation to respondent characteristics (i.e., professional experience, geographical region, research discipline, and focal freshwater ecosystem, biotic group, and ecosystem function). 3.Our literature review identified two well-established trait definitions, which emphasise individual phenotypic characteristics that influence either eco-evolutionary aspects (i.e., organism performance and fitness) or ecosystem dynamics and processes (i.e., responses to the environment and/or effects on ecosystem functioning). Publications used a range of trait-related terms and their frequency of use varied among scientific fields. The term functional trait dominated fields such as biodiversity conservation, environmental sciences and ecology, plant sciences and microbiology. In contrast, the terms biological trait, functional trait, and species trait were used with similar frequencies in fields such as entomology, fisheries, marine and freshwater biology, and zoology. We also found that well-established trait definitions are difficult to apply to freshwater unicellular organisms, colonial multicellular organisms, genomic information, and cultural traits. 4.Our survey revealed highly inconsistent use of trait terms among freshwater researchers. Terms including biological trait, functional trait, structural measure, and ecosystem function were commonly used to describe the same traits or functions. Variability in the use of terms was generally explained by research discipline, geographical region, and focal biotic group and ecosystem functions. 5.We propose making the trait concept flexible enough to be applicable to all freshwater biota and their characteristics, while keeping and integrating links to eco-evolutionary and ecosystem aspects. Specifically, our new definition expands the established functional trait definition by considering also supra-individual scales of trait measurement (colonial- or community-mean traits), genotypic traits (e.g., functional gene markers of enzymes) and cultural traits (e.g., feeding behaviours, communication skills). To reduce terminological ambiguity, we also recommend that researchers define trait terms, prioritising the use of functional trait as an overarching term over alternative terms (e.g., biological trait), and restricting specific terms (e.g., morphological trait) to situations in which such precision is desirable. The findings of our integrative study could help to improve terminological consistency across freshwater disciplines and to better recognise the potential of traits to elucidate the mechanisms behind ecological patterns

Ecological factors determining the distribution and assemblages of the aquatic Hemiptera (Gerromorpha & Nepomorpha) in the Segura River basin (Spain)

Although the Segura River basin is located in one of Europe's most arid regions, it features a wide variety of aquatic ecosystems, some of which are rare within the European continent. Assemblages of aquatic Hemiptera and their indicator species in the Segura River basin, as well as the key environmental factors that determine their distribution, were evaluated in this study. Between 1980 and 2010, a total of 38 species of aquatic Hemiptera were collected in 402 sites that have been classified into 12 types of habitats. Aquatic Hemiptera were well-represented among the different habitats and were widely distributed across the entire study area. Relationships between community structure and environmental variables were evaluated using multivariate analyses, including non-parametric multidimensional scaling (NMDS), principal components analysis (PCA) and distance-based redundancy analysis (dbRDA). Results revealed that the distribution of aquatic Hemiptera was influenced primarily by an environmental gradient from lotic and freshwater headwater environments to lentic and highly-mineralised waters in lower river sections. Hence, the lotic/lentic character of the habitat and its conductivity were the most important factors shaping the spatial distribution of the aquatic Hemiptera in the Segura River basin. Additionally, an indicator species analysis (IndVal) revealed four aquatic Hemiptera assemblage types: one was related with lotic headwater environments, a second was associated with rivers and reservoirs, a third was found primarily in lotic saline environments and a fourth transitional assemblage type was associated with microhabitat availability and included species with a widespread distribution. Defining Hemiptera assemblage types statistically, however, was difficult due to the widespread distribution of species caused by high dispersion capability and low microhabitat requirements.La Cuenca del Río Segura (SE España), a pesar de representar una de las áreas más áridas del Mediterráneo occidental, presenta una amplia variedad de ecosistemas acuáticos, algunos de los cuales son raros a nivel europeo. En dichos ecosistemas, englobados en 12 tipos de hábitats, se han registrado un total de 38 especies de hem'ıpteros acuáticos en 402 estaciones de muestreo entre 1980 y 2010. Se trata de un grupo bien representado en los diferentes hábitats tipo y ampliamente distribuido por el área de estudio. Las relaciones entre la estructura de la comunidad y las variables ambientales fueron estudiadas mediante diferentes técnicas de análisis multivariantes. El an'alisis de escalamiento multidimensional no paramétrico (NMDS), el análisis de componentes principales (PCA) y el análisis de redundancia basado en las distancias (dbRDA) revelaron que la distribución espacial de los hem'ıpteros acu'aticos en la cuenca del río Segura sigue un gradiente ambiental desde ambientes lóticos de agua dulce en cabecera hasta ambientes leníticos mineralizados en desembocadura. Por tanto, el tipo de h'abitat (lótico o lenítico) y la conductividad son los principales factores determinantes de la distribución de las especies. Finalmente, un análisis de especies indicadoras (IndVal) mostró 4 tipos de asociaciones de especies características de diferentes tipos de hábitats: ambientes lóticos de cabecera, ríos y embalses, ambientes leníticos salinos y un último grupo de transición entre los anteriores. A pesar de estos resultados, los análisis estadísticos muestran que es difícil definir grupos para los hemípteros acuáticos debido a que muchas de estas especies presentan una amplia distribución como resultado de su alta capacidad de dispersión y de su baja especificidad de hábitat

Populations of high-value predators reflect the traits of their prey

The extent to which prey traits combine to influence the abundance of predators is still poorly understood, particularly for mixed predators in sympatry and in aquatic ecosystems. In this study, we characterise prey use and distribution in iconic bird (grey wagtails and Eurasian dippers) and fish species (brown trout and Atlantic salmon) to assess whether prey traits could predict populations of these four riverine predators. Specifically, we hypothesised that: 1) prey key traits would predict predator populations more effectively than 2) diversity of prey traits, 3) the taxonomic abundance or richness of prey (known as traditional or mass‐effect types of biodiversity) or 4) the prevailing environmental conditions. Combined predator population sizes were predicted better by a few key traits – specifically those revealing prey habitat use, size and drifting behaviour – than by prey diversity or prey trait diversity or environmental conditions. Our findings demonstrate that the complex relationships between prey assemblages and multiple predator species can be represented mechanistically when the key prey traits that govern encounter and consumption rates are identified. Given their apparent potential to reveal trophic relationships, and to complement more traditional measures of prey abundance, we advocate further development of trait‐based approaches in predator–prey research

Evaluating riparian solutions to multiple stressor problems in river ecosystems - A conceptual study

Rivers are among the most sensitive of all ecosystems to the effects of global change, but options to prevent, mitigate or restore ecosystem damage are still inadequately understood. Riparian buffers are widely advocated as a cost-effective option to manage impacts, but empirical evidence is yet to identify ideal riparian features (e.g. width, length and density) which enhance ecological integrity and protect ecosystem services in the face of catchment-scale stressors. Here, we use an extensive literature review to synthesise evidence on riparian buffer and catchment management effects on instream environmental conditions (e.g. nutrients, fine sediments, organic matter), river organisms and ecosystem functions. We offer a conceptual model of the mechanisms through which catchment or riparian management might impact streams either positively or negatively. The model distinguishes scale-independent benefits (shade, thermal damping, organic matter and large wood inputs) that arise from riparian buffer management at any scale from scale-dependent benefits (nutrient or fine sediment retention) that reflect stressor conditions at broader (sub-catchment to catchment) scales. The latter require concerted management efforts over equally large domains of scale (e.g. riparian buffers combined with nutrient restrictions). The evidence of the relationships between riparian configuration (width, length, zonation, density) and scale-independent benefits is consistent, suggesting a high certainty of the effects. In contrast, scale-dependent effects as well as the biological responses to riparian management are more uncertain, suggesting that ongoing diffuse pollution (nutrients, sediments), but also sources of variability (e.g. hydrology, climate) at broader scales may interfere with the effects of local riparian management. Without concerted management across relevant scales, full biological recovery of damaged lotic ecosystems is unlikely. There is, nevertheless, sufficient evidence that the benefits of riparian buffers outweigh potential adverse effects, in particular if located in the upstream part of the stream network. This supports the use of riparian restoration as a no-regrets management option to improve and sustain lotic ecosystem functioning and biodiversity

Do all roads lead to Rome? Exploring community trajectories in response to anthropogenic salinization and dilution of rivers

Abiotic stress shapes how communities assemble and support ecological functions. However, it remains unclear whether artificially increasing or decreasing stress levels would lead to communities assembling predictably along a single axis of variation or along multiple context-dependent trajectories of change. In response to stress intensity alterations, we hypothesize that a single trajectory of change occurs when trait-based assembly prevails, while multiple trajectories of change arise when dispersal-related processes modify colonization and trait-filtering dynamics. Here, we tested these hypotheses using aquatic macroinvertebrates from rivers exposed to gradients of natural salinity and artificially diluted or salinized ion contents. Our results showed that trait-filtering was important in driving community assembly in natural and diluted rivers, while dispersal-related processes seemed to play a relevant role in response to salinization. Salinized rivers showed novel communities with different trait composition, while natural and diluted communities exhibited similar taxonomic and trait compositional patterns along the conductivity gradient. Our findings suggest that the artificial modification of chemical stressors can result in different biological communities, depending on the direction of the change (salinization or dilution), with trait-filtering, and organism dispersal and colonization dynamics having differential roles in community assembly. The approach presented here provides both empirical and conceptual insights that can help in anticipating the ecological effects of global change, especially for those stressors with both natural and anthropogenic origins