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Many inland waters exhibit complete or partial desiccation, or have vanished due to global change, exposing sediments to the atmosphere. Yet, data on carbon dioxide (CO2) emissions from these sediments are too scarce to upscale emissions for global estimates or to understand their fundamental drivers. Here, we present the results of a global survey covering 196 dry inland waters across diverse ecosystem types and climate zones. We show that their CO2 emissions share fundamental drivers and constitute a substantial fraction of the carbon cycled by inland waters. CO2 emissions were consistent across ecosystem types and climate zones, with local characteristics explaining much of the variability. Accounting for such emissions increases global estimates of carbon emissions from inland waters by 6% (~0.12 Pg C y−1). Our results indicate that emissions from dry inland waters represent a significant and likely increasing component of the inland waters carbon cycle., Many inland waters seasonally or permanently dry up, thus exposing sediments to the atmosphere. Here the authors show that a substantial amount of CO2 is emitted from these dry sediments, increasing current inland water carbon flux estimates by 6%.

International audience; Non-perennial rivers and streams (NPRS) cover > 50% of the global river network. They are particularly predominant in Mediterranean Europe as a result of dry climate conditions, climate change and land use development. Historically, both scientists and policy makers underestimated the importance of NRPS for nature and humans alike, mainly because they have been considered as systems of low ecological and economic value. During the past decades, diminishing water resources have increased the spatial and temporal extent of artificial NPRS as well as their exposure to multiple stressors, which threatening their ecological integrity, biodiversity and ecosystem services. In this paper, we provide a comprehensive overview of the structural and functional characteristics of NPRS in the European Mediterranean, and discuss gaps and problems in their management, concerning their typology, ecological assessment, legislative and policy protection, and incorporation in River Basin Management Plans. Because NPRS comprise highly unstable ecosystems, with strong and often unpredictable temporal and spatial variability - at least as far as it is possible to assess - we outline the future research needs required to better understand, manage and conserve them as highly valuable and sensitive ecosystems. Efficient collaborative activities among multidisciplinary research groups aiming to create innovative knowledge, water managers and policy makers are urgently needed in order to establish an appropriate methodological and legislative background. The incorporation of NPRS in EU-Med River Basin Management Plans in combination with the application of ecological flows is a first step towards enhancing NPRS management and conservation in order to effectively safeguard these highly valuable albeit threatened ecosystems

We define Dry Rivers as those whose usual habitat in space and time are dry channels where surface water may interrupt dry conditions for hours or a few days, primarily after heavy rainfall events that are variable in time and that usually lead to flash floods, disconnected from groundwater and thereby unable to harbor aquatic life. Conceptually, Dry Rivers would represent the extreme of the hydrological continuum of increased flow interruption that typically characterizes the non-perennial rivers, thus being preceded by intermittent and ephemeral rivers that usually support longer wet phases, respectively. This paper aims to show that Dry Rivers are ecosystems in their own right given their distinct structural and functional characteristics compared to other non-perennial rivers due to prevalence of terrestrial conditions. We firstly reviewed the variety of definitions used to refer to these non-perennial rivers featured by a predominant dry phase with the aim of contextualizing Dry Rivers. Secondly, we analyzed existing knowledge on distribution, geophysical and hydrological features, biota and biogeochemical attributes that characterize Dry Rivers. We explored the capacity of Dry Rivers to provide ecosystem services and described main aspects of anthropogenic threats, management challenges and the conservation of these ecosystems. We applied an integrative approach that incorporates to the limnological perspective the terrestrial view, useful to gain a better understanding of Dry Rivers. Finally, we drew main conclusions where major knowledge gaps and research needs are also outlined. With this paper, we ultimately expect to put value in Dry Rivers as non-perennial rivers with their own ecological identity with significant roles in the landscape, biodiversity and nutrient cycles, and society; thus worthy to be considered, especially in the face of exacerbated hydrological drying in many rivers across the world.

The dataset (Data File S1; DOI: 10.6084/m9.figshare.8863721) and the R code used to generate the results (Code S1; DOI: 10.6084/m9.figshare.8863655), including step by step explanations of the statistical tests, have been deposited in Figshare Digital Repository (https://figshare.com/projects/Sediment_Respiration_Pulses_in_Intermittent_Rivers_and_Ephemeral_Streams/66104)., Intermittent rivers and ephemeral streams (IRES) may represent over half the global stream network, but their contribution to respiration and carbon dioxide (CO2) emissions is largely undetermined. In particular, little is known about the variability and drivers of respiration in IRES sediments upon rewetting, which could result in large pulses of CO2. We present a global study examining sediments from 200 dry IRES reaches spanning multiple biomes. Results from standardized assays show that mean respiration increased 32–66-fold upon sediment rewetting. Structural equation modelling indicates that this response was driven by sediment texture and organic matter quantity and quality, which, in turn, were influenced by climate, land use and riparian plant cover. Our estimates suggest that respiration pulses resulting from rewetting of IRES sediments could contribute significantly to annual CO2 emissions from the global stream network, with a single respiration pulse potentially increasing emission by 0.2–0.7%. As the spatial and temporal extent of IRES increases globally, our results highlight the importance of recognizing the influence of wetting-drying cycles on respiration and CO2 emissions in stream networks., We thank Y. Etxeberria, L. Sánchez, C. Gutiérrez, G. LeGoff and B. Launay for laboratory support. DvS was supported by a Short-Term Scientific Mission of the COST Action CA15113 (SMIRES, Science and Management of Intermittent Rivers and Ephemeral Streams, www.smires.eu), supported by COST (European Cooperation in Science and Technology) and received additional funding from the EU’s 7th Framework Programme for research, technological development and demonstration under grant agreement No. 603629 (GLOBAQUA) and a Grant for Research Groups of the Basque University System (IT-951-16) funded by the Basque Government. RM and BO were supported by the Spanish Ministry of Science, Innovation and Universities through project C-HYDROCHANGE (CGL2017-86788-C3-2-P and CGL2017-86788-C3-3-P). FA was funded by the Swiss National Science Foundation grants No. PP00P3_150698 and PP00P3_179089. NC was supported by the EU project LIFE+ TRivers (LIFE13 ENV/ES/000341). SDL received funding from the EU’s Horizon 2020 research and innovation programme under the Marie Sklodowska-Curie grant agreement No 748625. PP and MP were supported by INTER-COST project LTC17017. The authors declare that they have no competing interests.

Intermittent rivers and ephemeral streams (IRES) are dynamic ecosystems that shift between aquatic and terrestrial states. IRES are widespread, abundant and increasing in extent, but developing biomonitoring programmes to determine their ecological quality is challenging. To date, quality assessments have focused on the aquatic organisms present during wet phases, whereas dry-phase communities remain poorly characterized. We examined multiple biotic groups present in dry IRES channels, to compare assemblages at sites impacted and unimpacted by human activity and to evaluate the potential of each group as an ecological quality indicator. We explored existing, unpublished data for three biotic groups: an aquatic microflora (diatoms), an aquatic fauna (the invertebrate ‘seedbank’), and a mixed flora (aquatic and terrestrial plants); notably, we did not source data for terrestrial assemblages with high potential to act as indicators. Diatom and plant assemblage composition differed between impacted and unimpacted sites, and the latter assemblages were more diverse and included more indicator taxa. Invertebrate seedbank taxa richness was higher at unimpacted sites but compositional differences were not detected, probably due to the coarse taxonomic resolution to which abundant taxa were identified. Performance of standard indices of ecological quality was variable, but differences were identified between impacted and unimpacted conditions for all biotic groups. Our results can inform the enhancement of biomonitoring programmes designed to characterize IRES ecological quality in relation to legislative targets. We highlight the need to integrate wet- and dry-phase survey data in holistic quality assessments. Although we suggest diatoms, aquatic plants and the aquatic invertebrate seedbank as having the potential to inform assessment of dry-phase ecological quality, we highlight the need for research to further characterize these aquatic groups and, crucially, to explore terrestrial assemblages with high potential to act as dry-phase quality indicators. ISSN:1470-160X ISSN:1872-7034

Climate change and human pressures are changing the global distribution and the extent of intermittent rivers and ephemeral streams (IRES), which comprise half of the global river network area. IRES are characterized by periods of flow cessation, during which channel substrates accumulate and undergo physico‐chemical changes (preconditioning), and periods of flow resumption, when these substrates are rewetted and release pulses of dissolved nutrients and organic matter (OM). However, there are no estimates of the amounts and quality of leached substances, nor is there information on the underlying environmental constraints operating at the global scale. We experimentally simulated, under standard laboratory conditions, rewetting of leaves, riverbed sediments, and epilithic biofilms collected during the dry phase across 205 IRES from five major climate zones. We determined the amounts and qualitative characteristics of the leached nutrients and OM, and estimated their areal fluxes from riverbeds. In addition, we evaluated the variance in leachate characteristics in relation to selected environmental variables and substrate characteristics. We found that sediments, due to their large quantities within riverbeds, contribute most to the overall flux of dissolved substances during rewetting events (56%–98%), and that flux rates distinctly differ among climate zones. Dissolved organic carbon, phenolics, and nitrate contributed most to the areal fluxes. The largest amounts of leached substances were found in the continental climate zone, coinciding with the lowest potential bioavailability of the leached OM. The opposite pattern was found in the arid zone. Environmental variables expected to be modified under climate change (i.e. potential evapotranspiration, aridity, dry period duration, land use) were correlated with the amount of leached substances, with the strongest relationship found for sediments. These results show that the role of IRES should be accounted for in global biogeochemical cycles, especially because prevalence of IRES will increase due to increasing severity of drying events., In this study we experimentally simulated under laboratory conditions rewetting of leaves, riverbed sediments, and epilithic biofilms collected during the dry phase from intermittent rivers and ephemeral streams covering global spatial scale. We determined the amounts and quality of the leached nutrients and dissolved organic matter, assessed their inter‐substrate and cross‐climate differences, and estimated areal fluxes from 1 m2 of riverbeds. In addition, we evaluated the variance in leachate characteristics related to selected environmental variables and substrate characteristics.

Debris piles and dry pools are common landscape elements along ephemeral river channels. However, their effects on resource availability and arthropod species diversity have been neglected so far. We investigated whether debris piles and pools along dry riverbeds generate nodes of environmental resources, influence microbial activity, and provide key habitats for ground-dwelling arthropods. We studied three different habitats (channel, debris pile and dry pool) to analyze organic matter, nutrients and microbial activity in dry sediments, and included two further habitats (riparian and upland areas) for ground-dwelling arthropod assemblages along a major ephemeral river (Ugab River, NW Namibia). We observed higher concentrations of organic matter and macro- and micro-nutrients in debris piles than in pools, and higher concentrations in both landscape elements than in the sandy channel matrix. Microbial activity showed a positive relationship with organic matter and nutrient availability in sediments. Debris piles, dry pools and channels harbored diverse and abundant ground-dwelling arthropod assemblages similar to adjacent terrestrial areas. Debris piles and pools differed in assemblage composition not only from riparian and upland areas but also from the sandy channel matrix. Overall, debris piles and dry pools acted as nodes of soil fertility, enhanced microbial activity, and shaped ground-dwelling arthropod assemblages.

Functional trait diversity can provide insight into ecosystem function beyond that provided by species diversity measures. The relationship between functional diversity and natural stressors has received less attention compared to anthropogenic stressors. In this study, we investigated how two natural stressors, water salinity and flow intermittence, affect functional richness and functional redundancy of aquatic invertebrate communities using seven biological traits and 39 modalities. For this purpose, we characterized these functional diversity measures in 22 Mediterranean streams with a gradient of natural salinity and flow intermittence. Our findings showed that both functional richness and functional redundancy decreased with increased stress by water salinity and flow intermittence for all the studied traits but more rapidly for the former, suggesting that water salinity is a stronger environmental stressor than flow intermittence. Our study also described an antagonistic interaction of the two study stressors, in which the net effect of both is less than the sum of their independent effects. This study emphasizes that in saline streams, characterized by lower functional richness and functional redundancy, the loss of any taxon can have a huge impact on community functioning. In particular, the functional singularity of saline intermittent streams makes them extremely sensitive to additional anthropogenic impacts. In the context of future global change scenarios, which predict higher flow intermittence and water salinity, this study gives a better understanding of the functional features of these types of ecosystems.

Saline streams occur naturally and they are distributed worldwide, particularly in arid and semiarid regions, but human activities have also increased their number in many parts of the world. Little attention has been paid to assess increasing salt effects on organic matter decomposition. The objectives of this study were to analyse wood breakdown rates and how salinity affects them in 14 streams that exemplify a natural salinity gradient. We also analysed the effect of this gradient on changes in wood chemical composition, fungal biomass and microbial activity. Our results showed low breakdown rates (0.0010-0.0032 d(-1)), but they fell within the same range as those reported in freshwater streams when a similar woody substrate was used. However, salinity had a negative effect on the breakdown rates and fungal biomass along the salinity gradient, and led to noticeable changes in wood composition. Water salinity did not affect microbial activity estimated using hydrolysis of fluorescein diacetate. Variation in breakdown rates and fungal biomass across streams was mediated mainly by salinity, and later by stream discharge. Despite the role of fungi in stick breakdown, the potential wood abrasion by salts must be analysed in detail to accurately understand the effect of increasing salinity on organic matter breakdown. Finally, our results indicate that increased salinity worldwide by human activities or by the global warming would imply organic matter breakdown and mineralisation slowing down, even in natural saline streams. However, because many variables are implicated, the final effect of climatic change on organic matter decomposition in streams is difficult to predict.

Water scarcity represents an important driver of stream microorganisms and N transformations rates. In semiarid regions many agricultural streams rich in NO 3 − undergo periods of flow cessation of severe duration and spatial extent. After drought, rainfalls tend to firstly rewet stream sediments by creating isolated pools that can remain in streambeds until the entire stream is reconnected. Sediment rewetting may lead the reactivation of nutrient processing, yet usually causes release of inorganic N. Thus, information about the role of sediments during this rewetting period on N processing and fluxes downstream is especially valuable when semiarid watercourses are affected by chronic NO 3 − exposure. We conducted a microcosm study to evaluate how a short-term sediment rewetting over a 7 days period in disconnected pools can control denitrification rates and N availability in an agriculturally impacted semiarid stream. We observed a significant stimulation of denitrification in dry sediments ( −2 h −1 ) upon rewetting with a maximum rate of 830 μg N m −2 h −1 after 24 h. In parallel, NO 3 − in dry sediments (1170 mg m −2 ) dropped considerably after 7 days of rewetting (72 mg m −2 ), despite the fact that denitrification was restrained from 24 h thereafter. However, from this time a notably increase of NH 4 + concentration was detected until the end, which suggests that at this point, NO 3 − is transformed into NH 4 + through dissimilatory reduction (DNRA) supported by low Eh (40 mV). If compared with NO 3 − , NH 4 + can be either more easily retained in sediments or assimilated preferably by organisms. We highlight that the diversification of N species provided by isolated pools creation just before flow resumption may progressively improve water quality in NO 3 − -rich streams, which is a key aspect in the face of future agriculture intensification and water scarcity scenario.

International audience; Perennial rivers and streams make a disproportionate contribution to global carbon (C) cycling. However, the contribution of intermittent rivers and ephemeral streams (IRES), which sometimes cease to flow and can dry completely, is largely ignored although they represent over half the global river network. Substantial amounts of terrestrial plant litter (TPL) accumulate in dry riverbeds and, upon rewetting, this material can undergo rapid microbial processing. We present the results of a global research collaboration that collected and analysed TPL from 212 dry riverbeds across major environmental gradients and climate zones. We assessed litter decomposability by quantifying the litter carbon-to-nitrogen ratio and oxygen (O2) consumption in standardized assays and estimated the potential short-term CO2 emissions during rewetting events. Aridity, cover of riparian vegetation, channel width and dry-phase duration explained most variability in the quantity and decomposability of plant litter in IRES. Our estimates indicate that a single pulse of CO2 emission upon litter rewetting contributes up to 10% of the daily CO2 emission from perennial rivers and stream, particularly in temperate climates. This indicates that the contributions of IRES should be included in global C-cycling assessments.

Intermittent streams are subject to high levels of environmental variation. However, little is known about how biota responds to river drying across the channel-to-upland habitat gradient. This is an important shortcoming because assumes that intermittent river habitats and metacommunities are static. Here we studied how river drying affects the spatial and temporal variation in ground-dwelling arthropod communities (spiders, beetles, and ants) across the lateral habitat gradient. We asked whether particular habitats, moments, or species contribute disproportionally to beta diversity. To this end, we monitored two perennial-intermittent reach pairs during an entire drying cycle, and applied beta-diversity partitioning methods to highly-resolved taxonomy data. We predicted that: (i) intermittent reaches would accumulate higher levels of diversity (alpha and beta) than perennial reaches; (ii) dry riverbeds would harbor more species and more unique species; (iii) alpha and beta diversity would be temporally more variable in intermittent than in permanent reaches; and (iv) species dispersal limitation would explain variation in contributions to beta diversity. Intermittent reaches presented higher alpha and beta diversity, with dry channels harboring more unique species and more species than any other habitat. Arthropod metacommunities were more variable across space than over time, and temporal turnover was significant—with species dispersal limitation being positively associated with contributions to beta diversity. Our results elevate the role of dry channels in intermittent river ecology, and show that previous research has likely underestimated their contributions to river-wide biodiversity. Our findings call for the need to integrate the dry phase of intermittent streams into monitoring and conservation programs.

The assessment of the ecological status of water bodies, as requires by the European Water Framework Directive, can raise a number of problems when applied to temporary streams. These problems are because of the particular physical, chemical and biological conditions resulting from the recurrent cessation of flow or even the complete drying of the stream beds. In such non-permanent water bodies, the reference quality standards developed for permanent streams may only be applicable under certain circumstances or may not be applicable at all. Work conducted within the collaborative EU-funded project Mediterranean Intermittent River ManAGEment (MIRAGE) has addressed most of these difficulties and has used diverse approachesto solve them. These approaches have been brought together inthe so-called MIRAGEToolbox. This toolbox consists of a series of methodologies that are designed to be used in a sequential manner to allow the establishment of the ecological and chemical status of temporary streams and to relate these findings to the hydrological status of the streams. The toolbox is intended to serve the following purposes: (i) the determination of the hydrological regime of the stream; (ii) the design of adequate schedules for biological andchemicalsamplingaccordingtotheaquaticstateofthestream;(iii)thefulfillmentofcriteriafordesigningreferenceconditionstations;(iv)the analysis of hydrological modifications of the stream regime (with the definition of the hydrological status); and (v) the development of new methods to measure the ecological status (including structural and functional methods) and chemical status when the stream’s hydrological conditions are far from those in permanent streams. Copyright © 2014 John Wiley & Sons, Ltd.

As saline streams are geographically widespread in arid and semiarid regions, flow intermittence frequently occurs as another stressor factor apart from salinity. Rambla Salada is a temporary naturally saline stream with an intermittent reach upstream. This stream is an ideal scenario to study the effects of intermittent stream flow on macroinvertebrate community composition and biological traits in a naturally saline Mediterranean stream. This study analysed three sites with different hydrological regimes (one intermittent and two perennials). Flow intermittence exerted low pressure on the macroinvertebrate composition and biological traits which led to the loss of drought-intolerant species and taxa rather than acting as a selective force to promote desiccation-resistant taxa. Macroinvertebrate community at the intermittent site was a subset of the community found in perennial sites, and the presence of flier taxa at this site helped avoid flow cessation. These minor changes have consequences to assess the ecological quality of these saline temporary streams in the context of the Water Framework Directive, given the major differences revealed by some indices between the intermittent and perennial sites as the former obtained lower values due to the presence of few desiccation-intolerant species, which significantly increased the value of those biological indices.

Most streams draining to the Mediterranean basin are temporary. As a result of their hydrological regime, temporary streams are affected by drying and rewetting periods. Drying can alter in-stream nitrogen (N) availability and reduce N processing rates and subsequent retention after re-wetting. We sought to determine if hydrologic drying modifies reach-scale sediment chemical properties and constrains the response of N processing to rewetting. We compared different abiotic characteristics of sediments and nitrification and denitrification rates between a perennial and intermittent reach in the same stream over a wet period, when surface water flowed in both reaches, and a dry period, when the intermittent reach dried up. We analyzed N processing rates by incubating sediments with stream water, thereby simulating a rewetting when sediments from the intermittent reach were dry. We found that drying increased the sediment nitrate (NO3−) content. Conversely, drying did not reduce the recovery of N processing rates to pre-dry levels after simulated flooding conditions. Our results suggest that dry reaches may act as a potential NO3− source by releasing downstream NO3− pulses after stream flow recovery. Given the European Water Framework Directive requirements to assess stream ecological status, these N pulses following rewetting should be considered when designing management plans in temporary streams. Our study highlights the rapid response of in-stream N processing to rewetting period following a drought. This high resilience to process N should be seen as a vital ecosystem service provided by temporary streams despite annual dry periods.

In 2011, the European Council stressed the significance of water quality for sustainable development in Europe and emphasized the need for better integration of the water policy objectives into the Common Agriculture Policy reform in rural areas. Since 2000, the Water Framework Directive (WFD) has required the EU Member States to target good ecological status for their water bodies. However, the implementation of the Directive for the numerous Mediterranean temporary streams has been delayed indicating the need for elaboration of the tools and methods that address the special characteristics of such water bodies. This requirement has been addressed by the recently completed MIRAGE project. In the context of the recent publication of the European Commission's Blueprint to Safeguard Europe's Waters, the MIRAGE-proposed framework for the characterization of the eco-hydrological dynamics and the systematic description of the measured impact for temporary rivers could bring considerable added value to the EU revision of all relevant water policies. The project recommends additions to WFD articles including an explicit definition of temporary rivers, adaptation of environmental objectives to their peculiarities and establishment of a proper method to determine the initial status and specific actions in River Basin Management Plans.

Intermittent rivers and ephemeral streams (IRES), by alternating flowing and dry phases, are coupled aquatic and terrestrial ecosystems hosting unique combinations of amphibian, reptile, bird, and mammal species (hereafter, wildlife) across different world regions. IRES are valuable ecosystems for wildlife providing essential ecological functions during both wet and dry phases. Concurrently, wildlife exerts direct and indirect biotic control of the structure (biodiversity and physical modification) and function (nutrient cycling and connectivity) of IRES, playing important ecological roles. Although flow intermittence has direct and indirect effects on the presence, abundance, and/or activity of vertebrate species, different adaptations allow them to cope with flow intermittence. Many anthropogenic activities (e.g., flow alteration, habitat degradation, water pollution) combined with the spread of invasive species, emerging diseases, and climate change threaten the local habitats and biological diversity of wildlife in IRES. Proper IRES management should involve a better knowledge of species’ flow requirements, as well as the preservation or restoration of natural flow regimes that ensure the functions of both aquatic and terrestrial ecological refuges.

The contribution of rare taxa to aquatic bioassessments remains a subject of debate, and generates contrasting positions among researchers. Very little is known about the effect of low-abundance taxa (LAT) for calculating both single and multimetric macroinvertebrate-based indices, as well as the ecological status classification. In this study, we aimed to: i) identify the aquatic macroinvertebrates that need special attention during index applications given their low abundance; ii) analyse the effect of excluding LAT on single (IBMWP and IASPT) and multimetric (STAR_ICMi) biological indices; and iii) investigate the influence of LAT on river ecological status assessments. To this end, two different river basins in SE Spain and N Italy with contrasting climatic conditions and river types were selected. Our results showed that almost all the taxa at the family level can act as low-abundance taxa. In particular, the LAT belonged mainly to orders Diptera, Trichoptera, Coleoptera, Gastropoda and Hemiptera. The IndVal analysis stressed Tabanidae, Cordulegasteridae and Hydroptilidae as the most characteristic low-abundance families in the Spanish data set, while Dryopidae and Athericidae were identified mostly in N Italy. Excluding LAT affected the studied index values and the resulting bioassessment classification, except for the IASPT index. Loss of the entire LAT pool reduced the ecological status for 78% of the samples for the IBMWP index. Changing took place in 41% of the samples when considering the STAR_ICM index. Relevant changes were detected even when considering loss of 50% of the LAT, especially with the IBMWP index. Similar values and patterns were obtained in each considered quality class and river type. Our results provide useful information about controversial taxa and stress the significance of LAT in river biomonitoring. Excluding LAT is discouraged, although different responses according to the considered index were detected. The IBMWP index always obtained lower values (coupled mainly with an underestimation of ecological classes), while STAR_ICMi and, especially the IASPT index, were less affected by excluding LAT. Paying special attention to all the protocol application stages is recommended, with emphasis placed when using the IBMWP index.

Autumnal input of leaf litter is a pivotal energy source in most headwater streams. In temporary streams, however, water stress may lead to a seasonal shift in leaf abscission. Leaves accumulate at the surface of the dry streambed or in residual pools and are subject to physicochemical preconditioning before decomposition starts after flow recovery. In this study, we experimentally tested the effect of photodegradation on sunlit streambeds and anaerobic fermentation in anoxic pools on leaf decomposition during the subsequent flowing phase. To mimic field preconditioning, we exposed Populus tremula leaves to UV-VIS irradiation and wet-anoxic conditions in the laboratory. Subsequently, we quantified leaf mass loss of preconditioned leaves and the associated decomposer community in five low-order temporary streams using coarse and fine mesh litter bags. On average, mass loss after approximately 45 days was 4 and 7% lower when leaves were preconditioned by irradiation and anoxic conditions, respectively. We found a lower chemical quality and lower ergosterol content (a proxy for living fungal biomass) in leaves from the anoxic preconditioning, but no effects on macroinvertebrate assemblages were detected for any preconditioning treatment. Overall, results from this study suggest a reduced processing efficiency of organic matter in temporary streams due to preconditioning during intermittence of flow leading to reduced substrate quality and repressed decomposer activity. These preconditioning effects may become more relevant in the future given the expected worldwide increase in the geographical extent of intermittent flow as a consequence of global change. © 2011 Springer Basel AG.

Biological indicators are increasingly being used as integrative measures of the ecosystem health in streams, particularly those using macroinvertebrate assemblage composition. Monitoring biological quality of rivers has not a long tradition in some Mediterranean European countries like Spain. Several macroinvertebrate metrics have been recently proposed to assess ecological status in Mediterranean streams, so it is necessary to compare the use of proposed biological quality metrics to select the most appropriate ones. In the present work, two classic richness metrics (total number of families and number of the Ephemeroptera, Plecoptera and Trichoptera families), three indices (IBMWP, IASPT and t-BMWQ) and two multimetric indices, recently proposed to be used in Mediterranean streams (ICM-9 and ICM-11a or IMMi-L), were compared by the analysis of the sensitivity of these metrics to a multiple stressor gradient which reflected the main pressures present in the study area. For this purpose, data from 193 sites sampled in spring (95 reference sites and 98 disturbed sites) belonging to five different Mediterranean stream types present in 35 basins were studied. The results showed that the adjusted regression coefficients (r2) for all seven metrics in the exponential regression models were higher than linear ones, thus indicating an exponential relationship between metrics and the environmental alteration. The two studied multimetric indices presented higher regression coefficients (r2 = 0.590–0.669) than the three indices (r2 = 0.524–0.574) and the two metrics (r2 = 0.471–0.525), therefore showing a better response to a stressor gradient in Mediterranean streams. Within the multimetric indices group, ICM-11a showed the highest regression coefficients. Based on the results obtained, we suggest using the ICM-11a, apart from the IBMWP, to assess ecological status in Mediterranean streams.

Correction to: Nature Geoscience https://doi.org/10.1038/s41561-018-0134-4, published online 21 May 2018. In the version of this Article originally published, the affiliation for M. I. Arce was incorrect; it should have been: 5Leibniz Institute of Freshwater Ecology and Inland Fisheries (IGB), Berlin, Germany. This has now been corrected in the online versions of the Article.

We investigated the seasonal changes in macroinvertebrate reference communities in four Mediterranean stream types (temporary, evaporite calcareous at medium altitude, siliceous headwaters at high altitude, and cal- careous headwaters at medium and high altitudes) and the interannual changes in the two headwaters stream types in Spain. Eighty-eight seasonal reference sites distributed into 23 basins were sampled on three occasions (spring, summer and autumn of 2003), and 18 interannual reference sites distributed in 6 basins were sampled in the autumn of 2003, 2004 and 2005 to examine this temporal variability. Interannual reference sites were a subset of seasonal reference sites. The analysis of similarity (ANOSIM) performed on Bray-Curtis similarity distances, using pres- ence-absence data, showed no seasonal or interannual changes in the macroinvertebrate communities. The infl uence of seasonal and interannual variability was also tested in all the stream types using 18 macroinvertebrate metrics classifi ed as richness, index, multimetric index, tolerance/intolerance and diversity. ANOVAs showed no seasonal differences in any of the studied metrics for temporary and evaporite calcareous at medium altitude and most of the metrics in the two headwaters stream types. This suggests the suitability of using a single season approach for the biomonitoring purposes of these metrics. Conversely, the seasonal differences detected in the metrics related with EPT (Ephemeroptera, Plecoptera and Trichoptera) and OCH (Odonata, Coleoptera and Heteroptera) taxa in calcareous headwaters indicate possible differences in the relative presence of macrohabitats (riffl es and pools) as a result of fl ow variation. No interannual changes were detected in any of the metrics except EPT/OCH in siliceous headwaters. However, the large variability in the annual rainfall in this study area suggests that this three-year study period may be too short to assess the effect of climatic variations on the ecological status assessment. In general, the lower temporal variability (measured as seasonal and interannual coeffi cients of variation) of the taxon richness metric (S) and the two studied indices (IBMWP and IASPT), compared with the other metrics, make them a priori robust indicators to assess ecological status in Mediterranean streams.

In Spain, a national project known as GUADALMED, focusing on Mediterranean streams, has been carried out from 1998 to 2005 to implement the European water framework directive (WFD) requirements. One of the main objectives of the second phase of the project (2002-2005) was to develop a predictive system for the Spanish Mediterranean aquatic macroinvertebrate communities. A combined-season (spring, summer, and autumn) predictive model was developed by using the latest improvements on the selection of best predictor variables. Overall model performance measures were used to select the best discriminant function (DF) models, and also to evaluate their biases and precision. The final predictive model was based on the best five DF models. Each one of these models involved eight environmental variables. Final observed (O), expected (E), and O/E values for the number of macroinvertebrate families (NFAM) and two biotic indices (IBMWP and IASPT) were calculated by averaging their values, previously weighted by the quality of each DF model. Regression analyses among the final O and E values for the calibration dataset showed a high proximity to the ideal theoretical model, where the final E values explained 73-84% of the variation present in the macroinvertebrate communities of the Spanish Mediterranean watercourses. The ANOVA performed among the reference (calibration and validation) and test datasets showed clear differences for the O/E values. Finally, the assessments carried out by the predictive model were sensitive to anthropogenic pressure present in the study area and allowed the definition of five ecological status classes according to the WFD requirements.

We explored the capacity of the biological and hydromorphological indices used in the Water Framework Directive (WFD) to assess ecosystem services by evaluating the ecological status of Spanish River Basins. This analysis relies on an exhaustive bibliography review which showed scientific evidence of the interlinkages between some ecosystem services and different hydromorphological and biological elements which have been used as indices in the WFD. Our findings indicate that, of a total of 38 ecosystem services analyzed, biological and hydromorphological indices can fully evaluate four ecosystem services. In addition, 18 ecosystem services can be partly evaluated by some of the analyzed indices, while 11 are not related with the indices. While Riparian Forest Quality was the index that was able to assess the largest number of ecosystem services (N = 12), the two indices of macrophytes offered very poor guarantees. Finally, biological indices related to diatoms and aquatic invertebrates and the Fluvial Habitat Index can be related with 7, 6, and 6 ecosystem services, respectively. Because the WFD indices currently used in Spain are not able to assess most of the ecosystem services analyzed, we suggest that there is potential to develop the second phase of the WFD implementation taking this approach into consideration. The incorporation of the ecosystem services approach into the WFD could provide the framework for assess the impacts of human activities on the quality of fluvial ecosystems and could give insights for water and watershed management in order to guarantee the delivery of multiple ecosystem services.

Summary 1. According to the guidelines of the European Water Framework Directive, assessment of the ecological quality of streams and rivers should be based on ecotype-specific reference conditions. Here, we assess two approaches for establishing a typology for Mediterranean streams: a top-down approach using environmental variables and bottom-up approach using macroinvertebrate assemblages. 2. Classification of 162 sites using environmental variables resulted in five ecotypes: (i) temporary streams; (ii) evaporite calcareous streams at medium altitude; (iii) siliceous headwater streams at high altitude; (iv) calcareous headwater streams at medium to high altitude and (v) large watercourses. 3. Macroinvertebrate communities of minimally disturbed sites (n = 105), grouped using UPGMA (unweighted pair-group method using arithmetic averages) on Bray–Curtis similarities, were used to validate four of the five ecotypes obtained using environmental variables; ecotype 5, large watercourses, was not included as this group had no reference sites. 4. Analysis of similarities (anosim) showed that macroinvertebrate assemblage composition differed among three of the four ecotypes, resulting in differences between the bottom-up and top-down classification approaches. Siliceous streams were clearly different from the other three ecotypes, evaporite and calcareous ecotypes did not show large differences in macroinvertebrate assemblages and temporary streams formed a very heterogeneous group because of large variability in salinity and hydrology. 5. This study showed that stream classification schemes based on environmental variables need to be validated using biological variables. Furthermore, our findings indicate that special attention should be given to the classification of temporary streams.

During 1994 and 1998–1999, temporal changes in the benthic organic matter (BOM) and its fractions (CBOM = coarse; FBOM = fine; UBOM = ultrafine) were studied in a 354-m reach of Chicamo stream, a saline (9.5 g l−1), temporary semi-arid stream located in southeast Spain. Both time periods differed in their frequency and intensity of spates occurring before sampling. BOM at all sites was dominated by FBOM, followed by UBOM and CBOM. Significant differences in total BOM and its fractions occurred among sampling sites, years, months and dates, illustrating the complex temporal variation of benthic organic matter. Positive correlations were found for FBOM and TBOM with discharge, for CBOM with water depth, and between total BOM, FBOM and UBOM and biomass of periphyton. Principal Component Analysis and Cluster Analysis revealed five different groups, each characterised by different patterns of dominance by fractions of BOM.

We examined the effect of spatial scale on aquatic macroinvertebrate communities in Mediterranean streams from six basins distributed across southern Europe, including Spain, France, Italy, and Greece. We classified the studied streams according to their long-term aquatic regime into the three following types: (i) permanent (P), (ii) intermittent with summer pools (I-P), and (iii) intermittent with summer dry channels (I-D). For each stream type, we analyzed taxonomic and trait diversity, as well as the composition of the macroinvertebrate community, following a spatially nested design at three spatial scales of analysis: microhabitat (substratum patches), mesohabitat (pools vs. riffles), and macrohabitat (streams). In order to assess intrinsic seasonal variability in streams from the Mediterranean region, 20 Surber samples were taken from each stream according to meso- and microhabitat frequency in the wet and the dry season during 2010. Given the need for adaptation to specific hydrological conditions and the fact that microhabitats should encompass the niche requirements of particular taxa, we hypothesized that this spatial scale would have a greater influence on macroinvertebrate taxa composition and biological traits than the other two larger spatial scales in intermittent streams. We observed that patterns in the relative importance of variance components across hierarchical spatial scales changed with time because low flow or droughts altered both mesohabitat prevalence and microhabitat composition. Our results confirm the importance of the microhabitat scale in I-P streams in the wet season but not in the dry one, when a loss of microhabitat diversity occurred. Stream-to-stream variability was more important in P and I-D streams. Our study also explored the relationships between traits and aquatic regimes. We found that aquatic macroinvertebrates inhabiting permanent streams exhibited traits related to the longer duration of life cycles in these rivers (e.g. large size of adult stages) and adaptations to flowing conditions (e.g. modes of aquatic dispersal), whereas aquatic macroinvertebrates inhabiting intermittent streams with summer pools had traits adapted to depositional conditions and ecological preferences for confined habitats (i.e. disconnected pools without flow). Finally, aquatic macroinvertebrates from intermittent streams with summer dry channels had adaptations conferring ability to survive periods with no water (e.g. modes of aerial dispersal, resistant stages). These results indicate that microhabitat conditions should not be neglected as they can play an important role in certain situations. © 2013 E. Schweizerbart'sche Verlagsbuchhandlung.

Type-specific physico-chemical reference conditions are required for the assessment of ecological status in the Water Framework Directive context, similarly to the biological and hydro-morphological elements. This directive emphasises that natural variability of quality elements in high status (reference condition) needs to be quantified. Mediterranean streams often present a marked seasonal pattern in hydrological, biological and geochemical processes which could affect physico-chemical reference conditions. This study establishes general physico-chemical reference conditions (oxygenation, nutrient, salinity and acidification conditions) for different Mediterranean stream types. 116 potential reference sites located in 23 Mediterranean catchments in Spain were sampled in spring, summer and autumn in 2003. All sites were subjected to a screening method for the selection of reference sites in Mediterranean streams (Mediterranean Reference Criteria) and classified using a pre-established stream typology that establishes five different stream types (temporary streams, evaporite-calcareous at medium altitude, siliceous headwaters, calcareous headwaters and large watercourses). Reference conditions (reference value and reference threshold equivalents to high-good class boundary) were calculated using two different methods according to the availability of reference sites: the reference site 75th percentile approach of all reference sites and the 25th percentile of the population approach. The majority of the studied potential reference sites (76 out of 116) were selected as reference sites. Regarding type-specific reference conditions, only siliceous headwaters could be considered different from the rest of stream types because lower conductivity and pH. All reference stream types presented seasonal differences as regards some parameters, except for temporary streams due to the high natural variation of this stream type. For those parameters which presented seasonal differences in a specific stream type, the least restrictive values were proposed as reference conditions.

The European Water Framework Directive establishes the need to define stream type-specific reference conditions to identify “high ecological status”. Methods for selecting reference sites using a priori criteria have been proposed by many authors. A review of these criteria revealed that the most relevant criteria for streams and rivers were those related to riparian vegetation, diffuse and point sources of pollution, river morphology and hydrological conditions and regulation. In this work, we propose 20 criteria that reflect the characteristics of Mediterranean streams and their most frequent disturbances for the selection of reference sites in Mediterranean streams in Spain. We studied 162 sites located in 33 Mediterranean basins belonging to five stream types. Of the locations, 57% were selected as a priori reference sites by having applied the proposed criteria. Reference sites were identified for all stream types except for “large watercourses” which includes the lower reaches of some rivers in this study area. This a priori selection of reference sites was subjected to validation using the macroinvertebrate community by applying of an IBMWP threshold, which is considered to be an indicator of undisturbed sites in Mediterranean streams. This approach determined that whole of this selection (100%) could be considered valid reference sites. Furthermore, we identified differences in the reference conditions for each stream type on the basis of macroinvertebrate assemblage composition.