Water availability and biological interactions shape amphibian abundance and diversity in Mediterranean temporary rivers.

Amphibians, the most threatened vertebrates globally, face risks due to climate change, habitat loss, and fragmentation. Their sensitivity to environmental changes highlights their importance as ecological indicators. Temporary rivers, influenced by geological, climatic, and anthropogenic factors, play a critical role in shaping biodiversity and community structure. Some species of amphibians may be adapted to these temporary waters, a fact reflected in their life cycles and various biological traits. However, to develop effective conservation strategies for amphibians, it is essential to address the knowledge gaps surrounding the complex interactions between biological dynamics and fluvial habitat conditions. In this study, we investigated how trophic interactions between amphibians and other aquatic organisms (diatoms, macroinvertebrates, and fish), coupled with environmental factors (water availability and riparian structure), can affect amphibian abundance and diversity in temporary rivers. The study was conducted in a Mediterranean river network located in Sant Llorenç del Munt i l'Obac Natural Park (Catalonia, Spain). Our expectations were that habitats suitable for egg deposition, lacking predators (e.g. tadpole-predators and fish), and abundant in food sources would likely support higher amphibian abundance and diversity. However, water availability was identified as a crucial factor shaping abundance and diversity in the studied amphibian communities, even if it correlated with fish presence, and especially impacting amphibian species usually linked to permanent water bodies. Concerning biotic interactions, while our results suggested that amphibian populations in temporary rivers are more dependent on top-down than bottom-up interactions, the presence of aquatic predators was not as conclusive as expected, suggesting that in temporary rivers the fish-avoiding amphibian species can survive using microhabitats or breeding opportunities linked to natural river dynamics. Overall, our findings highlight the importance of considering multi-trophic interactions, hydroperiod and habitat heterogeneity in temporary river ecosystems for effective amphibian conservation.

Unravelling large-scale patterns and drivers of biodiversity in dry rivers.

More than half of the world's rivers dry up periodically, but our understanding of the biological communities in dry riverbeds remains limited. Specifically, the roles of dispersal, environmental filtering and biotic interactions in driving biodiversity in dry rivers are poorly understood. Here, we conduct a large-scale coordinated survey of patterns and drivers of biodiversity in dry riverbeds. We focus on eight major taxa, including microorganisms, invertebrates and plants: Algae, Archaea, Bacteria, Fungi, Protozoa, Arthropods, Nematodes and Streptophyta. We use environmental DNA metabarcoding to assess biodiversity in dry sediments collected over a 1-year period from 84 non-perennial rivers across 19 countries on four continents. Both direct factors, such as nutrient and carbon availability, and indirect factors such as climate influence the local biodiversity of most taxa. Limited resource availability and prolonged dry phases favor oligotrophic microbial taxa. Co-variation among taxa, particularly Bacteria, Fungi, Algae and Protozoa, explain more spatial variation in community composition than dispersal or environmental gradients. This finding suggests that biotic interactions or unmeasured ecological and evolutionary factors may strongly influence communities during dry phases, altering biodiversity responses to global changes.

Evaluating the response of current biotic indices and functional metrics to natural and anthropogenic predictors in disconnected pools of temporary rivers.

Temporary rivers, forming the majority of river networks worldwide, are key biodiversity hotspots. Despite their great value for maintaining biodiversity and ecosystem functioning, they are often neglected in biomonitoring programs due to several challenges, such as their variable hydromorphology and the difficulty of establishing reference conditions given their dynamic nature, resulting in highly variable communities. Disconnected pools often form in temporary rivers when flow ceases, providing refuge for aquatic taxa. Given their importance for biodiversity conservation, revising and adapting biotic indices are needed. Here, we evaluate the performance of current biological indices designed for perennial rivers (macroinvertebrates, diatoms) and functional metrics (macroinvertebrates) in assessing biological quality of disconnected pools. We sampled 55 disconnected pools in Catalonia, NE Spain, covering local (e.g., physico-chemical variables, water chemistry) and regional (e.g., human influence, hydrological variables at the water body level) natural and anthropogenic gradients. Only a few macroinvertebrate biotic indices (e.g., family richness, EPT/EPT + OCH and OCH) showed strong responses to anthropogenic predictors and were unaffected by natural predictors at both local and regional scales, making them suitable for biomonitoring. Of the newly adopted functional metrics of macroinvertebrate communities tested, only two (i.e., functional redundancy of predators and response diversity based on the total community) responded strongly to anthropogenic predictors. The rest showed varying responses to the interactive effect of anthropogenic and natural predictors, requiring calibration efforts. Models assessing these metrics explained <40 % of the total variation, likely due to the interplay of colonization/extinction dynamics and density-dependent trophic interactions governing community assemblages in disconnected pools. Although some existing biological metrics could potentially be used to monitor the ecological status of disconnected pools, we call for further development of biomonitoring tools specifically designed for these habitats since they will become more widespread with global change.

Non-perennial segments in river networks.

Non-perennial river segments - those that recurrently cease to flow or frequently dry - occur in all river networks and are globally more abundant than perennial (always flowing) segments. However, research and management have historically focused on perennial river segments. In this Review, we outline how non-perennial segments are integral parts of river networks. Repeated cycles of flowing, non-flowing and dry phases in non-perennial segments influence biodiversity and ecosystem dynamics at different spatial scales, from individual segments to entire river networks. Varying configurations of perennial and non-perennial segments govern physical, chemical and ecological responses to changes in the flow regimes of each river network, especially in response to human activities. The extent of non-perennial segments in river networks has increased owing to warming, changing hydrological patterns and human activities, and this increase is predicted to continue. Moreover, the dry phases of flow regimes are expected to be longer, drier and more frequent, albeit with high regional variability. These changes will likely impact biodiversity, potentially tipping some ecosystems to compromised stable states. Effective river-network management must recognize ecosystem services (such as flood risk management and groundwater recharge) provided by non-perennial segments and ensure their legislative and regulatory protection, which is often lacking.

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.

Spatial priorities for freshwater biodiversity conservation in light of catchment protection and connectivity in Europe.

Freshwater ecosystems host disproportionately high numbers of species relative to their surface area yet are poorly protected globally. We used data on the distribution of 1631 species of aquatic plant, mollusc, odonate and fish in 18,816 river and lake catchments in Europe to establish spatial conservation priorities based on the occurrence of threatened, range-restricted and endemic species using the Marxan systematic conservation planning tool. We found that priorities were highest for rivers and ancient lakes in S Europe, large rivers and lakes in E and N Europe, smaller lakes in NW Europe and karst/limestone areas in the Balkans, S France and central Europe. The a priori inclusion of well-protected catchments resulted in geographically more balanced priorities and better coverage of threatened (critically endangered, endangered and vulnerable) species. The a priori exclusion of well-protected catchments showed that priority areas that need further conservation interventions are in S and E Europe. We developed three ways to evaluate the correspondence between conservation priority and current protection by assessing whether a cathment has more (or less) priority given its protection level relative to all other catchments. Each method found that priority relative to protection was high in S and E Europe and generally low in NW Europe. The inclusion of hydrological connectivity had little influence on these patterns but decreased the coverage of threatened species, indicating a trade-off between connectivity and conservation of threatened species. Our results suggest that catchments in S and E Europe need urgent conservation attention (protected areas, restoration, management, species protection) in the face of imminent threats such as river regulation, dam construction, hydropower development and climate change. Our study presents continental-scale conservation priorities for freshwater ecosystems in ecologically meaningful planning units and will thus be important in freshwater biodiversity conservation policy and practice, and water management in Europe.

Taxon-specific sensitivities to flow intermittence reveal macroinvertebrates as potential bioindicators of intermittent rivers and streams.

As complex mosaics of lotic, lentic, and terrestrial habitats, intermittent rivers and ephemeral streams (IRES) support high biodiversity. Despite their ecological importance, IRES are poorly represented in routine monitoring programs, but recent recognition of their considerable-and increasing-spatiotemporal extent is motivating efforts to better represent IRES in ecological status assessments. We examine response patterns of aquatic macroinvertebrate communities and taxa to flow intermittence (FI) across three European climatic regions. We used self-organizing map (SOM) to ordinate and classify sampling sites based on community structure in regions with continental, Mediterranean and oceanic climates. The SOM passively introduced FI, quantified as the mean annual % flow, and visualized its variability across classified communities, revealing a clear association between community structure and FI in all regions. Indicator species analysis identified taxa indicative of low, intermediate and high FI. In the continental region, the amphipod Niphargus was indicative of high FI and was associated with groundwater-fed IRES, whereas indicators of Mediterranean IRES comprised Odonata, Coleoptera and Heteroptera taxa, which favor lentic conditions. In the oceanic region, taxa indicative of relatively high FI included leuctrid stoneflies and a limnephilid caddisfly, likely reflecting the colonization of IRES by aerial adults from nearby perennial reaches. The Diptera families Chironomidae and Simuliidae showed contrasting FI preferences among regions, reflecting environmental heterogeneity between regions and the coarse taxonomic resolution to which these organisms were identified. These region-specific community and taxon responses of aquatic biota to FI highlight the need to adapt standard biotic indices to enable effective ecological status assessments in IRES.

DISPERSE, a trait database to assess the dispersal potential of European aquatic macroinvertebrates.

Dispersal is an essential process in population and community dynamics, but is difficult to measure in the field. In freshwater ecosystems, information on biological traits related to organisms' morphology, life history and behaviour provides useful dispersal proxies, but information remains scattered or unpublished for many taxa. We compiled information on multiple dispersal-related biological traits of European aquatic macroinvertebrates in a unique resource, the DISPERSE database. DISPERSE includes nine dispersal-related traits subdivided into 39 trait categories for 480 taxa, including Annelida, Mollusca, Platyhelminthes, and Arthropoda such as Crustacea and Insecta, generally at the genus level. Information within DISPERSE can be used to address fundamental research questions in metapopulation ecology, metacommunity ecology, macroecology and evolutionary ecology. Information on dispersal proxies can be applied to improve predictions of ecological responses to global change, and to inform improvements to biomonitoring, conservation and management strategies. The diverse sources used in DISPERSE complement existing trait databases by providing new information on dispersal traits, most of which would not otherwise be accessible to the scientific community.

A Metacommunity Approach to Improve Biological Assessments in Highly Dynamic Freshwater Ecosystems.

Rapid shifts in biotic communities due to environmental variability challenge the detection of anthropogenic impacts by current biomonitoring programs. Metacommunity ecology has the potential to inform such programs, because it combines dispersal processes with niche-based approaches and recognizes variability in community composition. Using intermittent rivers-prevalent and highly dynamic ecosystems that sometimes dry-we develop a conceptual model to illustrate how dispersal limitation and flow intermittence influence the performance of biological indices. We produce a methodological framework integrating physical- and organismal-based dispersal measurements into predictive modeling, to inform development of dynamic ecological quality assessments. Such metacommunity-based approaches could be extended to other ecosystems and are required to underpin our capacity to monitor and protect ecosystems threatened under future environmental changes.

Simulating rewetting events in intermittent rivers and ephemeral streams: A global analysis of leached nutrients and organic matter.

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.

Biomonitoring of intermittent rivers and ephemeral streams in Europe: Current practice and priorities to enhance ecological status assessments.

Intermittent rivers and ephemeral streams (IRES) are common across Europe and dominate some Mediterranean river networks. In all climate zones, IRES support high biodiversity and provide ecosystem services. As dynamic ecosystems that transition between flowing, pool, and dry states, IRES are typically poorly represented in biomonitoring programmes implemented to characterize EU Water Framework Directive ecological status. We report the results of a survey completed by representatives from 20 European countries to identify current challenges to IRES status assessment, examples of best practice, and priorities for future research. We identify five major barriers to effective ecological status classification in IRES: 1. the exclusion of IRES from Water Framework Directive biomonitoring based on their small catchment size; 2. the lack of river typologies that distinguish between contrasting IRES; 3. difficulties in defining the 'reference conditions' that represent unimpacted dynamic ecosystems; 4. classification of IRES ecological status based on lotic communities sampled using methods developed for perennial rivers; and 5. a reliance on taxonomic characterization of local communities. Despite these challenges, we recognize examples of innovative practice that can inform modification of current biomonitoring activity to promote effective IRES status classification. Priorities for future research include reconceptualization of the reference condition approach to accommodate spatiotemporal fluctuations in community composition, and modification of indices of ecosystem health to recognize both taxon-specific sensitivities to intermittence and dispersal abilities, within a landscape context.

TREHS: An open-access software tool for investigating and evaluating temporary river regimes as a first step for their ecological status assessment.

When the regime of a river is not perennial, there are four main difficulties with the use of hydrographs for assessing hydrological alteration: i) the main hydrological features relevant for biological communities are not quantitative (discharges) but qualitative (phases such as flowing water, stagnant pools or lack of surface water), ii) stream flow records do not inform on the temporal occurrence of stagnant pools, iii) as most of the temporary streams are ungauged, their regime has to be evaluated by alternative methods such as remote sensing or citizen science, and iv) the biological quality assessment of the ecological status of a temporary stream must follow a sampling schedule and references adapted to the flow- pool-dry regime. To overcome these challenges within an operational approach, the freely available software tool TREHS has been developed within the EU LIFE TRIVERS project. This software permits the input of information from flow simulations obtained with any rainfall-runoff model (to set an unimpacted reference stream regime) and compares this with the information obtained from flow gauging records (if available) and interviews with local people, as well as instantaneous observations by individuals and interpretation of ground-level or aerial photographs. Up to six metrics defining the permanence of water flow, the presence of stagnant pools and their temporal patterns of occurrence are used to determine natural and observed river regimes and to assess the degree of hydrological alteration. A new regime classification specifically designed for temporary rivers was developed using the metrics that measure the relative permanence of the three main phases: flow, disconnected pools and dry stream bed. Finally, the software characterizes the differences between the natural and actual regimes, diagnoses the hydrological status (degree of hydrological alteration), assesses the significance and robustness of the diagnosis and recommends the best periods for biological quality samplings.

Long-term changes in a population of an invasive bivalve and its effects.

Although the ecological and economic effects of non-native species probably often change through time, few studies have documented such effects. The zebra mussel (Dreissena polymorpha) is an important invader that has had large ecological and economic effects on the ecosystems it has invaded in North America and western Europe. Our 20-year study of the Hudson River, New York, showed that the characteristics of a zebra mussel population and its effects on other benthic animals both changed substantially through time. Over the period of study, annual survivorship of adult zebra mussels fell >100-fold, which caused the aggregate filtration rate of the population to fall by 82%. Population size and body size of zebra mussels may also have fallen. In the early years of the invasion, densities of nearly all benthic animals in deepwater sites fell steeply (by 80-99%). After about 8 years of decline, these populations began to recover, and are approaching pre-invasion densities. The littoral zoobenthos showed neither the initial decline nor the subsequent recovery. Although the mechanisms behind these changes are not fully clear, our study shows that the effects of an invader may change considerably over time.

Patterns of metal bioaccumulation in two filter-feeding macroinvertebrates: exposure distribution, inter-species differences and variability across developmental stages.

This study focused on the metal bioaccumulation of two aquatic insects (Ephoron virgo and Hydropsyche spp.) in order to evaluate the spatial distribution of metals, the interspecific differences between both filter-feeders and the bioaccumulation dynamics during E. virgo development stages. Hg, Cd, Ni, Cr, As, Pb, Cu, Ti, Zn and Mn were quantified in insects and in suspended particulate matter (SPM) sampled downstream and upstream of a chemical plant, where more than 300,000t of polluted sediments are deposited. Hg concentrations were one order of magnitude higher downstream of the sediment dump, which showed that the Hg pollution originated in the chemical plant. Cd, Ni, Cr, Pb, Ti, Zn and Mn in invertebrates revealed that metal pollution was present upstream in other parts of the river. Interspecific differences were observed for all metals but Mn; significantly higher concentrations were observed in E. virgo over Hydropsyche exocellata, except for Cd, which showed 10-fold higher values. Hg and Cd increased until E. virgo nymphs reached 11 mm and decreased afterwards in late instars when nymphs were about to emerge. Cr, Pb, Ti and Mn decreased along early instars followed by a steady state in late instars. Similar values were obtained for Cu, As and Zn along all instars. Sexual differences between males and females of E. virgo were observed for Cd, Cu and Mn. Hg and Cd persistence was strong across developmental stages since high concentrations were found in eggs and emerging adults. Because the behavior of different metals varied for the two species and during the developmental stages of E. virgo, care should be taken in the interpretation of insect metal concentrations when analyzing the food chain transfer of metals in river ecosystems.