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Flowing waters have a unique role in supporting global biodiversity, biogeochemical cycles and human societies(1-5). Although the importance of permanent watercourses is well recognized, the prevalence, value and fate of non-perennial rivers and streams that periodically cease to flow tend to be overlooked, if not ignored(6-8). This oversight contributes to the degradation of the main source of water and livelihood for millions of people(5). Here we predict that water ceases to flow for at least one day per year along 51-60 per cent of the world's rivers by length, demonstrating that non-perennial rivers and streams are the rule rather than the exception on Earth. Leveraging global information on the hydrology, climate, geology and surrounding land cover of the Earth's river network, we show that non-perennial rivers occur within all climates and biomes, and on every continent. Our findings challenge the assumptions underpinning foundational river concepts across scientific disciplines(9). To understand and adequately manage the world's flowing waters, their biodiversity and functional integrity, a paradigm shift is needed towards a new conceptual model of rivers that includes flow intermittence. By mapping the distribution of non-perennial rivers and streams, we provide a stepping-stone towards addressing this grand challenge in freshwater science.

Determining and understanding relationships between biodiversity and hydrology is a critical goal in ecology, particularly given biodiversity in the freshwater realm is in crisis. Despite the prevalence of rivers experiencing natural drying disturbances (which we collectively refer to as intermittent rivers), and projections of increased frequency and duration of drying events, the importance of drying relative to other flow-related determinants of river biodiversity remains understudied. We assessed the influence of drying on alpha- and beta-diversity using discharge and macroinvertebrate data collated from Australia and southwest Europe over broad spatial and temporal scales, providing information on current and past drying events, and combining a wide variety of flow metrics. We found clear evidence that drying acts as a strong environmental filter and is a primary hydrological determinant of alpha-diversity; even when considering both intermittent and perennial rivers, drying-event conditions were its most important predictors. Macroinvertebrate richness declined with increasing durations of drying over the long-term (Australia) and recent (Australia and southwest Europe) history of river discharge, and with decreasing predictability of event timing (Australia). Our analysis also revealed that: responses can be taxon specific due to variation in traits of resistance and resilience to drying; some taxa may respond just as or more strongly to variation in other discharge components (e.g. high- or low-flow events) than to drying; and together these phenomena may result in differing community-level responses within and across regions. Patterns of beta-diversity across the wide biogeographical range of our study suggested that convergent and divergent niche-selection processes may act in combination on aquatic communities of rivers experiencing drying disturbances. However, strong ability to disperse by flight (not by water) weakened beta-diversity patterning among rivers. Our findings can be used to improve understanding of biodiversity organisation in disturbed systems, notably in those with dendritic features, including intermittent rivers.

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.

Headwater streams represent a substantial proportion of river systems and many of them have intermittent flows due to their upstream position in the network. These intermittent rivers and ephemeral streams have recently seen a marked increase in interest, especially to assess the impact of drying on aquatic ecosystems. The objective of this paper is to quantify how discrete (in space and time) field observations of flow intermittence help to extrapolate over time the daily probability of drying (defined at the regional scale). Two empirical models based on linear or logistic regressions have been developed to predict the daily probability of intermittence at the regional scale across France. Explanatory variables were derived from available daily discharge and groundwater-level data of a dense gauging/piezometer network, and models were calibrated using discrete series of field observations of flow intermittence. The robustness of the models was tested using an independent, dense regional dataset of intermittence observations and observations of the year 2017 excluded from the calibration. The resulting models were used to extrapolate the daily regional probability of drying in France: (i) over the period 2011-2017 to identify the regions most affected by flow intermittence; (ii) over the period 1989-2017, using a reduced input dataset, to analyse temporal variability of flow intermittence at the national level. The two empirical regression models performed equally well between 2011 and 2017. The accuracy of predictions depended on the number of continuous gauging/piezometer stations and intermittence observations available to calibrate the regressions. Regions with the highest performance were located in sedimentary plains, --where the monitoring network was dense and --where the regional probability of drying was the highest. Conversely, the worst performances were obtained in mountainous regions. Finally, temporal projections (1989-2016) suggested the highest probabilities of intermittence (> 35 %) in 1989-1991, 2003 and 2005. A high density of intermittence observations improved the information provided by gauging stations and piezometers to extrapolate the temporal variability of intermittent rivers and ephemeral streams.

Community ecology recognizes today that local biological communities are not only affected by local biotic interactions and abiotic environmental conditions, but also by regional processes (e.g. dispersal). While much is known about how metacommunities are organized in space in terrestrial, marine and freshwater ecological systems, their temporal variations remain poorly studied. Here, we address the question of the dynamics of metacommunities in highly variable systems, using intermittent rivers (IRs), those rivers which temporarily stop flowing or dry up, as a model system. We first review how habitat heterogeneity in space and time influences metacommunity organization. Second, we compare the metacommunities in IRs to those in perennial rivers (PRs) and develop the idea that IRs could undergo highly dynamic shifts due to the temporal variability in local and regional community processes. Third, we develop the idea that in IRs, metacommunities of the wet and dry phases of IRs are closely intertwined, thereby increasing even more their respective temporal dynamics. Last, we provide a roadmap to stimulate further conceptual and empirical developments of metacommunity research and identify possible applications for improving the management of IRs and other highly dynamic ecological systems.

1. Intermittent rivers and ephemeral streams (IRES) are watercourses that cease flow at some point in time and space. Arguably Earth's most widespread type of flowing water, IRES are expanding where Anthropocenic climates grow drier and human demands for water escalate. 2. However, IRES have attracted far less research than perennial rivers and are undervalued by society, jeopardizing their restoration or protection. Provision of ecosystem services by IRES is especially poorly understood, hindering their integration into management plans in most countries. 3. We conceptualize how flow intermittence governs ecosystem service provision and transfers at local and river-basin scales during flowing, non-flowing and dry phases. Even when dry or not flowing, IRES perform multiple ecosystem services that complement those of nearby perennial rivers. 4. Synthesis and applications. Conceptualizing how flow intermittence in rivers and streams governs ecosystem services has applied a socio-ecological perspective for validating the ecosystem services of intermittent rivers and ephemeral streams. This can be applied at all flow phases and in assessing impacts of altered flow intermittence on rivers and their ecosystem services in the Anthropocene.

The current erosion of biodiversity is a major concern that threatens the ecological integrity of ecosystems and the ecosystem services they provide. Due to global change, an increasing proportion of river networks are drying and changes from perennial to non-perennial flow regimes represent dramatic ecological shifts with potentially irreversible alterations of community and ecosystem dynamics. However, there is minimal understanding of how biological communities respond functionally to drying. Here, we highlight the taxonomic and functional responses of aquatic macroinvertebrate communities to flow intermittence across river networks from three continents, to test predictions from underlying trait-based conceptual theory. We found a significant breakpoint in the relationship between taxonomic and functional richness, indicating higher functional redundancy at sites with flow intermittence higher than 28%. Multiple strands of evidence, including patterns of alpha and beta diversity and functional group membership, indicated that functional redundancy did not compensate for biodiversity loss associated with increasing intermittence, contrary to received wisdom. A specific set of functional trait modalities, including small body size, short life span and high fecundity, were selected with increasing flow intermittence. These results demonstrate the functional responses of river communities to drying and suggest that ongoing biodiversity reduction due to global change in drying river networks is threatening their functional integrity. These results indicate that such patterns might be common in these ecosystems, even where drying is considered a predictable disturbance. This highlights the need for the conservation of natural drying regimes of intermittent rivers to secure their ecological integrity.

Temporary rivers are increasingly common freshwater ecosystems, but there have been no global syntheses of their community patterns. In this study, we examined the responses of aquatic invertebrate communities to flow intermittence in 14 rivers from multiple biogeographic regions covering a wide range of flow intermittence and spatial arrangements of perennial and temporary reaches. Hydrological data were used to describe flow intermittence (FI, the proportion of the year without surface water) gradients. Linear mixed‐effects models were used to examine the relationships between FI and community structure and composition. We also tested if communities at the most temporary sites were nested subsets of communities at the least temporary and perennial sites. Taxon richness decreased as FI increased and invertebrate communities became dominated by ubiquitous taxa. The number of resilient taxa (with high dispersal capacities) decreased with increased FI, whereas the number of resistant taxa (with adaptations to desiccation) was not related to FI. River‐specific and river‐averaged model comparisons indicated most FI‐community relationships did not differ statistically among rivers. Community nestedness along FI gradients was detected in most rivers and there was little or no influence of the spatial arrangement of perennial and temporary reaches. These results indicate that FI is a primary driver of aquatic communities in temporary rivers, regardless of the biogeographic species pool. Community responses are largely due to resilience rather than resistance mechanisms. However, contrary to our expectations, resilience was not strongly influenced by spatial fragmentation patterns, suggesting that colonist sources other than adjacent perennial reaches were important.

The concept of metacommunity (i.e., a set of local communities linked by dispersal) has gained great popularity among community ecologists. However, metacommunity research mostly addresses questions on spatial patterns of biodiversity at the regional scale, whereas conservation planning requires quantifying temporal variation in those metacommunities and the contributions that individual (local) sites make to regional dynamics. We propose that recent advances in diversity-partitioning methods may allow for a better understanding of metacommunity dynamics and the identification of keystone sites. We used time series of the 2 components of beta diversity (richness and replacement) and the contributions of local sites to these components to examine which sites controlled source-sink dynamics in a highly dynamic model system (an intermittent river). The relative importance of the richness and replacement components of beta diversity fluctuated over time, and sample aggregation led to underestimation of beta diversity by up to 35%. Our literature review revealed that research on intermittent rivers would benefit greatly from examination of beta-diversity components over time. Adequately appraising spatiotemporal variability in community composition and identifying sites that are pivotal for maintaining biodiversity at the landscape scale are key needs for conservation prioritization and planning. Thus, our framework may be used to guide conservation actions in highly dynamic ecosystems when time-series data describing biodiversity across sites connected by dispersal are available.

Drying disturbances are the primary determinant of aquatic community biodiversity in dynamic river ecosystems. Research exploring how communities respond to disturbance has focused on benthic invertebrates in surface sediments, inadequately representing a connected community that extends into the subsurface. We compared subsurface and benthic invertebrate responses to drying, to identify common and context‐dependent spatial patterns. We characterized community composition, alpha diversity and beta diversity across a gradient of drying duration. Subsurface communities responded to drying, but these responses were typically less pronounced than those of benthic communities. Despite compositional changes and in contrast to reductions in benthic alpha diversity, the alpha diversity of subsurface communities remained stable except at long drying durations. Some primarily benthic taxa were among those whose subsurface frequency and abundance responded positively to drying. Collectively, changing composition, stable richness and taxon‐specific increases in occurrence provide evidence that subsurface sediments can support persistence of invertebrate communities during drying disturbances. Beta‐diversity patterns varied and no consistent patterns distinguished the total diversity, turnover or nestedness of subsurface compared to benthic communities. In response to increasing drying duration, beta diversity increased or remained stable for benthic communities, but remained stable or decreased for subsurface communities, likely reflecting contrasts in the influence of mass effects, priority effects and environmental filtering. Dissimilarity between subsurface and benthic communities remained stable or increased with drying duration, suggesting that subsurface communities maintain distinct biodiversity value while also supporting temporary influxes of benthic taxa during drying events. As temporary rivers increase in extent due to global change, we highlight that recognizing the connected communities that extend into the subsurface sediments can enable holistic understanding of ecological responses to drying, the key determinant of biodiversity in these dynamic ecosystems.