Climate change is poised to alter mountain stream ecosystem processes via organismal phenological shifts.

Climate change is affecting the phenology of organisms and ecosystem processes across a wide range of environments. However, the links between organismal and ecosystem process change in complex communities remain uncertain. In snow-dominated watersheds, snowmelt in the spring and early summer, followed by a long low-flow period, characterizes the natural flow regime of streams and rivers. Here, we examined how earlier snowmelt will alter the phenology of mountain stream organisms and ecosystem processes via an outdoor mesocosm experiment in stream channels in the Eastern Sierra Nevada, California. The low-flow treatment, simulating a 3- to 6-wk earlier return to summer baseflow conditions projected under climate change scenarios in the region, increased water temperature and reduced biofilm production to respiration ratios by 32%. Additionally, most of the invertebrate species explaining community change (56% and 67% of the benthic and emergent taxa, respectively), changed in phenology as a consequence of the low-flow treatment. Further, emergent flux pulses of the dominant insect group (Chironomidae) almost doubled in magnitude, benefitting a generalist riparian predator. Changes in both invertebrate community structure (composition) and functioning (production) were mostly fine-scale, and response diversity at the community level stabilized seasonally aggregated responses. Our study illustrates how climate change in vulnerable mountain streams at the rain-to-snow transition is poised to alter the dynamics of stream food webs via fine-scale changes in phenology-leading to novel predator-prey "matches" or "mismatches" even when community structure and ecosystem processes appear stable at the annual scale.

Groundwater is a hidden global keystone ecosystem.

Groundwater is a vital ecosystem of the global water cycle, hosting unique biodiversity and providing essential services to societies. Despite being the largest unfrozen freshwater resource, in a period of depletion by extraction and pollution, groundwater environments have been repeatedly overlooked in global biodiversity conservation agendas. Disregarding the importance of groundwater as an ecosystem ignores its critical role in preserving surface biomes. To foster timely global conservation of groundwater, we propose elevating the concept of keystone species into the realm of ecosystems, claiming groundwater as a keystone ecosystem that influences the integrity of many dependent ecosystems. Our global analysis shows that over half of land surface areas (52.6%) has a medium-to-high interaction with groundwater, reaching up to 74.9% when deserts and high mountains are excluded. We postulate that the intrinsic transboundary features of groundwater are critical for shifting perspectives towards more holistic approaches in aquatic ecology and beyond. Furthermore, we propose eight key themes to develop a science-policy integrated groundwater conservation agenda. Given ecosystems above and below the ground intersect at many levels, considering groundwater as an essential component of planetary health is pivotal to reduce biodiversity loss and buffer against climate change.

Climate and land-use changes interact to drive long-term reorganization of riverine fish communities globally.

As climate change unfolds, changes in population dynamics and species distribution ranges are expected to fundamentally reshuffle communities worldwide. Yet, a comprehensive understanding of the mechanisms and extent of community reorganization remains elusive. This is particularly true in riverine systems, which are simultaneously exposed to changing temperature and streamflow, and where land-use change continues to be a major driver of biodiversity loss. Here, we use the most comprehensive compilation of fish abundance time series to date to provide a global synthesis of climate- and LU-induced effects on riverine biota with respect to changes in species thermal and streamflow affinities. We demonstrate that fish communities are increasingly dominated by thermophilic (warm-water) and limnophilic (slow-water) species. Despite being consistent with trends in water temperature and streamflow observed over recent decades, these community changes appear largely decoupled from each other and show wide spatial variation. We further reveal a synergy among climate- and land use-related drivers, such that community thermophilization is heightened in more human-modified systems. Importantly, communities in which species experience thermal and flow regimes that approach or exceed their tolerance thresholds (high community sensitivity), as well as species-poor communities (low community resilience), also display faster rates of compositional change. This research illustrates that quantifying vulnerability of riverine systems to climate change requires a broadening from a narrower thermal focus to more integrative approaches that account for the spatially varying and multifaceted sensitivity of riverine organisms to the interactive effects of water temperature, hydrology, and other anthropogenic changes.

Scale of population synchrony confirms macroecological estimates of minimum viable range size.

Global ecosystems are facing a deepening biodiversity crisis, necessitating robust approaches to quantifying species extinction risk. The lower limit of the macroecological relationship between species range and body size has long been hypothesized as an estimate of the relationship between the minimum viable range size (MVRS) needed for species persistence and the organismal traits that affect space and resource requirements. Here, we perform the first explicit test of this assumption by confronting the MVRS predicted by the range-body size relationship with an independent estimate based on the scale of synchrony in abundance among spatially separated populations of riverine fish. We provide clear evidence of a positive relationship between the scale of synchrony and species body size, and strong support for the MVRS set by the lower limit of the range-body size macroecological relationship. This MVRS may help prioritize first evaluations for unassessed or data-deficient taxa in global conservation assessments.

Four decades of climatic fluctuations and fish recruitment stability across a marine-freshwater gradient.

Investigating the effects of climatic variability on biological diversity, productivity, and stability is key to understanding possible futures for ecosystems under accelerating climate change. A critical question for estuarine ecosystems is, how does climatic variability influence juvenile recruitment of different fish species and life histories that use estuaries as nurseries? Here we examined spatiotemporal abundance trends and environmental responses of 18 fish species that frequently spend the juvenile stage rearing in the San Francisco Estuary, CA, USA. First, we constructed multivariate autoregressive state-space models using age-0 fish abundance, freshwater flow (flow), and sea surface temperature data (SST) collected over four decades. Next, we calculated coefficients of variation (CV) to assess portfolio effects (1) within and among species, life histories (anadromous, marine opportunist, or estuarine dependent), and the whole community; and (2) within and among regions of the estuary. We found that species abundances varied over space and time (increasing, decreasing, or dynamically stable); and in 83% of cases, in response to environmental conditions (wet/dry, cool/warm periods). Anadromous species responded strongly to flow in the upper estuary, marine opportunist species responded to flow and/or SST in the lower estuary, and estuarine dependent species had diverse responses across the estuary. Overall, the whole community when considered across the entire estuary had the lowest CV, and life histories and species provided strong biological insurance to the portfolio (2.4- to 3.5-fold increases in stability, respectively). Spatial insurance also increased stability, although to a lesser extent (up to 1.6-fold increases). Our study advances the notion that fish recruitment stability in estuaries is controlled by biocomplexity-life history diversity and spatiotemporal variation in the environment. However, intensified drought and marine heatwaves may increase the risk of multiple consecutive recruitment failures by synchronizing species dynamics and trajectories via Moran effects, potentially diminishing estuarine nursery function.

The geography of metapopulation synchrony in dendritic river networks.

Dendritic habitats, such as river ecosystems, promote the persistence of species by favouring spatial asynchronous dynamics among branches. Yet, our understanding of how network topology influences metapopulation synchrony in these ecosystems remains limited. Here, we introduce the concept of fluvial synchrogram to formulate and test expectations regarding the geography of metapopulation synchrony across watersheds. By combining theoretical simulations and an extensive fish population time-series dataset across Europe, we provide evidence that fish metapopulations can be buffered against synchronous dynamics as a direct consequence of network connectivity and branching complexity. Synchrony was higher between populations connected by direct water flow and decayed faster with distance over the Euclidean than the watercourse dimension. Likewise, synchrony decayed faster with distance in headwater than mainstem populations of the same basin. As network topology and flow directionality generate fundamental spatial patterns of synchrony in fish metapopulations, empirical synchrograms can aid knowledge advancement and inform conservation strategies in complex habitats.

Drought effects on invertebrate metapopulation dynamics and quasi-extinction risk in an intermittent river network.

Ecological communities can remain stable in the face of disturbance if their constituent species have different resistance and resilience strategies. In turn, local stability scales up regionally if heterogeneous landscapes maintain spatial asynchrony across discrete populations-but not if large-scale stressors synchronize environmental conditions and biological responses. Here, we hypothesized that droughts could drastically decrease the stability of invertebrate metapopulations both by filtering out poorly adapted species locally, and by synchronizing their dynamics across a river network. We tested this hypothesis via multivariate autoregressive state-space (MARSS) models on spatially replicated, long-term data describing aquatic invertebrate communities and hydrological conditions in a set of temperate, lowland streams subject to seasonal and supraseasonal drying events. This quantitative approach allowed us to assess the influence of local (flow magnitude) and network-scale (hydrological connectivity) drivers on invertebrate long-term trajectories, and to simulate near-future responses to a range of drought scenarios. We found that fluctuations in species abundances were heterogeneous across communities and driven by a combination of hydrological and stochastic drivers. Among metapopulations, increasing extent of dry reaches reduced the abundance of functional groups with low resistance or resilience capacities (i.e. low ability to persist in situ or recolonize from elsewhere, respectively). Our simulations revealed that metapopulation quasi-extinction risk for taxa vulnerable to drought increased exponentially as flowing habitats contracted within the river network, whereas the risk for taxa with resistance and resilience traits remained stable. Our results suggest that drought can be a synchronizing agent in riverscapes, potentially leading to regional quasi-extinction of species with lower resistance and resilience abilities. Better recognition of drought-driven synchronization may increase realism in species extinction forecasts as hydroclimatic extremes continue to intensify worldwide.

Evidence for self-organization in determining spatial patterns of stream nutrients, despite primacy of the geomorphic template.

Nutrients in freshwater ecosystems are highly variable in space and time. Nevertheless, the variety of processes contributing to nutrient patchiness, and the wide range of spatial and temporal scales at which these processes operate, obfuscate how this spatial heterogeneity is generated. Here, we describe the spatial structure of stream nutrient concentration, quantify the relative importance of the physical template and biological processes, and detect and evaluate the role of self-organization in driving such patterns. We examined nutrient spatial patterns in Sycamore Creek, an intermittent desert stream in Arizona that experienced an ecosystem regime shift [from a gravel/algae-dominated to a vascular plant-dominated (hereafter, "wetland") system] in 2000 when cattle grazing ceased. We conducted high-resolution nutrient surveys in surface water along a 10-km stream reach over four visits spanning 18 y (1995-2013) that represent different successional stages and prewetland stage vs. postwetland state. As expected, groundwater upwelling had a major influence on nutrient spatial patterns. However, self-organization realized by the mechanism of spatial feedbacks also was significant and intensified over ecosystem succession, as a resource (nitrogen) became increasingly limiting. By late succession, the effects of internal spatial feedbacks and groundwater upwelling were approximately equal in magnitude. Wetland establishment influenced nutrient spatial patterns only indirectly, by modifying the extent of surface water/groundwater exchange. This study illustrates that multiple mechanisms interact in a dynamic way to create spatial heterogeneity in riverine ecosystems, and provides a means to detect spatial self-organization against physical template heterogeneity as a dominant driver of spatial patterns.

Detrimental effects of a novel flow regime on the functional trajectory of an aquatic invertebrate metacommunity.

Novel flow regimes resulting from dam operations and overallocation of freshwater resources are an emerging consequence of global change. Yet, anticipating how freshwater biodiversity will respond to surging flow regime alteration requires overcoming two challenges in environmental flow science: shifting from local to riverscape-level understanding of biodiversity dynamics, and from static to time-varying characterizations of the flow regime. Here, we used time-series methods (wavelets and multivariate autoregressive models) to quantify flow-regime alteration and to link time-varying flow regimes to the dynamics of multiple local communities potentially connected by dispersal (i.e., a metacommunity). We studied the Chattahoochee River below Buford dam (Georgia, U.S.A.), and asked how flow regime alteration by a large hydropower dam may control the long-term functional trajectory of the downstream invertebrate metacommunity. We found that seasonal variation in hydropeaking synchronized temporal fluctuations in trait abundance among the flow-altered sites. Three biological trait states describing adaptation to fast flows benefitted from flow management for hydropower, but did not compensate for declines in 16 "loser" traits. Accordingly, metacommunity-wide functional diversity responded negatively to hydropeaking intensity, and stochastic simulations showed that the risk of functional diversity collapse within the next 4 years would decrease by 17% if hydropeaking was ameliorated, or by 9% if it was applied every other season. Finally, an analysis of 97 reference and 23 dam-affected river sites across the U.S. Southeast suggested that flow variation at extraneous, human-relevant scales (12-hr, 24-hr, 1-week) is relatively common in rivers affected by hydropower dams. This study advances the notion that novel flow regimes are widespread, and simplify the functional structure of riverine communities by filtering out taxa with nonadaptive traits and by spatially synchronizing their dynamics. This is relevant in the light of ongoing and future hydrologic alteration due to climate non-stationarity and the new wave of dams planned globally.

Patterns and drivers of fish extirpations in rivers of the American Southwest and Southeast.

Effective conservation of freshwater biodiversity requires spatially explicit investigations of how dams and hydroclimatic alterations among climate regions may interact to drive species to extinction. We investigated how dams and hydroclimatic alterations interact with species ecological and life history traits to influence past extirpation probabilities of native freshwater fishes in the Upper and Lower Colorado River (CR), Alabama-Coosa-Tallapoosa (ACT), and Apalachicola-Chattahoochee-Flint (ACF) basins. Using long-term discharge data for continuously gaged streams and rivers, we quantified streamflow anomalies (i.e., departure "expected" streamflow) at the sub-basin scale over the past half-century. Next, we related extirpation probabilities of native fishes in both regions to streamflow anomalies, river basin characteristics, species traits, and non-native species richness using binomial logistic regression. Sub-basin extirpations in the Southwest (n = 95 Upper CR, n = 130 Lower CR) were highest in lowland mainstem rivers impacted by large dams and in desert springs. Dampened flow seasonality, increased longevity (i.e., delayed reproduction), and decreased fish egg sizes (i.e., lower parental care) were related to elevated fish extirpation probability in the Southwest. Sub-basin extirpations in the Southeast (ACT n = 46, ACF n = 22) were most prevalent in upland rivers, with flow dependency, greater age and length at maturity, isolation by dams, and greater distance upstream. Our results confirm that dams are an overriding driver of native fish species losses, irrespective of basin-wide differences in native or non-native species richness. Dams and hydrologic alterations interact with species traits to influence community disassembly, and very high extirpation risks in the Southeast are due to interactions between high dam density and species restricted ranges. Given global surges in dam building and retrofitting, increased extirpation risks should be expected unless management strategies that balance flow regulation with ecological outcomes are widely implemented.

Interpreting beta-diversity components over time to conserve metacommunities in highly dynamic ecosystems.

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.

Environmental filtering determines metacommunity structure in wetland microcrustaceans.

Metacommunity approaches are becoming popular when analyzing factors driving species distribution at the regional scale. However, until the popularization of the variation partitioning technique it was difficult to assess the main drivers of the observed patterns (spatial or environmental). Here we propose a new framework linking the emergence of different metacommunity structures (e.g., nested, Gleasonian, Clementsian) to spatial and environmental filters. This is a novel approach that provides a more profound analysis of how both drivers could lead to similar metacommunity structures. We tested this framework on 110 sites covering a strong environmental gradient (i.e., microcrustacean assemblages organized along a salinity gradient, from freshwater to brackish water wetlands). First we identified the metacommunity structure that better fitted these microcrustacean assemblages. Then, we used hierarchical variation partitioning to quantify the relative influences of environmental filters and the distance among wetlands on the identified structure. Our results showed that under strong environmental filtering metacommunity structures were non-random. We also noted that even passive dispersers, that are supposed to be poorly spatially filtered, showed spatial signals at a large geographical scale. However, some difficulties arose when inferring biotic interactions at finer-scale spatial signals. Overall, our study shows the potential of elements of metacommunity structure combined with variation partition techniques to detect environmental drivers and broadscale patterns of metacommunity structure, and that some caution is needed when interpreting finer-scale spatial signals.

Anomalous droughts, not invasion, decrease persistence of native fishes in a desert river.

Changing climate extremes and invasion by non-native species are two of the most prominent threats to native faunas. Predicting the relationships between global change and native faunas requires a quantitative toolkit that effectively links the timing and magnitude of extreme events to variation in species abundances. Here, we examine how discharge anomalies--unexpected floods and droughts--determine covariation in abundance of native and non-native fish species in a highly variable desert river in Arizona. We quantified stochastic variation in discharge using Fourier analyses on >15,000 daily observations. We subsequently coupled maximum annual spectral anomalies with a 15-year time series of fish abundances (1994-2008), using Multivariate Autoregressive State-Space (MARSS) models. Abiotic drivers (discharge anomalies) were paramount in determining long-term fish abundances, whereas biotic drivers (species interactions) played only a secondary role. As predicted, anomalous droughts reduced the abundances of native species, while floods increased them. However, in contrast to previous studies, we observed that the non-native assemblage was surprisingly unresponsive to extreme events. Biological trait analyses showed that functional uniqueness was higher in native than in non-native fishes. We also found that discharge anomalies influenced diversity patterns at the meta-community level, with nestedness increasing after anomalous droughts due to the differential impairment of native species. Overall, our results advance the notion that discharge variation is key in determining community trajectories in the long term, predicting the persistence of native fauna even in the face of invasion. We suggest this variation, rather than biotic interactions, may commonly underlie covariation between native and non-native faunas, especially in highly variable environments. If droughts become increasingly severe due to climate change, and floods increasingly muted due to regulation, fish assemblages in desert rivers may become taxonomically and functionally impoverished and dominated by non-native taxa.

Understanding temporal variability across trophic levels and spatial scales in freshwater ecosystems.

A tenet of ecology is that temporal variability in ecological structure and processes tends to decrease with increasing spatial scales (from locales to regions) and levels of biological organization (from populations to communities). However, patterns in temporal variability across trophic levels and the mechanisms that produce them remain poorly understood. Here we analyzed the abundance time series of spatially structured communities (i.e., metacommunities) spanning basal resources to top predators from 355 freshwater sites across three continents. Specifically, we used a hierarchical partitioning method to disentangle the propagation of temporal variability in abundance across spatial scales and trophic levels. We then used structural equation modeling to determine if the strength and direction of relationships between temporal variability, synchrony, biodiversity, and environmental and spatial settings depended on trophic level and spatial scale. We found that temporal variability in abundance decreased from producers to tertiary consumers but did so mainly at the local scale. Species population synchrony within sites increased with trophic level, whereas synchrony among communities decreased. At the local scale, temporal variability in precipitation and species diversity were associated with population variability (linear partial coefficient, ß = 0.23) and population synchrony (ß = -0.39) similarly across trophic levels, respectively. At the regional scale, community synchrony was not related to climatic or spatial predictors, but the strength of relationships between metacommunity variability and community synchrony decreased systematically from top predators (ß = 0.73) to secondary consumers (ß = 0.54), to primary consumers (ß = 0.30) to producers (ß = 0). Our results suggest that mobile predators may often stabilize metacommunities by buffering variability that originates at the base of food webs. This finding illustrates that the trophic structure of metacommunities, which integrates variation in organismal body size and its correlates, should be considered when investigating ecological stability in natural systems. More broadly, our work advances the notion that temporal stability is an emergent property of ecosystems that may be threatened in complex ways by biodiversity loss and habitat fragmentation.

Nestedness and successional trajectories of macroinvertebrate assemblages in man-made wetlands.

Current successional models, primarily those based on floral succession, propose several distinct trajectories based on the integration of two key hypotheses from succession theory: convergence versus divergence in species composition among successional sites, and progression towards versus deviation from a desired reference state. We applied this framework to faunal succession, including differential colonization between active and passive dispersers, and the nested patterns generated as a consequence of this peculiarity. Nine man-made wetlands located in three different areas, from 0-3 years from wetland creation, were assessed. In addition, 91 wetlands distributed throughout the region were used as references for natural macroinvertebrate communities. We predicted the following: (1) highly nested structures in pioneering assemblages will decrease to lower mid-term values due to a shift from active pioneering taxa to passive disperser ones; (2) passive idiosyncratic taxa will elicit divergent successional trajectories among areas; (3) the divergent trajectories will provoke lower local and higher regional diversity values in the mid-term assemblages than in pioneer assemblages. Our results were largely congruent with hypotheses (1) and (2), diverging from the anticipated patterns only in the case of the temporary wetlands area. However, overall diversity trends based on hypothesis (3) did not follow the expected pattern. The divergent successional trajectories did not compensate for regional biodiversity losses that occurred as a consequence of pioneering colonizer decline over time. Consequently, we suggest reconsidering wetland construction for mitigation purposes within mid-term time frames (≤ 3 years). Wetlands may not offset, within this temporal scenario, regional biodiversity loss because the ecosystem may not support idiosyncratic taxa from natural wetlands.

The color of environmental noise in river networks.

Despite its far-reaching implications for conservation and natural resource management, little is known about the color of environmental noise, or the structure of temporal autocorrelation in random environmental variation, in streams and rivers. Here, we analyze the geography, drivers, and timescale-dependence of noise color in streamflow across the U.S. hydrography, using streamflow time series from 7504 gages. We find that daily and annual flows are dominated by red and white spectra respectively, and spatial variation in noise color is explained by a combination of geographic, hydroclimatic, and anthropogenic variables. Noise color at the daily scale is influenced by stream network position, and land use and water management explain around one third of the spatial variation in noise color irrespective of the timescale considered. Our results highlight the peculiarities of environmental variation regimes in riverine systems, and reveal a strong human fingerprint on the stochastic patterns of streamflow variation in river networks.

Isolation controls reestablishment mechanisms and post-drying community structure in an intermittent stream.

Biota in disturbance-prone landscapes have evolved a variety of strategies to persist long term, either locally (resistance) or by regional recolonization (resilience). Habitat fragmentation and isolation can limit the availability of recolonization pathways, and thus the dynamics of post-disturbance community reestablishment. However, empirical studies on how isolation may control the mechanisms that enable community recovery remain scarce. Here, we studied a pristine intermittent stream (Chalone Creek, Pinnacles National Park, California) to understand how isolation (distance from a perennial pool) alters invertebrate community recolonization after drying. We monitored benthic invertebrate reestablishment during the rewetting phase along a ~2-km gradient of isolation, using mesh traps that selected for specific recolonization pathways (i.e., drift, flying, swimming/crawling, and vertical migration from the hyporheic). We collected daily emigration samples, surveyed the reestablished benthic community after 6 weeks, and compared assemblages across trap types and sites. We found that isolation mediated migration dynamics by delaying peak vertical migration from the hyporheic by ca. 1 day on average per 250 m of dry streambed. The relative importance of reestablishment mechanisms varied longitudinally-with more resistance strategists (up to 99.3% of encountered individuals) in the upstream reaches, and increased drift and aerial dispersers in the more fragmented habitats (up to 17.2% and 18%, respectively). Resistance strategists persisting in the hyporheic dominated overall (88.2% of individuals, ranging 52.9%-99.3% across sites), but notably most of these organisms subsequently outmigrated downstream (85.6% on average, ranging 52.1%-96% across sites). Thus, contrary to conventional wisdom, resistance strategists largely contributed to downstream resilience as well as to local community recovery. Finally, increased isolation was associated with a general decrease in benthic invertebrate diversity, and up to a 3-fold increase in the relative abundance of drought-resistant stoneflies. Our results advance the notion that understanding spatial context is key to predicting post-disturbance community dynamics. Considering the interaction between disturbance and fragmentation may help inform conservation in ecosystems that are subject to novel environmental regimes.

Energy limitation or sensitive predators? Trophic and non-trophic impacts of wastewater pollution on stream food webs.

Impacts of environmental stressors on food webs are often difficult to predict because trophic levels can respond in divergent ways, and biotic interactions may dampen or amplify responses. Here we studied food-web-level impacts of urban wastewater pollution, a widespread source of degradation that can alter stream food webs via top-down and bottom-up processes. Wastewater may (1) subsidize primary producers by decreasing nutrient limitation, inducing a wide-bottomed trophic pyramid. However, (2) wastewater may also reduce the quality and diversity of resources, which could decrease energy transfer efficiency by reducing consumer fitness, leading to predator starvation. Additionally, (3) if higher trophic levels are particularly sensitive to pollution, primary consumers could be released from predation pressure. We tested these hypotheses in 10 pairs of stream sites located upstream and downstream of urban wastewater effluents with different pollutant levels. We found that wastewater pollution reduced predator richness by ∼34%. Community size spectra (CSS) slopes were steeper downstream than upstream of wastewater effluents in all except one impact site where predators became locally extinct. Further, variation in downstream CSS slopes were correlated with pollution loads: the more polluted the stream, the steeper the CSS. We estimate that wastewater pollution decreased energy transfer efficiencies to primary consumers by ∼70%, limiting energy supply to predators. Additionally, traits increasing vulnerability to chemical pollution were overrepresented among predators, which presented compressed trophic niches (δ15 N-δ13 C) downstream of effluents. Our results show that wastewater pollution can impact stream food webs via a combination of energy limitation to consumers and extirpation of pollution-sensitive top predators. Understanding the indirect (biotically mediated) vs. direct (abiotic) mechanisms controlling responses to stress may help anticipating impacts of altered water quantity and quality, key signatures of global change.

River ecosystem conceptual models and non-perennial rivers: A critical review.

Conceptual models underpin river ecosystem research. However, current models focus on continuously flowing rivers and few explicitly address characteristics such as flow cessation and drying. The applicability of existing conceptual models to nonperennial rivers that cease to flow (intermittent rivers and ephemeral streams, IRES) has not been evaluated. We reviewed 18 models, finding that they collectively describe main drivers of biogeochemical and ecological patterns and processes longitudinally (upstream-downstream), laterally (channel-riparian-floodplain), vertically (surface water-groundwater), and temporally across local and landscape scales. However, perennial rivers are longitudinally continuous while IRES are longitudinally discontinuous. Whereas perennial rivers have bidirectional lateral connections between aquatic and terrestrial ecosystems, in IRES, this connection is unidirectional for much of the time, from terrestrial-to-aquatic only. Vertical connectivity between surface and subsurface water occurs bidirectionally and is temporally consistent in perennial rivers. However, in IRES, this exchange is temporally variable, and can become unidirectional during drying or rewetting phases. Finally, drying adds another dimension of flow variation to be considered across temporal and spatial scales in IRES, much as flooding is considered as a temporally and spatially dynamic process in perennial rivers. Here, we focus on ways in which existing models could be modified to accommodate drying as a fundamental process that can alter these patterns and processes across spatial and temporal dimensions in streams. This perspective is needed to support river science and management in our era of rapid global change, including increasing duration, frequency, and occurrence of drying.