Climatic aridity increases temporal nestedness of invertebrate communities in naturally drying rivers

Climate change is altering the water cycle globally, increasing the frequency and magnitude of floods and droughts. An outstanding question is whether biodiversity responses to hydrological disturbance depend on background climatic context – and if so, which contexts increase vulnerability to disturbance. Answering this question requires comparison of organismal responses across environmental gradients. However, opportunities to track disturbed communities against an undisturbed baseline remain rare. Here we gathered a global dataset capturing responses of aquatic invertebrate communities to river drying, which includes 112 sites spanning a gradient of climatic aridity. We measured the effects of river drying on taxonomic richness and temporal β‐diversity (turnover and nestedness components). We also measured the relative abundance of aquatic invertebrates with strategies that confer resilience (or resistance) to drying. Contrary to our expectations, we found that taxonomic richness recovered from drying similarly across the aridity gradient. The turnover component of β‐diversity (i.e. species replacements over time) largely accounted for differences in community composition before versus after drying. However, increasing aridity was associated with greater nestedness‐driven compositional changes at intermittent sites – that is, after drying communities became subsets of those before drying. These results show that climatic context can explain variation in community responses to the same hydrological disturbance (drying), and suggest that increased aridity will constrain biodiversity responses at regional scales. Further consideration of the climatic context in hydroecological research may help improve predictions of the local impacts of hydrological disturbance by identifying climate regions where communities are more (or less) sensitive to extremes, including river drying events

Protecting U.S. temporary waterways

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Direct observations of the effect of fine sediment deposition on the vertical movement of Gammarus pulex (Amphipoda: Gammaridae) during substratum drying

Benthic macroinvertebrates inhabit the streambed sediments of temporary streams during drying events. Fine sediment (< 2 mm in diameter) deposition and clogging of interstitial pathways reduces the connectivity between benthic and subsurface habitats, potentially inhibiting macroinvertebrate vertical movements. Direct observations within subsurface sediments are, however, inherently difficult. As a result, confirmation of macroinvertebrate vertical movement, and the effect of fine sediment, is limited. We used laboratory mesocosms containing transparent gravel sized particles (10–15 mm) to facilitate the direct observation and tracking of vertical movements by Gammarus pulex in response to water level reduction and sedimentation. Seven sediment treatments comprised two fine sediment fractions (small: 0.125–0.5 mm, coarse sand: 0.5–1 mm) deposited onto the surface of the substrate, and a control treatment where no fine sediment was applied. We found that G. pulex moved into the subsurface gravel sediments in response to drying, but their ability to remain submerged during water level reduction was impeded by fine sediment deposition. In particular deposition of the coarser sand fraction clogged the sediment surface, limiting vertical movements. Our results highlight the potential effect of sedimentation on G. pulex resistance to drying events in streams

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 ex‐ tent 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 (precon‐ ditioning), 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 experi‐ mentally simulated, under standard laboratory conditions, rewetting of leaves, river‐ bed sediments, and epilithic biofilms collected during the dry phase across 205 IRES from five major climate zones. We determined the amounts and qualitative character‐ istics 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 dis‐ solved substances during rewetting events (56%–98%), and that flux rates distinctly differ among climate zones. Dissolved organic carbon, phenolics, and nitrate contrib‐ uted 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 vari‐ ables expected to be modified under climate change (i.e. potential evapotranspiration, aridity, dry period duration, land use) were correlated with the amount of leached sub‐ stances, 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 event

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 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 organic matter. 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

Biological indices to characterize community responses to drying in streams with contrasting flow permanence regimes

Many river networks include temporary reaches that stop flowing and may dry during unpredictable droughts (near-perennial) or more frequently (intermittent). A few biological indices have been developed to assess invertebrate community responses to hydrological variability, including the instream conditions associated with drought, but their performance in temporary streams remains poorly known. We evaluated the ability of two such indices, the Lotic-invertebrate Index for Flow Evaluation (LIFE) and the Drought Effect of Habitat Loss on Invertebrates (DEHLI), to predict responses to flow cessation and drying in temporary streams with contrasting flow permanence regimes. We used a 26-year dataset comprising spring-season invertebrate community samples and daily discharge measurements from 46 sites in a cool, wet temperate region, to examine relationships between hydrological variables and changes in index scores. We also identified taxon-specific thresholds at which occurrence changed with increasing drying and flowing durations. Both indices effectively characterized responses to increasing no-flow durations. DEHLI also reflected community changes following flow resumptions, identified differences in responses among flow permanence groups, and was particularly able to predict community responses at near-perennial sites. DEHLI scores at near-perennial sites took on average three years after a drying event to return to values typical of perennial sites, whereas responses to increasing flow duration were more erratic at intermittent sites. Lotic specialists declined whereas lentic and semi-aquatic taxa increased in occurrence with no-flow duration after summers with <50 days without flow, due to changes in the availability of preferred habitat types. Community responses to drying events were less predictable among intermittent than near-perennial sites, likely because differences in habitat conditions and connectivity may lead intermittent communities to harbour contrasting pools of species with strategies that promote persistence during and/or recolonization after drying. We identify DEHLI as an index that can characterize community responses to drying in temporary streams with contrasting flow permanence regimes. We also recommend the development of new indices that include lentic, semi-aquatic and terrestrial as well as lotic taxa, to more comprehensively describe and predict community responses to changing instream conditions

Sediment respiration pulses in intermittent rivers and ephemeral streams

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

A global analysis of terrestrial plant litter dynamics in non-perennial waterways

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

Les communautés d'invertébrés à lit de gravier, rivières tressés sont très résistants à l'écoulement intermittence

International audienceIn naturally disturbed systems, harsh environmental conditions act as filters on the regional species pool, restricting the number of taxa able to form a local community to those with traits promoting resistance or resilience. Thus, communities in highly disturbed ecosystems may be less sensitive to a given disturbance than those in less disturbed ecosystems. We explored this idea by examining the response of aquatic invertebrate communities to flow intermittence in gravel-bed, braided rivers (BRs). Flow intermittence is considered a major driver of communities in rivers, but its influence on communities in BRs, which are recognized as naturally highly disturbed environments, is relatively unexplored. We used a multisite Before-After–Control- Impact (BACI) design to quantify the effects of drying events of different durations (moderate: 2–3 wk, severe:1–3 mo) on invertebrate communities in 8 BRs in southeastern France. As predicted, no effects of flow intermittence were detected 1 to 4 mo after flow resumption on taxonomic richness, composition, or functional diversity of communities facing moderate drying events. Communities subjected to severe drying events were similar to those in perennial reaches as few as 19 d after flow resumption. Moreover, communities showed functional redundancy and no loss of functional diversity after drying events. These results differ from those of studies in other river systems, where persistent effects of flow intermittence on communities generally have been found, and highlight the need for cross-system comparisons that explore the effects of drying on communities. Identifying the processes (e.g., niche selection, cotolerance) and habitat features (e.g., hyporheic zone refugia) that promote community resilience in BRs will advance our understanding of how anthropogenic stressors and climate change may affect communities in freshwater ecosystems

Gammarus pulex (Crustacea: Amphipoda) avoids increasing water temperature and intraspecific competition through vertical migration into the hyporheic zone: a mesocosm experiment

The saturated interstices below and adjacent to the riverbed (i.e., the hyporheic zone) can be a refuge for biota during low flows, flow cessation and river drying. Prior to complete drying, organisms are constrained by abiotic and biotic factors (e.g., water temperature, com- petition) and may respond through vertical migration into the hyporheic zone. However, it remains unclear when temperature and competition become harsh enough to trigger migration. Furthermore, potential consequences of using the hyporheic zone, which is often food-limited, on the survival, effects on ecosystem function and physiology of organisms are unknown. We tested the hypotheses that (1) Gammarus pulex, a widespread detritivore, migrates into the hyporheic zone to avoid increasing surface water temperature and intraspecific competition and (2) that these factors would reduce their survival, leaf mass consumption and energy stores. Using 36 mesocosms, three temperature (15, 20, 25 °C) and species density levels (low, medium, high) were manipulated in a factorial design over 15 days. Increasing temperature to 25 °C and a threefold increase in density both caused G. pulex to vertically migrate, and the interaction of these factors was additive, rather than antagonistic or synergistic. Importantly, survival, leaf consumption and glycogen content were reduced in high temperature and density treatments, suggesting tradeoffs between tolerating harsh surface conditions and limitations of inhabiting the hyporheic zone. Identifying that the hyporheic zone is used by G. pulex to avoid high water temperature and intraspecific competition is a key finding considering the global-scale increases in temperature and flow intermittence