Hyporheic invertebrates as bioindicators of ecological health in temporary rivers: A meta-analysis

Worldwide, many rivers cease flow and dry either naturally or owing to human activities such as water extraction. However, even when surface water is absent, diverse assemblages of aquatic invertebrates inhabit the saturated sediments below the river bed (hyporheic zone). In the absence of surface water or flow, biota of this zone may be sampled as an alternative to surface water-based ecological assessments. The potential of hyporheic invertebrates as ecological indicators of river health, however, is largely unexplored. We analysed hyporheic taxa lists from the international literature on temporary rivers to assess compositional similarity among broad-scale regions and sampling conditions, including the presence or absence of surface waters and flow, and the regional effect of hydrological phase (dry channel, non-flowing waters, surface flow) on richness. We hypothesised that if consistent patterns were found, then effects of human disturbances in temporary rivers may be assessable using hyporheic bioindicators. Assemblages differed geographically and by climate, but hydrological phase did not have a strong effect at the global scale. However, hyporheic assemblage composition within regions varied along a gradient of higher richness during wetter phases. This indicates that within geographic regions, hyporheic responses to surface drying are predictable and, by extension, hyporheic invertebrates are potentially useful ecological indicators of temporary river health. With many rivers now experiencing, or predicted to experience, lower flows and longer dry phases owing to climate change, the development of ecological assessment methods specific to flow intermittency is a priority. We advocate expanded monitoring of hyporheic zones in temporary rivers and recommend hyporheic invertebrates as potential bioindicators to complement surface water assessments.     

Oligochaete Assemblages in the Hyporheic Zone and Coarse Surface Sediments: Their Importance for Understanding of Ecological Functioning of Watercourses

The hyporheic zone and its interactions with coarse surface sediments is increasingly reported by aquatic ecologists because the water exchanges between surface and subsurface are important factors for the understanding of the ecosystem functioning. However, the hyproheic oligochaete assemblages have received less attention than other assemblages such as crustaceans. In addition, studies investigating the incidence of pollution in watercourses have mostly focused on the benthic zone and have neglected the hyporheic zone. Some examples are given from an unpolluted glacial river (Roseg), polluted plains rivers (Moselle, Rhône) and a protected wetland in an urbanized environment. The hyporheic zone kept the memory of past and present incidences of pollution, in particular when downwellings of polluted surface waters to the hyporheic zone predominated. The Active hydrologic Exchange Describers between surface and subsurface (AED oligochaete species) were the same in the glacial river Roseg, the rivers Rhône and Moselle and the urbanized wetland. The predominance of pollution-tolerant species like Limnodrilus hoffmeisteri was observed in polluted groundwater as well as in polluted surface coarse sediments. Moreover, the urbanized wetland exhibited a high species richness, suggesting that the hyporheic zone is a reservoir of species. The oligochaete communities enable biologists to simultaneously assess the pollution incidence, the permeability of coarse habitats, the water exchanges between surface and subsurface, and give an approximate measure of the metabolic activities in the sediments. Consequently, the simultaneous study of surface and hyporheic oligochaete assemblages is of great interest when considering the ecological functioning of watercourses and the incidence of pollution inputs.     

The ecological significance of exchange processes between rivers and groundwater

1. This review focuses on the connectivity between river and groundwater ecosystems, viewing them as linked components of a hydrological continuum. Ecological processes that maintain the integrity of both systems and those that are mediated by their ecotones are evaluated. 2. The hyporheic zone, as the connecting ecotone, shows diverse gradients. Thus it can be characterized by hydrological, chemical, zoological and metabolic criteria. However, the characteristics of the hyporheic zone tend to vary widely in space and time as well as from system to system. The exact limits are difficult to designate and the construction of static concepts is inadequate for the representation of ecological processes. The hyporheic interstices are functionally a part of both the fluvial and groundwater ecosystems. 3. The permeability of the ecotone depends on the hydraulic conductivity of the sediment layers which, because of their heterogeneity, form many flowpath connections between the stream and the catchment, from the small scale of a single microhabitat to the large scale of an entire alluvial aquifer. Local up- and downwellings are determined by geomorphologic features such as streambed topography, whereas large-scale exchange processes are determined mainly by the geological properties of the catchment. Colmation—clogging of the top layer of the channel sediments—includes all processes leading to a reduction of pore volume, consolidation of the sediment matrix, and decreased permeability of the stream bed. Consequently, colmation can hinder exchange processes between surface water and groundwater. 4. Physicochemical gradients in the interstices result from several processes: (i) hyporheic flow pattern and the different properties of surface and groundwaters; (ii) retention, caused by the filtering effect of pore size and lithologic sorption as well as the transient storage of solutes caused by diminished water velocities; (iii) biogeochemical transformations in conjunction with local residence time. Each physicochemical parameter may develop its own vertical dynamics laterally from the active channel into the banks as well as longitudinally because of geomorphologic changes. 5. The river–groundwater interface can act as a source or sink for dissolved organic matter, depending on the volume and direction of flow, dissolved organic carbon concentrations and biotic activity. Interstitial storage of particulate organic matter is influenced mainly by grain size distribution and by spates involving bedload movement that may import or release matter, depending on the season. After initial transient and abiotic storage, hyporheic organic matter is mobilized and transformed by the biota. Micro-organisms account for over 90% of the community respiration. In subterranean waters most bacteria are attached to surfaces and remain in a biofilm. 6. Hyporheic interstices are functionally significant for phreatic and riverine metazoans because they act as a refuge against adverse conditions. The net flow direction exerts a dominant influence on interstitial colonization, but many other factors also seem to be important in structuring the hyporheos. 7. The hyporheic corridor concept emphasizes connectivity and interactions between subterranean and surface flow on an ecosystem level for floodplain rivers. It is a complementary concept to others which focus on surficial processes in the lateral and longitudinal dimensions. 8. The ecological integrity of groundwater and fluvial systems is often threatened by human activities: (i) by reducing connectivity; (ii) by altering exchange processes; and (iii) by toxic or organic contamination.     

Restoring freshwater ecosystems in riverine landscapes: the roles of connectivity and recovery processes

1. This paper introduces key messages from a number of papers emanating from the Second International Symposium on Riverine Landscapes held in August 2004 in Sweden, focusing on river restoration. Together these papers provide an overview of the science of river restoration, and point out future research needs. 2. Restoration tests the feasibility of recreating complex ecosystems from more simple and degraded states, thereby presenting a major challenge to ecological science. Therefore, close cooperation between practitioners and scientists would be beneficial, but most river restoration projects are currently performed with little or no scientific involvement. 3. Key messages emanating from this series of papers are: The scope, i.e. the maximum and minimum spatial extent and temporal duration of habitat use, of species targeted for restoration should be acknowledged, so that all relevant stages in their life cycles are considered. Species that have been lost from a stream cannot be assumed to recolonise spontaneously, calling for strategies to ensure the return of target species to be integrated into projects. Possible effects of invasive exotic species also need to be incorporated into project plans, either to minimise the impact of exotics, or to modify the expected outcome of restoration in cases where extirpation of exotics is impractical. 4. Restoration of important ecological processes often implies improving connectivity of the stream. For example, longitudinal and lateral connectivity can be enhanced by restoring fluvial dynamics on flood‐suppressed rivers and by increasing water availability in rivers subject to water diversion or withdrawal, thereby increasing habitat and species diversity. Restoring links between surface and ground water flow enhances vertical connectivity and communities associated with the hyporheic zone. 5. Future restoration schemes should consider where in the catchment to locate projects to make restoration most effective, consider the cumulative effects of many small projects, and evaluate the potential to restore ecosystem processes under highly constrained conditions such as in urban areas. Moreover, restoration projects should be properly monitored to assess whether restoration has been successful, thus enabling adaptive management and learning for the future from both successful and unsuccessful restorations.     

Longitudinal patterns of invertebrates in the hyporheic zone of a glacial river

1. Longitudinal changes in physicochemical factors and the composition of the invertebrate community were examined in the hyporheic zone of a glacial river (Val Roseg, Switzerland) over a distance of 11 km from the glacier terminus. Multivariate analysis was used to determine the habitat preferences of taxa along an upstream‐downstream gradient of increasing temperature and groundwater contribution to river flow. 2. The hyporheos conformed to the longitudinal distribution model described for zoobenthic communities of glacial rivers in that taxonomic richness increased with distance from the glacier terminus. Spatial variation in taxonomic richness was best explained by temperature, the influence of groundwater, and the amount of organic matter. The overriding importance of these variables on the distribution of taxa was confirmed by the multivariate analysis. 3. The hyporheic zone contributed significantly to the overall biodiversity of the Roseg River. Whereas insect larvae were predominant in the benthos, hyporheic invertebrates were dominated by taxa belonging to the true groundwater fauna and the permanent hyporheos. Several permanently aquatic taxa (e.g. Nematoda, Ostracoda, Cyclopoida, Harpacticoida, Oligochaeta) appeared exclusively in the hyporheic zone or they extended farther upstream in the hyporheic layer than in the benthic layer. Leuctridae, Nemouridae, and Heptageniidae colonised hyporheic sediments where maximum water temperature was only 4 °C. 4. Despite strong seasonal changes in river discharge and physicochemistry in hyporheic water, the density and distribution of the hyporheos varied little over time. 5. Taxonomic richness increased markedly in the downstream part of a floodplain reach with an extensive upwelling zone. Upwelling groundwater not only maintained a permanent flow of water but also created several species‐rich habitats that added many species to the community of the main channel.     

The role of micro-organisms in the ecological connectivity of running waters

1. Riparian zones hold a central place in the hydrological cycle, owing to the prevalence of surface and groundwater interactions. In riparian transition zones, the quality of exfiltrating water is heavily influenced by microbial activities within the bed sediments. This paper reviews the role of micro-organisms in biogeochemical cycling in the riparian-hyporheic ecotone. 2. The production of organic substances, such as cellulose and lignin, by riparian vegetation is an important factor influencing the pathways of microbial processing in the riparian zone. For example, anaerobic sediment patches, created by entrainment of allochthonous organic matter, are focal sites of microbial denitrification. 3. The biophysical structure of the riparian zone largely influences in-stream microbial transformations through the retention of organic matter. Particulate and dissolved organic matter (POM and DOM) is retained effectively in the hyporheic zone, which drives biofilm development and associated microbial activity. 4. The structure of the riparian zone, the mechanisms of POM retention, the hydrological linkages to the stream and the intensity of key biogeochemical processes vary greatly along the river continuum and in relation to the geomorphic setting. However, the present state of knowledge of organic matter metabolism in the hyporheic zone suggests that lateral ecological connectivity is a basic attribute of lotic ecosystems. 5. Due to their efficiency in transforming POM into heterotrophic microbial biomass, attached biofilms form an abundant food resource for an array of predators and grazers in the interstitial environments of rivers and streams. The interstitial microbial loop, and the intensity of microbial production within the bed sediments, may be a primary driver of the celebrated high productivity and biodiversity of the riparian zone. 6. New molecular methods based on the analysis of the low molecular weight RNA (LMW RNA) allow unprecedented insights into the community structure of natural bacterial assemblages and also allow identification and study of specific strains hitherto largely unknown. 7. Research is needed on the development and evaluation of sampling methods for interstitial micro-organisms, on the characterization of biofilm structure, on the analysis of the biodegradable matter in the riparian-hyporheic ecotone, on the regulation mechanisms exerted on microbiota by interstitial predators and grazers, and on measures of microbial respiration and other key activities that influence biogeochemical cycles in running waters. 8. Past experiences from large-scale alterations of riparian zones by humans, such as the River Rhine in central Europe, undeniably demonstrate the detrimental consequences of disconnecting rivers from their riparian zones. A river management approach that uses the natural services of micro-organisms within intact riparian zones could substantially reduce the costs of clean, sustainable water supplies for humans.     

Benthic and hyporheic invertebrate assemblages along a flow intermittence gradient: effects of duration of dry events

1. A large proportion of the total river length on Earth comprises rivers that are temporary in nature. However, the effects of periodical dry events have received far less attention from ecologists than those of floods and low flows. 2. This study concomitantly examined the effects of flow intermittence on invertebrates from the streambed surface and from a depth of 30 cm in the hyporheic zone. Invertebrates were collected during 3 years in the Albarine River, France, before and after summer dry events from 18 sites (seven were perennial) distributed along a longitudinal flow intermittence gradient. 3. I predicted benthic and hyporheic density and taxonomic richness to decrease, and assemblage composition to shift from desiccation‐sensitive to desiccation‐resistant taxa with increased dry event duration. Second, I predicted benthic and hyporheic assemblages from sites that dried for longer periods to be nested subsets of assemblages from sites that dried for shorter periods. Last, I predicted a convergence in benthic and hyporheic assemblage composition with increasing duration of dry events, resulting from increased vertical migration of benthic taxa into the hyporheic sediments to cope with dry events. 4. Increased dry event duration in the Albarine River led to a decrease in both benthic and hyporheic density and taxonomic richness. Invertebrate assemblage composition shifted along the gradient of increasing flow intermittence, but broad taxonomic overlap between perennial and temporary reaches and nestedness patterns indicated that these shifts were because of the loss of taxa susceptible to drying rather than selection for desiccation‐resistant specialists. 5. Assemblage composition between benthic and hyporheic invertebrates diverged with increasing dry event duration, suggesting that the hyporheic zone did not act as a refuge during dry events in this river. 6. Quantitative studies on the relationships between ecology and intermittence are still rare but are needed to predict the consequences of future changes in flow intermittence. The relationships found in this study should be tested across a wide range of temporary rivers to better evaluate the generality of these findings.     

Influence of hyporheic zone characteristics on the structure and activity of microbial assemblages

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Biogeochemical Constraints on the Ecological Rehabilitation of Wetland Vegetation in River Floodplains

The European policy for river management during peak discharge periods is currently changing from exclusion strategies (reinforcement of dykes) to allowing a more natural situation by creating more floodplain space to reduce water levels during peak discharges. In addition, water retention and water storage areas have been created. The new measures are generally being combined with nature development strategies. Up till now, however, ecological targets of broadscale floodplain wetland restoration including sedge marshes, species-rich floodplain forests and carrs, riparian mesotrophic grasslands and other biodiverse riverine ecosystems, have hardly developed in these areas. Most studies on the conditions needed for sustainable ecological development of floodplains have focused on hydrological and geomorphological rather than biogeochemical issues (including nutrient availability and limitation). There are, however, large differences in the composition of river water and groundwater and in sediment quality between rivers in densely populated areas and those in more pristine areas, which serve as a reference. It is very likely that these factors, in combination with heavily altered hydrological regimes and the narrow areas confined between the dykes on both sides of the rivers, impose major constraints on sustainable ecological development of riverine areas. Another issue is that existing wetlands are generally considered to be very appropriate for water retention and conservation, although recent research has shown that this may pose a serious threat to their biodiversity. The present paper reviews the biogeochemical constraints on the combination of floodplain rehabilitation, water conservation and the conservation and development of wetlands. It is concluded that biogeochemical problems (mainly related to eutrophication) predominantly arise in less dynamic parts of the river system, to which the flood-pulse concept applies less. Sound knowledge of the biogeochemical processes involved will contribute to greater efficiency and a better prediction of the opportunities for restoration and development of riverine wetlands. This information can be directly applied in nature management, water management, policy-making and consultancy.     

Patchiness of River–Groundwater Interactions within Two Floodplain Landscapes and Diversity of Aquatic Invertebrate Communities

In fluvial systems, the interactions between rivers and groundwater significantly affect various ecological structures (for example, riparian vegetation) and functions. To examine the effects of hydrological exchange between groundwater and surface water on the distribution of aquatic invertebrates within a riverine landscape, we investigated the main stem, tributaries, and various surface and subsurface waters of two floodplains of a southern Alpine river (Brenno, Switzerland) in terms of their physicochemical, hydraulic, substratum, and faunal characteristics. The origins of the water were investigated by analyzing geomorphic settings and physicochemical variables. The two floodplains had different hydrological regimes. The middle floodplain was dominated by lateral inputs and exfiltration of hillslope groundwater from two different subcatchments. Bank filtration of river water sustained subsurface water only close to the channel. The aquatic habitats of the middle floodplain formed a rather homogeneous group with high taxon richness and intrahabitat diversities. These aquatic habitats resembled mountain springbrooks in their physicochemical characteristics and faunal compositions. In the lower floodplain, the exchange between river water and groundwater was more extensive. The aquatic floodplain habitats of the lower floodplain were fed mainly by deep and shallow alluvial groundwater, hyporheic exfiltration, and partly by surface water. In contrast to aquatic habitats of the middle floodplain, habitats of the lower floodplain showed a low intrahabitat and a high interhabitat diversity in terms of both substrate characteristics and faunal compositions. For both floodplains, ordination analyses showed a high concordance between the structure of the invertebrate community and the characteristics of the environmental habitat, including chemical, geomorphic, and hydraulic variables. Ordinations grouped aquatic habitats according to the origins of the waters. Taxon richness was related to local structural diversity, but species turnover was related to differential vertical and lateral connectivity. Exfiltration of groundwaters provided aquatic floodplain habitats for several specialized species. The results of this study show the significance of the river–groundwater connectivity for the creation of the habitat mosaic that sustains biodiversity in floodplains and thus have important implications for managing the ecological integrity of floodplains.     

Responses of hyporheic invertebrate assemblages to large-scale variation in flow permanence and surface–subsurface exchange

1. Flow permanence (the proportion of time that flowing water is present) strongly influences benthic invertebrate assemblages in ephemeral and intermittent river reaches. Effects of varying flow permanence on hyporheic invertebrate assemblages are not well understood, and have not previously been studied at large spatial scales. 2. We used a 52‐km long flow‐permanence gradient in the alluvial Selwyn River, New Zealand to assess hyporheic assemblage responses to variation in flow permanence and surface–subsurface exchange. The Selwyn mainstem consists of perennial and temporary reaches embedded in longer downwelling (losing) and upwelling (gaining) sections. 3. We predicted that hyporheic invertebrate diversity, density and assemblage stability would increase with increasing flow permanence. We further predicted that assemblage structure would be influenced by the relative contribution of downwelling and upwelling water at the reach‐scale. 4. Hyporheic invertebrates were collected at 15 river cross‐sections over a 13‐month period. As predicted, hyporheic taxon richness, density and assemblage stability varied directly with flow permanence. The distribution of taxa along the flow permanence gradient appeared to be related to desiccation resistance. However, it is possible that proximity to colonist sources also contributed to distribution patterns. 5. Taxon richness was significantly higher at sites in the gaining section compared with the losing section. Sites with high flow permanence in the gaining and losing sections supported distinct hyporheic assemblages, characterised by amphipods and isopods in the gaining section, and ostracods, Hydra sp. and the mayfly Deleatidium spp. in the losing section. 6. Results of the study suggest an expansion of the scope of the Hyporheic Corridor Concept to include large hyporheic flowpaths associated with unbounded alluvial plains rivers. Hyporheic assemblages in alluvial rivers are strongly influenced by large‐scale flow permanence gradients, large‐scale surface water–groundwater exchange, and their interactions.     

Influence of streambed sediment clogging on microbial processes in the hyporheic zone

1. The hyporheic zone plays a key role in hydrological exchange and biogeochemical processes in streambed sediments. The clogging of sediments caused by the deposition of particles in the bed of streams and rivers can decrease sediment permeability and hence greatly affect hyporheic microbial processes. 2. The main objective of this study was to determine the influence of sediment clogging on hyporheic microbial processes in three French rivers (the Usses, Drôme and Isère). In each river, microbial abundance and activity were studied at three depths (10, 30 and 50 cm) in the sediment at one unclogged (high porosity) and one clogged site (low porosity). 3. The results showed that the sediment clogging had inconsistent effects on microbial processes in the three rivers. Increases (Usses) or decreases (Drôme and Isère) in both aerobic and anaerobic processes were detected at the clogged sites compared to unclogged sites. These results suggest that microbial changes because of the sediment clogging are mainly mediated by the residence time of water within the hyporheic sediments. 4. A single model predicting the effect of clogging on hyporheic microbial processes cannot be applied generally to all rivers because the degree of clogging creates heterogeneous effects on flow rates between surface and interstitial waters. As a consequence, the influence of heterogeneous clogging on surface water–hyporheic exchanges needs to be evaluated by water tracing and hydraulic modelling to determine the links between microbial processes and hydraulic heterogeneity induced by clogging in hyporheic sediments.     

The influence of changes to river hydrology on freshwater fish in regulated rivers of the Murray–Darling basin

Freshwater fish are often used as an indicator of the response of the ecosystem to the restoration of river flows or the provision of environmental flows. The ability to model the biological response of fish depends on the capacity to establish clear relationships between changes in river hydrology and the fish assemblages in a river. The fish assemblage structure and the abundances of individual fish species were examined in relation to a hydrological index that described hydrological change in six regulated rivers in the Murray–Darling Basin. The hydrological index explained only a small amount of variation in fish assemblage structure. In addition, the abundances of individual fish species were only weakly correlated with the index of flow deviation. It is suggested that these results make the modelling of responses of fish assemblages to environmental water allocations unfeasible at a large scale and that future studies should concentrate on potentially more simple responses, such as the relationships of fish spawning and recruitment to specific aspects of river hydrology.     

溪流潜流层大型无脊椎动物生态学研究进展

溪流潜流层是溪流表层水和地下水相互作用的群落交错区,生物多样性丰富,是溪流生态系统的重要组成部分.大型无脊椎动物位于潜流层食物网的顶层,直接影响着潜流层物质和能量动态,是河流健康潜在的指示生物,调节着潜流层的环境净化和生态缓冲功能,对溪流生态系统发挥着至关重要的作用.潜流层大型无脊椎动物类群按生活史划分为偶入动物、非典型潜流层动物和典型潜流层动物.潜流层孔隙大小、孔隙水流速、溶解氧、温度、可利用的食物源、渗透系数和水力停留时间是影响大型无脊椎动物在潜流层分布的主要因素.对于潜流层这样一个特殊的生态界面,针对不同的研究目的应该选择合适的取样和调查方法.潜流层大型无脊椎动物的生活史和生活史对策,在溪流生态系统物质循环和能量流动中作用的定量化分析,基于潜流层大型无脊椎动物的河流健康评价体系,以及潜流层作为“庇护地”对于大型无脊椎动物分布和进化的生态学意义,都值得进一步关注和深入研究.     

Integrating and extending ecological river assessment: Concept and test with two restoration projects

While the number of river restoration projects is increasing, studies on their success or failure relative to expectations are still rare. Only a few decision support methodologies and integrative methods for evaluating the ecological status of rivers are used in river restoration projects, thereby limiting informed management decisions in restoration planning as well as success control. Moreover, studies quantifying river restoration effects are often based on the assessment of a single organism group, and the effects on terrestrial communities are often neglected. In addition, potential effects of water quality or hydrological degradation are often not considered for the evaluation of restoration projects.We used multi-attribute value theory to re-formulate an existing river assessment protocol and extend it to a more comprehensive, integrated ecological assessment program. We considered habitat conditions, water quality regarding nutrients, micropollutants and heavy metals, and five instream and terrestrial organism groups (fish, benthic invertebrates, aquatic vegetation, ground beetles and riparian vegetation). The physical, chemical and biological states of the rivers were assessed separately and combined to value the overall ecological state.The assessment procedure was then applied to restored and unrestored sites at two Swiss rivers to test its feasibility in quantifying the effect of river restoration. Uncertainty in observations was taken into account and propagated through the assessment framework to evaluate the significance of differences between the ecological states of restored and unrestored reaches. In the restored sites, we measured a higher width variability of the river, as well as a higher width of the riparian zone and a higher richness of organism groups. According to the ecological assessment, the river morphology and the biological states were significantly better at the restored sites, with the largest differences detected for ground beetles and fish communities, followed by benthic invertebrates and riparian vegetation. The state of the aquatic vegetation was slightly lower at the restored sites. According to our assessment, the presence of invasive plant species counteracted the potential ecological gain. Water quality could be a causal factor contributing to the absence of larger improvements.Overall, we found significantly better biological and physical states, and integrated ecological states at the restored sites. Even in the absence of comprehensive before-after data, based on the similarity of the reaches before restoration and mechanistic biological knowledge, this can be safely interpreted as a causal consequence of restoration. An integrative perspective across aquatic and riparian organism groups was important to assess the biological effects, because organism groups responded differently to restoration. In addition, the potential deteriorating effect of water quality demonstrates the importance of integrated planning for the reduction of morphological, water quality and hydrological degradation.     

A landscape perspective of surface–subsurface hydrological exchanges in river corridors

1. River corridors can be visualised as a three‐dimensional mosaic of surface–subsurface exchange patches over multiple spatial scales. Along major flow paths, surface water downwells into the sediment, travels for some distance beneath or along the stream, eventually mixes with ground water, and then returns to the stream. 2. Spatial variations in bed topography and sediment permeability result in a mosaic of patch types (e.g. gravel versus sandy patches) that differ in their hydrological exchange rate with the surface stream. Biogeochemical processes and invertebrate assemblages vary among patch types as a function of the flux of advected channel water that determines the supply of organic matter and terminal electron acceptors. 3. The overall effect of surface–subsurface hydrological exchanges on nutrient cycling and biodiversity in streams not only depends on the proportion of the different patch types, but also on the frequency distribution of patch size and shape. 4. Because nutrients are essentially produced or depleted at the downwelling end of hyporheic flow paths, reach‐scale processing rates of nutrients should be greater in stretches with many small patches (e.g. short compact gravel bars) than in stretches with only a few large patches (e.g. large gravel bars). 5. Based on data from the Rhône River, we predict that a reach with many small bars should offer more hyporheic refugia for epigean fauna than a reach containing only a few large gravel bars because benthic organisms accumulate preferentially in sediments located at the upstream and downwelling edge of bars during floods. However, large bars are more stable and may provide the only refugia during severe flood events. 6. In river floodplain systems exhibiting pronounced expansion/contraction cycles, hyporheic assemblages within newly created patches not only depend on the intrinsic characteristics of these patches but also on their life span, hydrological connection with neighbouring patches, and movement patterns of organisms. 7. Empirical and theoretical evidence illustrate how the spatial arrangement of surface–subsurface exchange patches affects heterogeneity in stream nutrient concentration, surface water temperature, and colonisation of dry reaches by invertebrates. 8. Interactions between fluvial action and geomorphic features, resulting from seasonal and episodic flow pulses, alter surface–subsurface exchange pathways and repeatedly modify the configuration of the mosaic, thereby altering the contribution of the hyporheic zone to nutrient transformation and biodiversity in river corridors.     

Biogeochemical implications of climate change for tropical rivers and floodplains

Large rivers of the tropics, many of which have extensive floodplains and deltas, are important in the delivery of nutrients and sediments to marine environments, in methane emission to the atmosphere and in providing ecosystem services associated with their high biological productivity. These ecosystem functions entail biogeochemical processes that will be influenced by climate change. Evidence for recent climate-driven changes in tropical rivers exists, but remains equivocal. Model projections suggest substantial future climate-driven changes, but they also underscore the complex interactions that control landscape water balances, river discharges and biogeochemical processes. The most important changes are likely to involve: (1) aquatic thermal regimes, with implications for thermal optima of plants and animals, rates of microbially mediated biogeochemical transformations, density stratification of water bodies and dissolved oxygen depletion; (2) hydrological regimes of discharge and floodplain inundation, which determine the ecological structure and function of rivers and floodplains and the extent and seasonality of aquatic environments; and (3) freshwater–seawater gradients where rivers meet oceans, affecting the distribution of marine, brackish and freshwater environments and the biogeochemical processing as river water approaches the coastal zone. In all cases, climate change affects biogeochemical processes in concert with other drivers such as deforestation and other land use changes, dams and other hydrological alterations and water withdrawals. Furthermore, changes in riverine hydrology and biogeochemistry produce potential feedbacks to climate involving biogeochemical processes such as decomposition and methane emission. Future research should seek improved understanding of these changes, and long-term monitoring should be extended to shallow waters of wetlands and floodplains in addition to the larger lakes and rivers that are most studied.     

Use of Shields stress to reconstruct and forecast changes in river metabolism

1. Discharge patterns of streams and rivers may be substantially affected by changes in water management, land use, or climate. Such hydrological alterations are likely to influence biotic processes, including overall ecosystem metabolism (photosynthesis and respiration). One regulator of aquatic ecosystem metabolism directly tied to hydrology is movement of bed sediments. 2. We propose that ecosystem metabolism can be reconstructed or predicted for any suite of hydrological conditions through the use of quantitative relationships between discharge, bed movement and metabolism. We tested this concept on a plains reach of the South Platte River in Colorado. 3. Movement of bed sediments was predicted from river discharge and the Shields stress, a ratio of velocity‐induced stress to sediment grain size. Quantitative relationships were established empirically between metabolic response to bed movement and recovery from bed movement, thus linking metabolism to hydrology. 4. The linkage of metabolism to hydrology allowed us to reconstruct daily photosynthesis and respiration over the 70‐year period for which discharge is known at our study site on the South Platte River. The reconstruction shows major ecological change caused by hydrological manipulation: the river has lost two‐thirds of its photosynthetic potential, and the ratio of photosynthesis to respiration is now much lower than it was prior to 1960. 5. The same approach could be used to anticipate ecological responses to proposed hydrological manipulations, to quantify benefits of hydrological restoration, or to illustrate potential effects of change in climate or land use on flowing‐water ecosystems.     

Interpreting spatial patterns in redox and coupled water–nitrogen fluxes in the streambed of a gaining river reach

Water pathways through permeable riverbeds are multi-dimensional, including lateral hyporheic exchange flows as well as vertical (upwelling and downwelling) fluxes. The influence of different pathways of water on solute patterns and the supply of nitrate and other redox-sensitive chemical species in the riverbed is poorly understood but could be environmentally significant. For example, nitrate-rich upwelling water in the gaining reaches of groundwater-fed rivers has the potential to supply significant quantities of nitrate through the riverbed to surface waters, constraining opportunities to deliver the goals of the EU Water Framework Directive to achieve ‘good ecological status’. We show that patterns in porewater chemistry in the armoured river bed of a gaining reach (River Leith, Cumbria) reflect the spatial variability in different sources of water; oxic conditions being associated with preferential discharge from groundwater and reducing conditions with longitudinal and lateral fluxes of water due to water movement from riparian zones and/or hyporheic exchange flows. Our findings demonstrate the important control of both vertical and lateral water fluxes on patterns of redox-sensitive chemical species in the river bed. Furthermore, under stable, baseflow conditions (<Q₉₀) a zone of preferential discharge, comprising 20 % of the reach by area contributes 4–9 % of the total nitrate being transported through the reach in surface water, highlighting the need to understand the spatial distribution of such preferential discharge locations at the catchment scale to establish their importance for nitrate delivery to the stream channel.     

河岸带生态系统植被与土壤对水文变化的响应研究进展

河岸带的植被与土壤是生态系统重要组成部分,对于维持河岸带的生态健康、生态系统服务与可持续性具有至关重要的作用。水文变化是河岸带生态系统的首要干扰因子,系统总结了水文变化对河岸带植被的特征以及植被形态、群落分布、繁殖、生存策略的影响,并阐述了河岸带水文和植被对土壤氮磷迁移转化的影响机制。根系作为土壤与植物地上部分之间物质、能量流动与信号传导的关键纽带,目前对根系的研究还较欠缺,需要加强水文变化对河岸带湿地植物根系形态、结构、功能特征的影响机理研究,以及湿地植物对水文变化的适应机制和耐受阈值方面的探究。在微观方面,应加强水文变化与植被等多因素耦合对土壤氮磷迁移转化过程的机理研究。河流形态和土壤的多样性决定着河岸带水文作用特征的复杂性,今后需注重河岸带个性特征与水文响应的关系研究。河岸带是横向的水陆生态过渡带和河流上下游的纵向生态廊道,亟需综合考虑和模拟流域土壤、植被与水文、人类活动之间的耦合关系,预测未来气候与社会经济情境下的河岸带生态系统演变规律,为河岸带生态系统的生态调节、生物多样性保护与生态恢复等提供理论依据与技术支撑。