Microform bed clusters: are they preferred habitats for invertebrates in a flood‐prone stream?

SUMMARY 1. Microform bed clusters are bedform microunits in streams. They consist of an obstacle clast (a large surface stone) against which other stones are stacked in a specific manner. Stream ecologists have suggested that these clusters are better flood refugia and more valuable habitats for invertebrates than single surface stones because of their higher stability during high‐flow events and their greater diversity of microhabitats available for colonisation. 2. To test these predictions in the Schmiedlaine, a flood‐prone prealpine stream, we sampled invertebrates on clustered and single stones on a total of seven occasions including before and after a moderate spate, two major floods and a minor spate. We also monitored surface particle stability during the second major flood and the minor spate. 3. Before the second flood (return period 1–2 years), we determined the exact positions of 60 clusters (60 obstacle clasts with 136 associated smaller particles) and 50 single stones in a 250‐m reach. We sampled the fauna on stones in the clusters, on well‐embedded single stones, and on loose single stones. Only six obstacle clasts and one single stone remained after the flood, implying movement of almost the entire surface layer. Therefore, we sampled the invertebrates on stones in newly formed clusters, on well‐embedded and single loose stones 3 and 11 days after the flood. The minor spate, in contrast to the second flood, moved only one of 495 monitored surface particles. 4. Three days after the flood, invertebrate density on clustered stones and both single stone categories were similar and equalled 34% of the mean pre‐flood density. Eight days later, density had almost doubled. The relatively high survival and rapid recovery suggest that invertebrates found refugia during the flood. However, the very low stability of clusters and single stones implied that surface particles were unimportant as refugia. 5. Total invertebrate density and taxon richness were never higher on clustered than on single stones (regardless of the timing of sampling relative to the last previous high‐flow event). Densities of the seven most common taxa and invertebrate community structure were also generally similar between particle types. We conclude that microform bed clusters cannot be regarded as more valuable invertebrate habitats than single surface stones in the Schmiedlaine.     

Patch- and reach-scale dynamics of a macrophyte-invertebrate system in a New Zealand lowland stream

Abundant growths of macrophytes are a common feature of streams in open lowland areas of New Zealand during summer, but the values of these to aquatic biota are poorly understood. We studied the temporal dynamics of, and associations amongst, elements of a macrophyte-invertebrate system to provide an improved information base for lowland stream management. The biomass of macrophytes increased significantly over the four quarterly sampling occasions from 43.8 g m^-2 in June to 370.8 g m^-2 in March; biomass was dominated by Egeria densa on all dates, except in December when Potamogeton crispus was dominant. We did not detect strong associations between epiphyton biomass and invertebrate abundance in our study, but this may reflect the fact that we sampled loosely-adhering epiphyton on young, surface-reaching shoots whereas invertebrates were collected from macrophytes growing through the water column. Density of some invertebrate species per gram dry weight of plant material varied by macrophyte type, with the chironomids Tanytarsus vespertinus and Naonella forsythi displaying positive correlations with Egeria and Potamogeton biomass, respectively. The shrimp Paratya curvirostris accounted for 50% of phytophilous invertebrate biomass, with Chironomidae the only other group to comprise more than 9%. Abundance of total phytophilous invertebrates displayed a positive linear relationship with macrophyte biomass in a sample (0.1 m²), and a humped relationship with species richness, such that highest numbers of taxa occurred at macrophyte biomass levels around 400 g dw m^-2. Our study suggests that intermediate macrophyte biomass levels are likely to enhance macroinvertebrate biodiversity in sandy-bottomed lowland streams. This revised version was published online in July 2006 with corrections to the Cover Date.     

Flow intermittency alters longitudinal patterns of invertebrate diversity and assemblage composition in an arid‐land stream network

1. Temporary streams comprise a large proportion of the total length of most stream networks, and the great majority of arid‐land stream networks, so it is important to understand their contribution to biotic diversity at both local and landscape scales. 2. In late winter 2010, we sampled invertebrate assemblages in 12 reaches of a large arid‐land stream network (including perennial and intermittent headwaters, intermittent middle reaches and perennial rivers) in south‐east Arizona, U.S.A. Intermittent reaches had then been flowing for c. 60 days, following a dry period of more than 450 days. We sampled a subset of the perennial study reaches three more times between 2009 and 2011. Since intermittent reaches were dry during these additional sampling periods, we used assemblage data from two other intermittent streams in the study network (sampled in 2004–05 and 2010) to explore interannual variability in intermittent stream assemblage composition. 3. Invertebrate richness was lowest in intermittent reaches, despite their often being connected to species‐rich perennial reaches. The assemblages of these intermittent reaches were not simply a subset of the species in perennial streams, but rather were dominated by a suite of stoneflies, blackflies and midges with adaptations to intermittency (e.g. egg and/or larval diapause). On average, 86% of individuals in these samples were specialists or exclusive to intermittent streams. Predators were 7–14 times more abundant in perennial than in intermittent reaches. 4. Despite being separated by long distances (12–25 km) and having very different physical characteristics, the assemblages of perennial headwaters and rivers were more similar to one another than to intervening intermittent reaches, emphasising the prime importance of local hydrology in this system. 5. The duration and recurrence intervals of dry periods, and the relative importance of dispersal from perennial refuges, probably influence the magnitude of biological differences between neighbouring perennial and temporary streams. Although perennial headwaters supported the highest diversity of invertebrates, intermittent reaches supported a number of unique or locally rare species and as such contribute to regional species diversity and should be included in conservation planning.     

Macroinvertebrate survival during cessation of flow and streambed drying in a lowland stream

相似文献   

Spatial and temporal patterns of invertebrate drift in streams draining a Neotropical landscape

1. Invertebrate drift in streams draining a tropical landscape in Costa Rica was studied to assess differences in assemblage composition above and below a major gradient break in geomorphic landform and to assess temporal patterns of drift in lowland reaches below the gradient break. The gradient break (～50 m a.s.l.) is the point at which the foothills of the Costa Rican Cordillera Central (piedmont) merge with the Caribbean coastal plain (lowlands).
2. Spatial patterns were assessed along two streams by sampling drift over 24 h once a month for 3 months in both the piedmont (90 m a.s.l.) and lowlands (30 m a.s.l.). Temporal patterns of drift were assessed through monthly diel sampling of three lowland sites over 8–10 months, encompassing both ‘dry’ (<400 mm precipitation per month, November to May) and wet (July to October) seasons.
3. Drift composition was insect dominated in piedmont sites and larval shrimp dominated in the lowlands. Percent similarity of assemblages between piedmont and lowland sites was low (range 26–43%) because of high larval shrimp densities in lowland versus piedmont sites.
4. Drift densities were higher during night than day, with peaks at sunset on all dates and at all sites. Diel patterns in drift agree with previous observations for the study area and support the ‘risk of predation’ hypothesis.
5. Analysis of monthly patterns in lowland sites showed high variability in drift densities; however, all major taxa were found every month. Overall, there was a trend for high invertebrate densities during the ‘dry’ season, but these trends were not significant.
6. Observed changes in drift composition support the concept of river zonation, which predicts a change in community composition along the stream continuum due to geomorphic features. Drift at lowland sites below the gradient break was dominated by shrimps, which are linked to marine environments via their migratory behaviour.     

Do stream macroinvertebrates use instream refugia in response to severe short-term flow reduction in New Zealand streams?

1. Demand for water is increasing and water managers need to know how much they can remove from a stream before there are significant detrimental effects on its biological integrity. Flow reduction alters a number of habitat variables known to be important to aquatic invertebrates such as depth, velocity, temperature and fine sediment accumulation. Some taxa may attempt to use instream refugia to mitigate the effects of flow reduction.
2. We experimentally manipulated flows by constructing weirs and diversions in three small New Zealand streams. Discharge was reduced by 88–96%. We tested the hypothesis that macroinvertebrates would use pools and the hyporheic zone as refugia during short-term (1-month) periods of reduced flow.
3. We sampled hyporheic invertebrates with colonization chambers and pool invertebrates with kick nets within a before-after, control-impact (BACI) experimental design. A suite of physicochemical parameters was measured concurrently including surface and hyporheic temperatures.
4. Flow reduction significantly decreased velocity (60–69%) in all streams. Depth (18–61%) and wetted width (24–31%) tended to decrease but these changes were not always significant. Sediment cover increased the most in farmland streams (10–80%). Apart from decreasing temperature range (18–26%), flow reduction had little impact on the surface water temperatures.
5. Flow reduction had no impact on the abundance of common pool macroinvertebrates or on the abundance, vertical distribution or community composition of hyporheic macroinvertebrates.
6. Our results suggest that aquatic macroinvertebrates are resistant to short-term, severe flow reduction as long as some water remains.     

Persistence and stability of lotic invertebrate communities in New Zealand

1. Persistence and stability of lotic invertebrate communities were determined at an annual time scale over a 9‐year period (1990–98) at 26 river sites over the northern half of New Zealand. In addition, a number of water quality variables were measured monthly and flow information collected continuously over the same period at each site. 2. The aim of the study was to describe the levels of interannual variability in invertebrate communities, and relate community changes to variability in environmental conditions. The consequences of this temporal variability for the sensitivity of predictive models were also investigated. 3. Levels of change in environmental conditions varied significantly between years, but were relatively similar across sites. In contrast, community persistence (similarity between years in species assemblage composition), and stability (similarity between years with respect to relative abundance of species in the community) both varied significantly between sites, but changes between years were similar. Community stability was highest at sites with relatively harsh flow conditions (high coefficient of variation, high relative size of floods), and was also greater in communities dominated by Ephemeroptera. 4. Relationships between change in environmental conditions and changes in community composition and structure were relatively weak for most individual sites. However, when average levels of change for each of the 26 sites were used, communities showed greater persistence under conditions where flow conditions remained relatively constant. Water quality changes had no significant effect on community persistence when assessed for all 26 sites combined. 5. Results from this study suggest that lotic invertebrate communities fluctuate around a relatively stable state, at least over a 9‐year period. However, the extent of interannual variation in community composition and structure observed, along with the relatively low degree of cluster fidelity observed within a single region, suggests that predictive models based on reference site conditions extrapolated over several years should be applied with caution in New Zealand streams.     

Biological,chemical and physical characteristics of downwelling and upwelling zones in the hyporheic zone of a north-temperate stream

Along a single stream riffle, there is a typical flow pattern in which surface water enters the hyporheic zone in a downwelling zone at the head of the riffle and hyporheic water returns to the stream surface in an upwelling zone at the tail of the riffle. Distinct patterns of physical and chemical conditions in the hyporheic zone are likely to determine patterns of microbial activity and occurrence of hyporheic fauna. Interstitial water and core samples were taken at three depths in the downwelling and upwelling zones of a single riffle in the Speed River, Southern Ontario, Canada. Physical and chemical characteristics of the hyporheic water, bacterial density, protein content, detritus content and faunal composition of the hyporheic sediment were analysed. The downwelling and upwelling zones differed significantly in temperature, pH, redox potential, dissolved oxygen and nitrate with significant positive correlations occurring among the latter three. There were no differences in bacterial density or detritus content between the two zones nor between depths in either zone, but protein content, considered to be a measure of biofilm biomass, was significantly higher in the downwelling zone. Total density of hyporheic fauna and the number of taxa decreased with increasing depth in both upwelling and downwelling zones, and were positively correlated with surface water characteristics (oxygen, temperature and nitrate), sediment protein content and detritus; however, only a weak correlation was found with zone. The composition of taxa differed between the two zones, and faunal distribution was correlated with dissolved oxygen, detritus, protein content and depth.     

Effects of pulsed and pressed disturbances on the benthic invertebrate community following a coal spill in a small stream in northeastern USA

In September 1999, a coal-carrying train derailed and spilled 180,000–270,000 kg of coal into the Cayuga Inlet near Ithaca, New York. This study determined the immediate effects of the coal spill and the clean up procedures on the aquatic invertebrate community, and whether the stream recovered from this event after 2 years. Benthic invertebrate samples were taken both upstream and downstream of the coal spill immediately following clean-up efforts and two years later. Just after the coal spill, the total abundance and species richness of aquatic invertebrates were significantly lower downstream of the spill, suggesting that the disturbance caused increased mortality and/or emigration compared to a reference site upstream. Taxa affected most were grazers and turbidity-susceptible invertebrates. Two years later the invertebrate communities were similar upstream and downstream of the spill, except for an increase in the percent of the dominant genus, Hydropsyche(Trichoptera: Hydropsychidae). We speculate that long-term effects of channelization of the stream that occurred during the clean-up prevented the invertebrate assemblage from returning to the conditions observed in a reference site upstream of the coal spill. We propose that large scale environmental clean-ups should be designed to avoid altering ecosystems permanently, and that streams should be allowed to recover naturally without destructive human intervention.     

Invertebrate colonization and processing of maple leaf litter in a forested and an agricultural reach of a stream

Processing of maple leaf (Acer saccharum Marsh) packs, their colonization by invertebrates and nutrient dynamics in leaves were investigated in a forested reach and agricultural reach of Canagagigue Creek, Ontario. Shredders, Pycnopsyche, and collectors, Ephemerella subvaria, Stenonema vicarium and Baetis were significantly more numerous in packs at the forest site than in packs at the agricultural site, whereas filter feeders, especially blackflies, were significantly more numerous in packs at the agricultural site. Weight loss of litter packs was nearly equivalent at the two sites. However, there were major differences in the mechanism of processing between the sites. Physical abrasion and microbial activity governed weight loss of maple leaf packs at the agricultural site, whereas processing was governed mainly by microbial and invertebrate activity and, to a much lesser extent, by physical abrasion at the forest site. Both shredders and collector species played an important role in the processing of leaf material at the forest site. Greater uptake of N and P (P<0.05 in spring) and higher C concentrations were observed in leaf packs at the forest site than the agricultural site. Therefore, the results support the concept of retention of nutrients in forested areas and their export in deforested (agricultural) areas. Findings also indicated that the processing of leaf litter is not an efficient means of monitoring changes in stream ecosystems since leaf processing is affected by many factors, particularly physical abrasion.     

Hyporheic community composition in a gravel-bed stream: influence of vertical hydrological exchange, sediment structure and physicochemistry

Summary 1. We studied the relative contributions of the magnitude and direction of vertical hydrological exchange, subsurface sediment composition and interstitial physicochemistry in determining the distribution of hyporheic invertebrates in the Kye Burn, a fourth order gravel‐bed stream in New Zealand. 2. In winter 2000 and summer 2001, we measured vertical hydrological gradient (VHG), dissolved oxygen, water temperature and water chemistry using mini‐piezometers, each installed in a different upwelling or downwelling zone. Next to every piezometer, a freeze core sample was taken to quantify the sediment, particulate organic matter and invertebrates. 3. Dissolved oxygen concentration at 25 cm was high on both occasions (>9 mg L^?1) but was higher in winter than summer. Interstitial water temperature was higher in down than upwellings and was substantially higher in summer than winter. Other features of the subsurface sediments and interstitial nitrate–nitrite concentrations were similar on both occasions and in up and downwellings. Interstitial ammonium and soluble reactive phosphorous concentrations were higher in winter than summer and ammonium was higher in up than downwelling areas. 4. The proportion of fine sediment (63 μm–1 mm), sediment heterogeneity and VHG accounted for the greatest proportion of variance in invertebrate distributions in both summer and winter. 5. The hyporheos was numerically dominated by early instar leptophlebiid mayfly nymphs and asellotan isopods. Water mites were a taxonomically diverse group with 13 genera. Taxonomic diversity (Shannon–Weaver), but not taxon richness, was higher in upwelling areas, reflecting lower numerical dominance by a few taxa in these locations. 6. Sediment composition (particularly the amount of fine sediments) and vertical hydrological exchange determined the composition and distribution of the hyporheos. Patchiness in these factors is important in planning sampling regimes or field manipulations in the hyporheic zone.     

Composition and zonation of benthic invertebrate communities in a New Zealand kauri forest stream

SUMMARY. The macro-invertebrate fauna of Waitakere River and its tributaries, a northern New Zealand kauri forest stream, was surveyed in January 1974 and January and June 1975, as part of a study on stream community dynamics. This survey and regular samples in the study area provided 144 taxa, almost three times the maximum number previously recorded from a New Zealand stream. Many species were restricted in distribution to a high gradient tributary, or the low gradient mainstream. These did not follow the division into rhithron and potamon families suggested for New Zealand.     

The hyporheic zone as a source of dissolved organic carbon and carbon gases to a temperate forested stream

The objective of this study was to examine chemical changes in porewaters that occur over small scales (cm) as groundwater flows through the hyporheic zone and discharges to a stream in a temperate forest of northern Wisconsin. Hyporheic-zone porewaters were sampled at discrete depths of 2, 10, 15, 61, and 183 cm at three study sites in the study basin. Chemical profiles of dissolved organic carbon (DOC), CO₂, CH₄, and pH show dramatic changes between 61 cm sediment depth and the water-sediment interface. Unless discrete samples at small depth intervals are taken, these chemical profiles are not accounted for. Similar trends were observed at the three study locations, despite each site having very different hydraulic-flow regimes. Increases in DOC concentration by an order of magnitude from 61 to 15 cm depth with a corresponding decrease in pH and rapid decreases in the molecular weight of the DOC suggest that aliphatic compounds (likely organic acids) are being generated in the hyporheic zone. Estimated efflux rates of DOC, CO₂, and CH₄ to the stream are 6.2, 0.79, 0.13 moles m² d^-1, respectively, with the vast majority of these materials produced in the hyporheic zone. Very little of these materials are accounted for by sampling stream water, suggesting rapid uptake and/or volatilization.     

The potential importance of unburned islands as refugia for the persistence of wildlife species in fire‐prone ecosystems

相似文献   

The response of hyporheic invertebrate communities to a large flood in the Hunter River, New South Wales

Previous studies on recovery in hyporheic communities have found that communities rapidly return to pre-disturbance levels. However, most of these studies have concentrated on small floods or ones with short return periods. I studied the impact of a large 1 in 6 year flood on the hyporheic community at 2 sites in the Hunter River, a large coastal river in New South Wales with a mean daily flow of 15 m³ s⁻¹. The flood peaked at 1270 m³ s⁻¹ and afterwards invertebrate densities at the 2 sites were 83 and 67% less than they were before. Recovery to pre-flood densities was slow but was aided by increases in the oligochaete and cyclopoid populations. At Site 1, there was a boom in oligochaete and cyclopoid numbers 61 d after the flood, but the communities resumed their pre-flood densities by Day 139. Recovery at Site 2 took 139 d. Most groundwater taxa (stygobites) living in the hyporheic zone did not recover from the disturbance when compared to non-stygobites. Apart from Microturbellaria and the harpacticoid Parastenocaris sp., numbers of all stygobite taxa continued to decline after the flood, becoming absent after 61 d. The poor recovery of stygobites is probably due to their adaptations for survival in the relatively stable groundwater environment. This study shows that hyporheic communities are sensitive to large bed-moving floods and supports the hypothesis that ecotonal species with a strong affinity to one ecosystem can be poor at recovering from disturbances that occur in an adjacent ecosystem.     

Effects of channelisation on stream habitat in relation to a spate and flow refugia for macroinvertebrates in northern Japan

1. The effects of channelisation on macroinvertebrates were examined in relation to a spate and flow refugia. Habitat components that can function as flow refugia were identified in a small, low‐gradient stream in northern Hokkaido, Japan. 2. Macroinvertebrates and their habitat characteristics (depth, current velocity and substratum) were sampled and measured in natural and channelised sections on three occasions: before, during and immediately after a spate. For macroinvertebrate sampling and habitat measurements, five (riffle, glide, pool, backwater and inundated habitats) and three (channelised‐mid, channelised‐edge and inundated habitats) habitat types were classified in the natural and channelised section, respectively. 3. The rate of velocity increase with discharge was compared among habitat types to determine which habitat types were less affected by increased discharge. The rate was the highest in riffles followed by glides and channelised‐mids. Backwaters maintained low current velocity even at high flow. In addition, current velocity in both natural and channelised inundated habitats was low relative to other habitat types during the spate. 4. Through the spate, total density of macroinvertebrates in channelised‐mids and taxon richness in both channelised‐mids and edges decreased. In the natural section, however, such a significant decrease was not found except for taxon richness in pools. This indicated that the spate had a greater impact on assemblages in the channelised section. Riffle assemblages exhibited a rapid recovery immediately after the spate, suggesting the existence of flow refugia in the natural section. Among the habitat types we examined, backwaters and inundated habitats appeared to have acted as flow refugia, because these habitats accumulated macroinvertebrates during the spate. 5. The lower persistence of the macroinvertebrate assemblage in the channelised section was attributable to the lower availability of flow refugia such as backwaters and inundated habitats. Our results emphasised the importance of considering flow fluctuations and refugia in assessing the effects of channelisation. In addition, the lateral heterogeneity of stream channels should be considered in stream restoration and management.     

山地河流浅滩生境潜流层大型无脊椎动物群落拓殖

通过包埋人工基质法研究大型无脊椎动物在山地河流潜流层中的拓殖过程。结果表明:群落个体密度在7—29 d呈"J"型增长,在29 d后骤然降低,55 d后呈波动趋势;物种丰富度在1—29 d呈增加趋势,29 d后呈波动状态;群落的生物量总体呈增加趋势。群落的物种丰富度、密度和生物量在第29、71和83天时没有显著性差异(P0.05),Shannon-Wiener多样性指数和Pielou均匀度指数在第55、71、83天没有显著性差异(P0.05),综合不同拓殖时间段物种的主成分分析,表明潜流层大型无脊椎动物群落在55 d后趋于稳定。群落优势种为摇蚊(Camptochironomus sp.)、河蚬(Corbicula fluminea)、四节蜉(Baetis sp.)、动蜉(Cinygmina sp.)、纹石蛾(Hydropsyche sp.)和扁泥甲科的一种(Psephenidae)。滤食者和收集者在整个拓殖过程中均是优势功能摄食群。群落拓殖过程是一个群落自身恢复能力和外部环境影响相互作用的过程,拓殖初期潜流层的结构是影响着无脊椎动物迁入的主要因素,中期动物的生活史特征起主要作用,稳定期之后群落可能受到各因素的综合影响。     

Flow- and substratum-mediated movement by a stream insect

1. In streams subject to frequent hydrologic disturbance, the ability of benthic invertebrates to disperse within the channel is key to understanding the mechanisms of flow refugium use and population persistence. This study focuses on crawling invertebrates, the effects on movement of abiotic factors (specifically, flow near the stream bed and bed micro‐topography) and the consequences for dispersal. 2. In a large flume, we observed individual cased caddisflies, Potamophylax latipennis, moving in fully turbulent flows over a precise replica of a water‐worked surface. From maps of movement paths, we quantified crawling behaviour and entrainment, and the influence of bed micro‐topography. We manipulated discharge and tested its effect on movement, linear displacement and areal dispersal. The highest discharge treatment was a disturbance to the caddis; the lowest discharge was not. Crawling behaviours were used to parameterise random walk models and estimate population dispersal, and to test the effects of abiotic factors on movement. 3. Bed micro‐topography influenced crawling in several ways. Caddis spent most of their time at the junctions between proud particles and the adjacent plane bed. The frequency distribution of turn angles was bimodal, with modal values approximating the angle required to travel around median‐sized particles. Larvae generally crawled downstream, but crawling direction relative to the flow was skewed by bed micro‐topography and was not directly downstream, unlike drift. 4. Caddis crawled for most of the time and discharge affected almost every aspect of their movement. As discharge increased, caddis crawled less often, more slowly and over shorter distances; they also became entrained more frequently and over greater distances. With increased discharge, caddis spent proportionately less time at the junctions between proud particles and the adjacent plane bed, and more time on the tops and sides of proud clasts. This is curious as most entrainment occurred from the tops and sides of clasts and entrainment is generally considered to be disadvantageous during disturbances. 5. Linear displacement (drift and entrainment combined) was downstream, but the relation between total displacement and discharge was complex. Total displacement decreased at intermediate discharge as crawling decreased, but increased at high discharge as entrainment and drift played a greater role in movement. 6. Within‐stream dispersal via crawling contained elements of both a correlated random walk (we observed directional persistence in turn angles) and a biased random walk (we observed downstream bias in move direction angles) and was best described as a biased correlated random walk. Dispersal was inversely related to discharge, suggesting that the ability of P. latipennis to crawl into flow refugia on the streambed is reduced at high flow.     

Hyporheic zone hydrology and nitrogen dynamics in relation to the streambed topography of a N-rich stream

The influence of riffle-pool units on hyporheic zone hydrology and nitrogen dynamics was investigated in Brougham Creek, a N-rich agricultural stream in Ontario, Canada. Subsurface hydraulic gradients, differences in background stream and groundwater concentrations of conservative ions, and the movement of a bromide tracer indicated the downwelling of stream water at the head of riffles and upwelling in riffle-pool transitions under base flow conditions. Channel water also flowed laterally into the floodplain at the upstream end of riffles and followed a subsurface concentric flow path for distances of up to 20 m before returning to the stream at the transition from riffles to pools. Differences in observed vs predicted concentrations based on background chloride patterns indicated that the hyporheic zone was a sink for nitrate and a source for ammonium. The removal of nitrate in the streambed was confirmed by the loss of nitrate in relation to co-injected bromide in areas of downwelling stream water in two riffles. Average stream water nitrate-N concentrations of 1.0 mg/L were often depleted to <0.005 mg/L near the sediment-water interface. Consequently, an extensive volume of the hyporheic zone in the streambed and floodplain had a large unused potential for nitrate removal. Conceptual models based mainly on studies of streams with low nutrient concentrations have emphasized the extent of surface-subsurface exchanges and water residence times in the hyporheic zone as important controls on stream nutrient retention. In contrast, we suggest that nitrate retention in N-rich streams is influenced more by the size of surface water storage zones which increase the residence time of channel water in contact with the major sites of rapid nitrate depletion adjacent to the sediment-water interface.