Benthic and hyporheic invertebrate assemblages along a gradient of increasing streambed colmation by fine sediment

Streambed colmation by fine sediment, e.g. the deposition, accumulation and storage of fines in the substrate, is a major environmental concern throughout the world. Nevertheless, the ecological effects of streambed colmation on both benthic and hyporheic invertebrate assemblages have rarely been considered simultaneously. We studied a continuum of a naturally increasing percentage of fine sediment in three temperate rivers and hypothesized that the increasing percentage of fine sediment would decrease both benthic and hyporheic invertebrate densities and diversities, and reduce the similarities between them. To test these hypotheses, we first compared heavily, moderately and lightly clogged reaches located in downwelling areas and sampled invertebrates in the benthic zone and at 3 different depths (10, 30 and 50 cm) in the hyporheic zone. Secondly, we modified the sediment grain size distribution experimentally by increasing the percentage of fine sediment and using artificial substrates. The increasing colmation halved the hyporheic taxonomic richness and reduced benthic and hyporheic densities to a third. Some taxa were found in both zones, mainly in high colmation (e.g. Baetidae) or low colmation contexts (e.g. Orthocladiinae, Cyclopoida and Harpacticoida). The dissimilarity between benthic and hyporheic fauna (only at ?50 cm) was significantly higher in heavily clogged reaches than in moderately and lightly clogged ones, suggesting reduced vertical exchange of invertebrates or differential impacts between zones. The total abundance, taxonomic richness, percentage of EPT taxa and densities of most organisms observed using the artificial substrates decreased linearly with the increasing percentage of fine sediment in the experiment. Only the Ephemeroptera Caenis spp. and Heptageniidae disappeared above 30 and 50 % of fine sediment, respectively, suggesting that the response to increasing colmation are strongly taxon-specific. High amount of fine sediments within the substrate significantly decreased habitat quality for benthic and hyporheic invertebrates and thus limit the production of streams and their capacity to recover after disturbance. Moreover, the use of hyporheic invertebrates seems more relevant than benthic invertebrates to assess the effect of colmation and thus could be tested in future research as indicators.     

Focused Groundwater Controlled Feedbacks into the Hyporheic Zone During Baseflow Recession

Groundwater surface water interaction in the hyporheic zone remains an important challenge for water resources management and ecosystem restoration. In heterogeneous stratified glacial sediments, reach‐scale environments contain an uneven distribution of focused groundwater flow occurring simultaneously with diffusely discharging groundwater. This results in a variation of stream‐aquifer interactions, where focused flow systems are able to temporally dominate exchange processes. The research presented here investigates the direct and indirect influences focused groundwater discharge exerts on the hyporheic zone during baseflow recession. Field results demonstrate that as diffuse sources of groundwater deplete during baseflow recession, focused groundwater discharge remains constant. During baseflow recession the hyporheic zone is unable to expand, while the high nitrate concentration from focused discharge changes the chemistry of the stream. The final result is a higher concentration of nitrate in the hyporheic zone as this altered surface water infiltrates into the subsurface. This indirect coupling of focused groundwater discharge and the hyporheic zone is unaccounted for in hyporheic studies at this time. Results indicate important implications for the potential reduction of agricultural degradation of water quality.     

The role of the hyporheic zone across stream networks

Many hyporheic papers state that the hyporheic zone is a critical component of stream ecosystems, and many of these papers focus on the biogeochemical effects of the hyporheic zone on stream solute loads. However, efforts to show such relationships have proven elusive, prompting several questions: Are the effects of the hyporheic zone on stream ecosystems so highly variable in place and time (or among streams) that a consistent relationship should not be expected? Or, is the hyporheic zone less important in stream ecosystems than is commonly expected? These questions were examined using data from existing groundwater modelling studies of hyporheic exchange flow at five sites in a fifth‐order, mountainous stream network. The size of exchange flows, relative to stream discharge (Q_HEF:Q), was large only in very small streams at low discharge (area ≈ 100 ha; Q < 10 l/s). At higher flows (flow exceedance probability > 0·7) and in all larger streams, Q_HEF:Q was small. These data show that biogeochemical processes in the hyporheic zone of small streams can substantially influence the stream's solute load, but these processes become hydrologically constrained at high discharge or in larger streams and rivers. The hyporheic zone may influence stream ecosystems in many ways, however, not just through biogeochemical processes that alter stream solute loads. For example, the hyporheic zone represents a unique habitat for some organisms, with patterns and amounts of upwelling and downwelling water determining the underlying physiochemical environment of the hyporheic zone. Similarly, hyporheic exchange creates distinct patches of downwelling and upwelling. Upwelling environments are of special interest, because upwelling water has the potential to be thermally or chemically distinct from stream water. Consequently, micro‐environmental patches created by hyporheic exchange flows are likely to be important to biological and ecosystem processes, even if their impact on stream solute loads is small. Published in 2011 by John Wiley & Sons, Ltd.     

Impact of debris dams on hyporheic interaction along a semi‐arid stream

Hyporheic exchange increases the potential for solute retention in streams by slowing downstream transport and increasing solute contact with the substrate. Hyporheic exchange may be a major mechanism to remove nutrients in semi‐arid watersheds, where livestock have damaged stream riparian zones and contributed nutrients to stream channels. Debris dams, such as beaver dams and anthropogenic log dams, may increase hyporheic interactions by slowing stream water velocity, increasing flow complexity and diverting water to the subsurface. Here, we report the results of chloride tracer injection experiments done to evaluate hyporheic interaction along a 320 m reach of Red Canyon Creek, a second order stream in the semi‐arid Wind River Range of Wyoming. The study site is part of a rangeland watershed managed by The Nature Conservancy of Wyoming, and used as a hydrologic field site by the University of Missouri Branson Geologic Field Station. The creek reach we investigated has debris dams and tight meanders that hypothetically should enhance hyporheic interaction. Breakthrough curves of chloride measured during the field experiment were modelled with OTIS‐P, a one‐dimensional, surface‐water, solute‐transport model from which we extracted the storage exchange rate α and cross‐sectional area of the storage zone A_s for hyporheic exchange. Along gaining reaches of the stream reach, short‐term hyporheic interactions associated with debris dams were comparable to those associated with severe meanders. In contrast, along the non‐gaining reach, stream water was diverted to the subsurface by debris dams and captured by large‐scale near‐stream flow paths. Overall, hyporheic exchange rates along Red Canyon Creek during snowmelt recession equal or exceed exchange rates observed during baseflow at other streams. Copyright © 2005 John Wiley & Sons, Ltd.     

Alteration of physico-chemical and microbial properties in freshwater substrates by burrowing invertebrates

The hyporheic interstitial provides habitat for many different organisms – from bacteria to burrowing invertebrates. Due to their burrowing and sediment reworking behaviour, these ecosystem engineers have the potential to affect hyporheic processes such as respiration and nutrient cycling. However, there is a lack of studies that characterize the interactions between bioturbators, physico-chemical habitat properties and microbial communities in freshwater substrates. In a standardized laboratory experiment, we investigated the effects of three functionally different bioturbators, duck mussels (Anodonta anatina, Linnaeus 1758), mayfly nymphs (Ephemera danica, Müller 1764) and tubificid worms (Tubifex tubifex, Müller 1774), on the physico-chemical conditions and bacterial communities in hyporheic substrates. We hypothesized that different invertebrates distinctly alter habitat conditions and thus microbial community composition, depending on the depth and the manner of burrowing. A. anatina and E. danica caused an increase in interstitial oxygen concentration, whereas strong declines in oxygen concentration and redox potential were detected in the T. tubifex treatment. These effects on physico-chemical habitat properties were even detectable in open water. Mussels and tubificid worms also significantly influenced the composition of bacterial communities in the hyporheic zone. A loss or replacement of bioturbators in stream ecosystems due to anthropogenic habitat alterations is expected to result in shifts in microbial community compositions, with effects on nutrient fluxes, pollutant degradation and benthic food webs. An understanding of the effects of functionally different native and invasive bioturbators is crucial to predict changes in stream ecosystem functioning.     

Active reintroduction of benthic invertebrates to increase stream biodiversity

A large number of restoration projects aims to improve the ecological quality of streams and rivers by focusing on the stream structure. However, improved habitat heterogeneity often does not lead to natural recolonization by sensitive freshwater macroinvertebrate communities, particularly when the recolonization potential is low and source populations are absent. In preliminary studies we tested whether natural substrate exposures could be used to sample and transport benthic macroinvertebrates. In this pilot study we used these previously tested natural substrate exposures to sample freshwater invertebrates in a donor stream in order to actively (re-)colonize a recipient stream. In the course of three reintroduction campaigns, we were able to accumulate over 350,000 benthic invertebrates, including 25 indicator taxa of the orders Ephemeroptera, Plecoptera and Trichoptera and 30 taxa scoring positive in the German Fauna Index. In total, 45 taxa, which did not occur in the recipient stream before, were reintroduced. They were transported gently within natural substrate exposures and released on a stream bottom area of 500 m² in the recipient stream. We intended to study if an increase of benthos fauna in a recipient stream is possible, and if this increase will eventually improve the ecological status. So far, the natural substrate exposure-method demonstrated to be an adequate tool to accumulate and transport benthic macroinvertebrates and, in general, has the potential to increase the biodiversity of streams when used as assisted migration measure.     

Determining long time‐scale hyporheic zone flow paths in Antarctic streams

In the McMurdo Dry Valleys of Antarctica, glaciers are the source of meltwater during the austral summer, and the streams and adjacent hyporheic zones constitute the entire physical watershed; there are no hillslope processes in these systems. Hyporheic zones can extend several metres from each side of the stream, and are up to 70 cm deep, corresponding to a lateral cross‐section as large as 12 m², and water resides in the subsurface year around. In this study, we differentiate between the near‐stream hyporheic zone, which can be characterized with stream tracer experiments, and the extended hyporheic zone, which has a longer time‐scale of exchange. We sampled stream water from Green Creek and from the adjacent saturated alluvium for stable isotopes of D and ¹⁸O to assess the significance and extent of stream‐water exchange between the streams and extended hyporheic zones over long time‐scales (days to weeks). Our results show that water residing in the extended hyporheic zone is much more isotopically enriched (up to 11‰ D and 2·2‰ ¹⁸O) than stream water. This result suggests a long residence time within the extended hyporheic zone, during which fractionation has occurred owing to summer evaporation and winter sublimation of hyporheic water. We found less enriched water in the extended hyporheic zone later in the flow season, suggesting that stream water may be exchanged into and out of this zone, on the time‐scale of weeks to months. The transient storage model OTIS was used to characterize the exchange of stream water with the extended hyporheic zone. Model results yield exchange rates (α) generally an order magnitude lower (10^?5 s^?1) than those determined using stream‐tracer techniques on the same stream. In light of previous studies in these streams, these results suggest that the hyporheic zones in Antarctic streams have near‐stream zones of rapid stream‐water exchange, where ‘fast’ biogeochemical reactions may influence water chemistry, and extended hyporheic zones, in which slower biogeochemical reaction rates may affect stream‐water chemistry at longer time‐scales. Copyright © 2003 John Wiley & Sons, Ltd.     

Ecological impact of urban stormwater runoff studied in experimental flumes: Population loss by drift and availability of refugial space

Urban stormwater runoff discharged through sewer systems into streams causes flush spills of water and pollutants in the receiving water. To make the right decisions in future plannings of the very costly rehabilitation of sewer systems, a solid ecological data base on the critical parameters of sewer overflows is badly needed. Therefore, we designed a laboratory flume which was operated in circular flow mode (to ensure adaptation of the test organisms) and in flow-through mode during the simulation of sewer overflows (to allow a proper evaluation of population loss by drift). Examples on the behaviour during the adaptation phase and the population loss during the exposure to flush spills of water and/or a mixture of sewage and clean water of a benthic invertebrate (Gammarus pulex) demonstrate the potential of the flume to identify critical parameters of sewer overflows at quasireal-world-conditions. We found clear evidence for synergetic effects since the exposure to high flow and sewage caused higher population loss ofGammarus than the sum of population loss at exposure to only high flow or only sewage. Population loss considerably depended on the availability of refugial space: if the interstices of the gravel in the flume were silted, this loss was higher than at open interstices. Only ten minutes of movement of the material forming the flume bottom reduced the population ofGammarus to about 60 or 50% of its initial size. Hence, our data strongly suggest that the characteristics of the receiving stream (refugial space, bed stability) play an important role for the potential ecological impact of a sewer overflow. Changes of stream morphology and/or creation of refugial space plus an appropriate technical solution for overflow treatment may be less costly and more effective than a large-scale technical project. Thus, the stream itself should be a major element in future management decisions.     

Contrasting effects of cross-ecosystem subsidies and predation on benthic invertebrates in two Pacific coastal streams

Cross-ecosystem subsidies, such as terrestrial invertebrates and leaf litter falling into water as resources for aquatic communities, can vary across environmental gradients. We examined whether the effect of terrestrial subsidy inputs on benthic invertebrates was mediated by resident coastal cutthroat trout (Oncorhynchus clarki) in two representative streams. We experimentally manipulated the input rates (reduced, ambient) of terrestrial subsidies (terrestrial invertebrates and leaf litter) as well as the presence or absence of cutthroat trout in the two streams. The hypothesis that the reduction of terrestrial subsidies to the stream influences benthic invertebrate assemblages was supported by experimental results. The treatments of terrestrial subsidy reduction and cutthroat trout presence had a significant negative effect on benthic invertebrate community biomass and shredder biomass in East Creek with high natural terrestrial subsidy input and small amount of large wood in channel. In contrast, results from Spring Creek with low subsidy input and large amount of large wood in channel showed that only the terrestrial subsidy reduction significantly reduced the biomass of shredders. The effects of the terrestrial subsidy input and trout predation on benthic invertebrate communities varied between the two streams. Our results indicate that a subsidy effect on benthic communities can vary between nearby streams differing in canopy and habitats. This study, with the major finding of highly context-dependent effects of spatial subsidies, suggests that the interplay of resource subsidies and predators on invertebrate community assemblages can be site-specific and context-dependent on habitat features.     

A modelling study of hyporheic exchange pattern and the sequence,size, and spacing of stream bedforms in mountain stream networks,Oregon, USA

Studies of hyporheic exchange flows have identified physical features of channels that control exchange flow at the channel unit scale, namely slope breaks in the longitudinal profile of streams that generate subsurface head distributions. We recently completed a field study that suggested channel unit spacing in stream longitudinal profiles can be used to predict the spacing between zones of upwelling (flux of hyporheic water into the stream) and downwelling (flux of stream water into the hyporheic zone) in the beds of mountain streams. Here, we use two‐dimensional groundwater flow and particle tracking models to simulate vertical and longitudinal hyporheic exchange along the longitudinal axis of stream flow in second‐, third‐, and fourth‐order mountain stream reaches. Modelling allowed us to (1) represent visually the effect that the shape of the longitudinal profile has on the flow net beneath streambeds; (2) isolate channel unit sequence and spacing as individual factors controlling the depth that stream water penetrates the hyporheic zone and the length of upwelling and downwelling zones; (3) evaluate the degree to which the effects of regular patterns in bedform size and sequence are masked by irregularities in real streams. We simulated hyporheic exchange in two sets of idealized stream reaches and one set of observed stream reaches. Idealized profiles were constructed using regression equations relating channel form to basin area. The size and length of channel units (step size, pool length, etc.) increased with increasing stream order. Simulations of hyporheic exchange flows in these reaches suggested that upwelling lengths increased (from 2·7 m to 7·6 m), and downwelling lengths increased (from 2·9 m to 6·0 m) with increase in stream order from second to fourth order. Step spacing in the idealized reaches increased from 5·3 m to 13·7 m as stream size increased from second to fourth order. Simulated downwelling lengths increased from 4·3 m in second‐order streams to 9·7 m in fourth‐order streams with a POOL–RIFFLE–STEP channel unit sequence, and increased from 2·5 m to 6·1 m from second‐ to fourth‐order streams with a POOL–STEP–RIFFLE channel unit sequence. Upwelling lengths also increased with stream order in these idealized channels. Our results suggest that channel unit spacing, size, and sequence are all important in determining hyporheic exchange patterns of upwelling and downwelling. Though irregularities in the size and spacing of bedforms caused flow nets to be much more complex in surveyed stream reaches than in idealized stream reaches, similar trends emerged relating the average geomorphic wavelength to the average hyporheic wavelength in both surveyed and idealized reaches. This article replaces a previously published version (Hydrological Processes, 19 (17), 2915–2929 (2005) [ DOI:10.1002/hyp.5790 ]. See also retraction notice DOI:10.1002/hyp.6350 Copyright © 2006 John Wiley & Sons, Ltd.     

Riparian influence on hyporheic‐zone formation downstream of a small dam in the Blackland Prairie region of Texas

Small‐order streams have highly variable flows that can result in large temporal and spatial variation of the hyporheic zone. Dam construction along these intermittent headwater streams alters downstream flow and influences the hydrologic balance between stream water and the adjacent riparian zone. A 3‐year site study was conducted along an impounded second‐order stream to determine the water balance between stream, unsaturated zone, groundwater and riparian vegetation. The presence of the upstream impoundment provided near‐perennial water flow in the stream channel. The observed woody plant transpiration accounted for 71% of average annual water loss in the site. The overall contribution of stream water via the hyporheic zone to site water balance was 73 cm, or 44% of total inputs. This exceeded both rainfall and upland subsurface contribution to the site. A highly dynamic hyporheic zone was indicated by high water use from woody plants that fluctuated seasonally with stream water levels. We found leaf area development in the canopy layer to be closely coupled with stream and groundwater fluctuations, indicating its usefulness as a potential indicator of site water balance for small dam systems. The net result of upstream impoundment increased riparian vegetation productivity by influencing movement of stream water to storage in the groundwater system. Copyright © 2006 John Wiley & Sons, Ltd.     

Distribution and species composition of macroinvertebrates in the hyporheic zone of bed sediment

The hyporheic zone is a layer of substrate on a river bed where benthic animals normally live,grow,feed,reproduce,and exist for any portion of their life cycle.The hyporheic zone was studied by samplin...     

Hyporheic exchange flows induced by constructed riffles and steps in lowland streams in southern Ontario,Canada

Stream–subsurface water interaction induced by natural riffles and constructed riffles/steps was examined in lowland streams in southern Ontario, Canada. The penetration of stream water into the subsurface was analysed using hydrometric data, and the zone of > 10% stream water was calculated from a chemical mixing equation using tracer injection of bromide and background chloride concentrations. The constructed riffles studied induced more extensive hyporheic exchange than the natural riffles because of their steeper longitudinal hydraulic head gradients and coarser streambed sediments. The depth of > 10% stream water zone in a small and a large constructed riffle extended to > 0·2 m and > 1·4 m depths respectively. Flux and residence time distribution of hyporheic exchange were simulated in constructed riffles using MODFLOW, a finite‐difference groundwater flow model. Hyporheic flux and residence time distribution varied along the riffles, and the exchange occurring upstream from the riffle crest was small in flux and had a long residence time. In contrast, hyporheic exchange occurring downstream from the riffle crest had a relatively short residence time and accounted for 83% and 70% of total hyporheic exchange flow in a small and large riffle respectively. Although stream restoration projects have not considered the hyporheic zone, our data indicate that constructed riffles and steps can promote vertical hydrologic exchange and increase the groundwater–surface water linkage in degraded lowland streams. Copyright © 2006 John Wiley & Sons, Ltd.     

Influence of macroinvertebrates on physico‐chemical and microbial processes in hyporheic sediments

The objective of this study was to measure the effects of invertebrates on the physical characteristics and microbial processes in hyporheic sediments. We investigated the impacts of an assemblage of three taxa (asellids, chironomid larvae, and tubificid worms) on sediment distribution, water fluxes, sediment organic carbon, biofilm (attached bacteria) characteristics, and O₂, dissolved organic carbon NO₃^?, NO₂^?, and NH₄⁺ concentrations in slow filtration sand–gravel columns. The results showed that invertebrates clearly modified the distribution of particles in the sediment column, probably because of the structures (tubes, macropores, and faecal pellets) produced by the three taxa in the sediment. Our assessment of water fluxes indicated that invertebrate activities led to an increase in the porosity of the sediment columns. In addition, aerobic (O₂ consumption) and anaerobic (denitrification and fermentative decomposition of organic matter) microbial processes occurring in the sediment were stimulated in the presence of invertebrates. Finally, the present study demonstrates that invertebrates can act as ecosystem engineers in heterogeneous sediments that are under the influence of an advective flux of water. The solute residence time increased in columns containing the faunal assemblage. Micro‐organisms used more dissolved organic matter and nutrients in the presence of invertebrates because invertebrate activities increased the contact between the biofilm and water. We conclude that engineering by invertebrates in natural conditions modifies characteristics of the hyporheic zone and thereby enhances both the porosity of the sediment and the solute transport across the benthic interface. Copyright © 2002 John Wiley & Sons, Ltd.     

Interstitial flow through preferential flow paths in the hyporheic zone of the Oberer Seebach,Austria

We investigated interstitial flow velocities in the Oberer Seebach, Austria, with NaCl tracer injections at a sediment depth of 30 cm to estimate the hydraulic conditions experienced by invertebrates inhabiting the hyporheic zone. Flow velocity measured with tracers is taken as travel time of the water along a straight line between injection and sampling points, although the water flows around sediment particles, and thus travels a somewhat longer distance. From sections of stream sediment in which the interstitial spaces were replaced by concrete, we estimated that this difference amounts, on average, to 27% and used this factor to correct the results of our velocity measurements. Corrected interstitial water velocities ranged from 0.01 to 1.32 cm s^-1 and were independent of surface discharge. We also studied spatial flow patterns in the bed sediments with long-term tracer injections. The three-dimensional distribution of tracer concentrations 24 hours after the start of the injection indicated that interstitial water preferentially flows in a complex network of areas of high hydraulic connectivity. Reynolds numbers for flow in the hyporheic pore space ranged from 0.1 to 489, implying that the flow environment varies from laminar up to the zone of transition to turbulent flow. Therefore, invertebrates may have a size-related active choice of areas where either friction drag or pressure drag predominates. The consequence of flow patterns, such as those observed in our study, is that small-scale variability of hydraulic conditions may be an important determinant of the patchy invertebrate distribution in bed sediments.     

A temporal stable isotopic (δ18O, δD,d‐excess) comparison in glacier meltwater streams,Taylor Valley,Antarctica

In this paper, we describe the importance of hyporheic dynamics within Andersen Creek and Von Guerard Stream, Taylor Valley, Antarctica, from the 2010–2011 melt season using natural tracers. Water collection started at flow onset and continued, with weekly hyporheic‐zone sampling. The water δ¹⁸O and δD values were isotopically lighter in the beginning and heavier later in the season. D‐excess measurements were used as an indicator of mixing because an evaporative signature was evident and distinguishable between 2 primary end‐members (glacier meltwater and hyporheic zone). Hyporheic‐zone influence on the channel water was variable with a strong control on streamwater chemistry, except at highest discharges. This work supports previous research indicating that Von Guerard Stream has a large, widespread hyporheic zone that varies in size with time and discharge. Andersen Creek, with a smaller hyporheic zone, displayed hyporheic‐zone solute interaction through the influence from subsurface hypersaline flow. Overall, the evolution of Taylor Valley hyporheic‐zone hydrology is described seasonally. In mid‐December, the hyporheic zone is a dynamic system exchanging with the glacier meltwater in the channel, and with diminishing flow in January, the hyporheic zone drains back into the channel flow also impacting stream chemistry. This work adds new information on the role of hyporheic zone–stream interaction in these glacier meltwater streams.     

Patterns in stream longitudinal profiles and implications for hyporheic exchange flow at the H.J. Andrews Experimental Forest,Oregon, USA

There is a need to identify measurable characteristics of stream channel morphology that vary predictably throughout stream networks and that influence patterns of hyporheic exchange flow in mountain streams. In this paper we characterize stream longitudinal profiles according to channel unit spacing and the concavity of the water surface profile. We demonstrate that: (1) the spacing between zones of upwelling and downwelling in the beds of mountain streams is closely related to channel unit spacing; (2) the magnitude of the vertical hydraulic gradients (VHGs) driving hyporheic exchange flow increase with increasing water surface concavity, measured at specific points along the longitudinal profile; (3) channel unit spacing and water surface concavity are useful metrics for predicting how patterns in hyporheic exchange vary amongst headwater and mid‐order streams. We use regression models to describe changes in channel unit spacing and concavity in longitudinal profiles for 12 randomly selected stream reaches spanning 62 km² in the H.J. Andrews Experimental Forest in Oregon. Channel unit spacing increased significantly, whereas average water surface concavity (AWSC) decreased significantly with increasing basin area. Piezometer transects installed longitudinally in a subset of stream reaches were used to measure VHG in the hyporheic zone, and to determine the location of upwelling and downwelling zones. Predictions for median pool length and median distance between steps in piezometer reaches bracketed the median distance separating zones of upwelling in the stream bed. VHG in individual piezometers increased with increasing water surface concavity at individual points in the longitudinal profile along piezometer transects. Absolute values of VHG, averaged throughout piezometer transects, increased with increasing AWSC, indicating increased potential for hyporheic exchange flow. These findings suggest that average hyporheic flow path lengths increase—and the potential for hyporheic exchange flow in stream reaches decreases—along the continuum from headwater to mid‐order mountain streams. Copyright © 2005 John Wiley & Sons, Ltd.     

How representative are subfossil assemblages of Chironomidae and common benthic invertebrates for the living fauna of Lake De Waay,the Netherlands?

The distribution of benthic invertebrates and their subfossil remains was examined within the basin of De Waay, a dimictic, eutrophic lake in the Netherlands. We focused on Chironomidae, but also report the abundances of 11 invertebrate groups that potentially produce chitinous remains that are preserved in the fossil record, although their remains could only be identified at a coarser taxonomic resolution. Most living invertebrates sampled in different seasons were constrained to the littoral zone, with the exception of a few taxa (Ceratopogonidae, Chaoborus flavicans, and Chironomus) that are adapted to low oxygen conditions in the seasonally anoxic profundal zone. In contrast, assemblages of invertebrate remains in lake surface sediments were similar in the entire lake basin, suggesting that considerable numbers of invertebrate remains are transported and redeposited off-shore in Lake De Waay, due to its steep bathymetry. These results indicate that a single sediment sample obtained from the centre of this lake contains subfossil invertebrate remains originating from the entire lake basin. In Lake De Waay, the majority of taxa found in the living assemblages were identified as remains in lake surface sediments, at least for the Chironomidae that could be identified at a similar taxonomic level in living and subfossil assemblages. Of the total 44 chironomid taxa found in Lake De Waay, 35 taxa occurred in the living assemblages and 34 taxa occurred in the subfossil assemblages. Thirty chironomid taxa occurred both as living and subfossil specimens, and on average these 30 taxa represent 94% of the specimens encountered in a sediment sample. Five rare chironomid taxa present as living larvae were not detected in the subfossil assemblages. Conversely, eight rare and four common chironomid taxa were found in subfossil remains, but not in living assemblages. Our results indicate that subfossil assemblages in surface sediment samples provide spatially integrated and representative samples of the living assemblage. However, a combined approach examining both the living benthic invertebrate fauna and invertebrate remains in lake surface sediments will potentially give a more complete and detailed overview of benthic invertebrates in a lake ecosystem than an approach based exclusively on one of these groups.