Effects of buried leaf litter and vertical hydrologic exchange on hyporheic water chemistry and fauna in a gravel-bed river in northern New South Wales, Australia

1. Large amounts of coarse particulate organic matter (CPOM) are buried in the sand and gravel beds of many rivers during spates. The effects of these patchily distributed resources on hyporheic invertebrates and water chemistry are poorly understood. Buried CPOM may provide local ‘hot-spots’ of food for hyporheic detritivores and their predators, alter nutrient supply to nearby sediment biofilms, and generate habitat for some invertebrates. 2. To examine potential short-term effects on hyporheic water chemistry, nutrient concentrations and invertebrate assemblage composition, leaf packs were buried in downwelling (surface water infiltrating the hyporheic zone) and upwelling (hyporheic water emerging to the surface) zones at two sites along a gravel-bed river in northern New South Wales. At one site, pits were excavated to simulate leaf burial (procedural control) and plastic ‘leaves’ were buried to test whether invertebrates might respond to leaves as refuges rather than food. Hyporheic CPOM, sediment size fractions, and interstitial silt content were also quantified at these sites. 3. Dry weights of naturally buried CPOM (leaf litter and wood fragments) varied substantially (0.6–71.7 g L^–1 sediment). Amounts of CPOM did not differ between up- vs. downwelling zones or between sites. Hyporheic dissolved oxygen saturation was generally high (> 75%), and was lower in upwelling zones. The hyporheos was dominated taxonomically by water mites (≈ 20 species), whereas small oligochaetes were most abundant (40% of total abundance). Tiny instars of elmid beetle larvae and leptophlebiid mayfly nymphs were also common. Before experimental manipulation, faunal composition differed between up- and downwelling zones. In upwelling zones, bathynellaceans and blind peracarids were found, whereas small individuals of the surface benthos were common in samples from downwelling zones. This validated stratification of the experiment across zones of hydrologic exchange. 4. Twenty days after leaf burial, there was no effect of the treatments at either site on changes in most variables, including mean numbers of taxa and individuals per sample. Similarly, changes in faunal composition of the hyporheos in the treatments paralleled those in the controls except for a weak response in the buried leaves treatment in the upwelling zone at site 1. Artificially buried leaf litter does not seem to influence hyporheic water chemistry or fauna at these two sites. It is probable that naturally buried leaf litter is swiftly processed soon after entrainment and that repeating this experiment immediately after a flood may yield different results.     

Faunal response to benthic and hyporheic sedimentation varies with direction of vertical hydrological exchange

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Spatial responses of hyporheic invertebrates to seasonal changes in environmental parameters

1. Spatial relationships between hyporheic invertebrates and subsurface water flow patterns, sediment characteristics, water physicochemical parameters and several possible food sources were compared over three seasons at one site beneath a riffle. Measures of food sources included particulate organic matter (POM), bacterial activity (aerobic respiration, nitrate respiration and mineralisation of organic nitrogen) and microbial abundance. 2. Patterns of water flow changed significantly over the 9‐month study period, from predominantly upwelling beneath the entire riffle in spring, to distinct differentiation between downwelling and upwelling zones in summer and autumn. Water physicochemical parameters changed accordingly, showing gradually weaker correlations with depth and stronger correlations with zone between spring and autumn. 3. Despite these changes, depth remained the strongest predictor of invertebrate richness, density and taxon composition throughout the study period. However, invertebrate distributions were secondarily correlated with water physicochemical parameters, and a minor gradient in invertebrate distributions between downwelling and upwelling zones became stronger from spring to summer. 4. The correlations between invertebrates and physicochemical parameters changed in both magnitude and direction with season. In spring, invertebrates showed a negative correlation with surface water infiltration, whereas in summer and autumn, the correlation was positive. Correlations were strongest in summer, when interstitial dissolved oxygen concentrations were lowest. 5. No relationships were found between hyporheic invertebrates and POM, microbial abundance or activity. This suggests that at this site, proximity to the streambed surface and physicochemical variables are more important than the abundance of food in controlling invertebrate distributions.     

Benthic and hyporheic macroinvertebrate distribution within the heads and tails of riffles during baseflow conditions

The distribution of lotic fauna is widely acknowledged to be patchy reflecting the interaction between biotic and abiotic factors. In an in situ field study, the distribution of benthic and hyporheic invertebrates in the heads (downwelling) and tails (upwelling) of riffles were examined during stable baseflow conditions. Riffle heads were found to contain a greater proportion of interstitial fine sediment than riffle tails. Significant differences in the composition of benthic communities were associated with the amount of fine sediment. Riffle tail habitats supported a greater abundance and diversity of invertebrates sensitive to fine sediment such as EPT taxa. Shredder feeding taxa were more abundant in riffle heads suggesting greater availability of organic matter. In contrast, no significant differences in the hyporheic community were recorded between riffle heads and tails. We hypothesise that clogging of hyporheic interstices with fine sediments may have resulted in the homogenisation of the invertebrate community by limiting faunal movement into the hyporheic zone at both the riffle heads and tails. The results suggest that vertical hydrological exchange significantly influences the distribution of fine sediment and macroinvertebrate communities at the riffle scale.     

The effect of in-stream gravel extraction in a pre-alpine gravel-bed river on hyporheic invertebrate community

We investigated the effect of in-stream gravel extraction in a pre-alpine gravel-bed river on hyporheic invertebrate community, together with changes in the hyporheic geomorphology, physico-chemistry and biofilm activity. Hyporheic invertebrates were collected, together with environmental data, on seven sampling occasions from June 2004 to May 2005, at two river reaches—at the site of in-stream gravel extraction and at a site 2.5 km upstream. The hyporheic samples were taken from the river bed and from the gravel bars extending laterally from the stream channel. The invertebrate community was dominated by insect larvae (occasional hyporheos), followed by meiofauna (permanent hyporheos). Stygobionts were present at low species richness and in low densities. Gravel extraction from the stream channel led to changes in the patterns of water exchange between surface and subsurface and changes in the sediment composition at the site. Immediate reductions in density and taxonomic richness of invertebrates were observed, together with changes in their community composition. The hyporheic invertebrate community in the river recovered relatively fast (in 2.5 months) by means of density and taxonomic richness, while by means of community composition invertebrates needed 5–7 months to recover. The impact of fine sediments (<0.1 mm) on biofilm activity measured through ETS activity and hyporheic invertebrate density and taxonomic richness was strongly confirmed in this study.     

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.     

Benthic algal response to hyporheic-surface water exchange in an alluvial river

We studied the response of benthic algae to points of hyporheic-surface water exchange in the main channel of the Middle Fork Flathead River within the Nyack Flood Plain, Montana. We examined hyporheic exchange at 120 sites using piezometers and measuring vertical hydraulic gradient (VHG), hydraulic conductivity, and vertical discharge. We removed benthic algae from a single cobble at each site, and we used VHG to group sampling sites for statistical analysis. Algal cell density and chlorophyll a concentration were significantly higher at sites with hyporheic discharge (+VHG, upwelling) compared to both sites with hyporheic recharge (−VHG, downwelling) and sites with no hyporheic-surface water exchange (=VHG, neutral) (ANOVA, P < 0.05). The assemblages of algae at upwelling sites were also significantly different from downwelling and neutral exchange sites (ANOSIM, P < 0.05). Filamentous green algae Stigeoclonium sp. and Zygnema sp. and a chrysophyte, Hydrurus foetidus (Villars) Trevisan were abundant at upwelling sites, whereas an assemblage of diatoms Achnanthidium minutissimum (Kützing) Czarnecki, Cymbella excisa Kützing, Diatoma moniliformis Kützing, and Gomphonema olivaceoides Hustedt, were the most abundant taxa at downwelling and neutral exchange sites, occurring attached to, or in close association with the stalks of Didymosphenia geminata (Lyngbye) Schmidt. These data show that benthic algal communities are structured differently depending on the direction of hyporheic flux in the main channel of a large alluvial river, suggesting that hyporheic-surface exchange may influence the spatial distribution of main-channel benthic algae in rivers with hyporheic-surface water connectivity. Handling editor: J. Padisak     

Flood effects on invertebrates, sediments and particulate organic matter in the hyporheic zone of a gravel-bed stream

1. We investigated the effects of a flood on the fauna and physical habitat of the hyporheic zone of the Kye Burn, a fourth order gravel‐bed stream in New Zealand. 2. Freeze core hyporheic samples (to 50 cm depth) and benthic samples (to 10 cm) were taken, along with measurements of vertical hydrological gradient, before, 2 days after and 1 month after the flood (estimated return period: 1.5 years, estimated Q_max = 10.4 m³ s^?1). 3. The composition of the hyporheos differed over the three sampling occasions with fewer taxa collected immediately postflood than preflood. The equitability of the community was higher on both postflood occasions, consistent with the reduced densities of two abundant taxa (Leptophlebiidae and Copepoda). 4. Total invertebrate abundance was lower on the postflood occasions than preflood in both benthic (0–10 cm) and hyporheic (10–50 cm) sediments. Several taxa, including asellotan isopods and amphipods, recovered within 1 month of the event. Hyporheic densities of larval Hydora and nematodes did not differ among the three sampling occasions, but the water mite Pseudotryssaturus was more abundant 1 month after the flood than preflood. There was no evidence of vertical movements (to 50 cm) by any taxa in response to the flood. 5. The proportion of fine sediments (<1 mm) in the subsurface sediments (10–50 cm) increased over the three sampling occasions and median particle size declined, but sediment porosity did not change. More particulate organic matter was found in the sediments after the flood. 6. Our study provides little evidence that the hyporheic zone (to 50 cm) acted as a significant refuge during the flood event, although movements to or recolonisation from sediments deeper than 50 cm could explain the recovery of many crustacean and mite taxa within 1 month.     

Beneath the surface: Application of transparent super absorbent polymer substrates to track faunal activity within the sediment layer

1. Tracking the movement of organisms is a fundamental goal of many ecological studies. Several techniques exist in the study of terrestrial and aquatic fauna; however, to date, the ability to monitor aquatic fauna within the sediment layer efficiently and in multiple dimensions is lacking. Given the importance of subsurface sediments in supporting ecosystem functioning, this inability to observe organism behaviour represents a fundamental gap in our knowledge and limits our capability to holistically characterise the response of freshwater systems to stressors.
2. Here we present an experimental study that employs novel transparent super absorbent polymer substrates (c. 8–12 mm in diameter) in combination with computer vision technology, which enables, for the first time, real-time observation and tracking of organisms within the sediment layer under lotic flow conditions. Use of these substrates allowed the successful extraction of organism trajectories, which enabled the velocity and body orientation of a freshwater amphipod (Gammarus fossarum) in the sediment layer to be calculated in response to a number of vertical hydrological exchange treatments (upwelling, downwelling, and no vertical exchange).
3. Results indicate that under vertical hydrological exchange, a higher proportion of fast velocities (both horizontal and vertical) were recorded for G. fossarum in the sediment layer compared to no vertical exchange (control) conditions. This increase was most marked for upwelling flow exchange. We also observed a change in the body orientation of individuals in the sediment layer from a vertical alignment under no vertical exchange to a more horizontal one under downwelling and more notably upwelling flow exchange. This shift in body position was exacerbated under stronger vertical exchange rates.
4. We identified that following the flow transition of downwelling to upwelling conditions, there was an immediate shift (0–2 min) in both the orientation angle and activity level of individuals. This increased rate of activity was maintained for the individuals' velocity but not for their changing orientation angle. These trends were not apparent within the flow transition of no vertical exchange to downwelling flow.
5. Our new methodological approach enables vital insights into the behaviour of organisms within the sediment layer. Use of super absorbent polymer substrates allows real-time multi-directional tracking of multiple organisms in parallel. We believe the method represents an innovative tool that can be employed to tackle a wide range of ecological questions and thereby improve our mechanistic understanding of ecological responses to biotic and abiotic processes/stressors.

     

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.     

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.     

Hyporheic rehabilitation in rivers: restoring vertical connectivity

1. The hyporheic zone below the channel and banks of many rivers where surface water and ground water exchanges plays a crucial functional role in the biogeochemical transformation of water, mediated by active microbial biofilms. This zone also harbours assemblages of invertebrates that graze biofilms, contribute to secondary production, and can alter the porosity of the hyporheic zone through their movement or burrowing activities. 2. Many human activities cause interstitial sedimentation or disrupt surface–groundwater hydrological linkages, impacting upon ecological processes in the hyporheic zone. However, strategies for river rehabilitation seldom explicitly consider the hyporheic zone or seek to restore lost vertical linkages with groundwater. Instead, restoration goals target surface, riparian or floodplain features even though current river ecosystem theory emphasises the three dimensions of hydrological connectivity. To guide effective, holistic river restoration, scientists and managers therefore need information on the mechanisms by which energy and material are transferred in the hyporheic zone and which ecosystem services are thus provided. 3. Other gaps in our understanding of hyporheic zone rehabilitation include recruitment processes of the hyporheos and the relative importance of groups of hyporheic invertebrates in rivers differing in substratum size, disturbance frequency and groundwater linkages. Carefully designed experiments that assess responses to hyporheic rehabilitation strategies will provide valuable data at varying scales (e.g. distribution of hyporheic habitat types at the reach scale) for management as well as providing insights into the mechanisms controlling hyporheic invertebrate assemblages and ecological processes. Fully successful river rehabilitation must include restoration of vertical linkages between the river and its shallow groundwater aquifers.     

Surface–subsurface water exchange rates along alluvial river reaches control the thermal patterns in an Alpine river network

1. Water temperature is a key characteristic of stream ecosystems that is gaining scientific and managerial relevance as maximum temperatures in aquatic ecosystems increase worldwide.
2. To assess the effect of surface–subsurface water exchange on stream water temperature patterns, four alluvial reaches in the Tagliamento River basin (NE Italy), constrained by geomorphic knickpoints at the upper and lower end, and two to four hyporheic flowpaths within each reach, were continuously studied during summer 2007 and winter 2007–08. Water temperature was continuously monitored at the upstream and downstream knickpoints of the floodplains, as well as at discrete upwelling areas within each reach. Discharge and vertical hydraulic gradient were measured along the alluvial reaches, and the residence time and chemistry of upwelling water were assessed four times during the study.
3. Discharge variation along the study reaches revealed that massive hyporheic exchange occurred in all sites, ranging from 21% in reach 2–52% in reach 1. End member mixing analysis showed little influence of ground water, as almost all upwelling water was freshly infiltrated hyporheic water. Importantly, hyporheic exchange flows shaped surface temperature at the upwelling locations in all study reaches, providing potential thermal refugia for aquatic biota. At sites with highest hyporheic flow rates, net temperature change was also reflected at the floodplain scale.
4. The magnitude of the thermal change along a hyporheic flowpath was not related to the flowpath length but to the estimated ²²²Rn water age. Reduction in the diel thermal amplitude by hyporheic flows rather than net temperature change, reduced temperature extremes. Therefore, restoration activities to create thermal refugia should consider the role of hyporheic flows and enhance the exchange between surface and hyporheic waters.     

Subsurface hydrology and degree of burial affect mass loss and invertebrate colonisation of leaves in a woodland stream

SUMMARY 1. In deciduous forest streams, fallen leaves form a large component of the total organic matter budget, and many leaves become buried within stream sediments. We examined the processing of buried leaves as compared with those at the surface, and the influence of subsurface hydrology on processing rates.
2. Leaf packs were secured on the streambed surface or buried 10 cm deep in upwelling and downwelling reaches of a second-order stream in Michigan, U.S.A. Mass loss and invertebrate colonisation were measured from October to February.
3. Leaves buried in upwelling reaches lost mass more slowly (exponential decay coefficient, k =−0.0097) than did leaves from the other treatments (buried downwelling: −0.017; surface upwelling: −0.022; surface downwelling: −0.021).
4. Initially, more invertebrates colonised surface leaf packs than buried packs. During the remainder of the study, however, hydrology had a greater effect on invertebrate abundance than did burial, as more invertebrates were found in packs in downwelling reaches than in upwelling reaches.
5. Local subsurface hydrology and degree of burial, factors rarely considered in studies of detritus processing, can significantly influence mass loss and invertebrate colonisation of fallen leaves in streams. Furthermore, because of slower processing, subsurface zones may function as organic matter reservoirs that gradually 'spiral' carbon to downstream subsurface and surface habitats.     

Alternative tactics in spawning site selection by brook trout (Salvelinus fontinalis) related to incubation microhabitats in a harsh winter environment

1. Species with distributions that span a broad range of latitudes may have populations that exhibit distinct life history traits associated with environmental gradients. The majority of previous studies have indicated a strong association between spawning site selection by brook trout (Salvelinus fontinalis) and the presence of upwelling groundwater, but does this generalisation extend to the thermal regimes experienced at northern sites? 2. We investigated the role of hyporheic flow in redd site selection by brook trout in a relatively high‐latitude boreal system. Hyporheic flows through streambed substratums can be dominated by groundwater or surface water and may be influenced by the presence of morphological features. For autumn spawners such as brook trout, embryos situated in microhabitats where hyporheic flow in the shallow substratum is groundwater dominant (i.e. warmer in winter) are likely to experience accelerated development rates relative to embryos in redds where there is downwelling surface water (i.e. colder in winter). 3. We measured vertical hydraulic gradients (VHG) at the microhabitat scale and spatial and temporal variation in upwelling/downwelling flow and thermal regimes in brook trout spawning/incubation habitats. Additionally, we noted the proximity of redd sites to stream morphological features (e.g. riffle crests). 4. Our results indicate that upwelling flow was not a decisive cue in redd site selection at the microhabitat scale (10⁰ m) as an approximately equal number of redds were situated in microhabitats with upward flow as compared to microhabitats with downward flow through the redd. Redds situated on bedforms with convex longitudinal profiles (e.g. riffle crests, log steps) were associated with downward flow, whereas redds not immediately adjacent to these bedform features were associated with upward flow. Winter streambed temperatures confirmed that both sites with steady upwelling (i.e. warm incubation regime) and downwelling (i.e. cold incubation regime) were indeed selected by spawners. 5. Our observations that spawners utilised both cold‐regime and warm‐regime sites suggests the existence of distinct reproductive tactics related to hyporheic flow patterns in this boreal system. As temperature is the dominant factor controlling rates of embryonic development, the use of spawning microhabitats with distinct thermal regimes implies substantial differences in the timing of hatching and the phenology of emergence.     

Responses of Hyporheic Meiofauna to Habitat Manipulation

Interactions between interstitial meiofauna and physicochemical parameters of the hyporheic zone were examined via an in situ experiment on the Speed River, Ontario. The manipulation comprised reversing upwelling and downwelling zones at the riffle scale, and was maintained for 1 month. Significant differences in physicochemical parameters were detected between zones and between treatments (control vs. manipulated). Depth-related variables, such as sediment particle size, were most important in structuring the hyporheic community during pre- and post-manipulation phases. Flow reversal was largely successful, with more significant changes occurring in the original downwelling zone. For example, change from downwelling to upwelling resulted in decreased larval chironomid dominance but an increase in the numbers of oligochaetes, nematodes, mites, and copepods. However, under field conditions, it was difficult to keep other variables, such as water temperature, constant and some of these may have contributed to the changes seen in the meiofauna.     

The Effects of an Environmental Flow Release on Water Quality in the Hyporheic Zone of the Hunter River,Australia

Environmental flow releases have been advocated as a useful rehabilitation strategy for improving river condition but assessments of their success have typically focused on surface water quality and biota. In this study, we investigated the impacts of an environmental flow release on water temperature, conductivity, dissolved oxygen, and nitrate concentrations in surface and subsurface (hyporheic) water at upwelling and downwelling zones in three sites along the Hunter River, New South Wales, Australia. We hypothesised that the flow pulse would ‘flush’ the sediments with oxygenated water, stimulating hyporheic microbial activity and nitrification, enhancing nitrate concentrations over time. Surface and subsurface samples were collected before, 7 days after, and 49 days after an environmental flow release of 5000 Ml for a period of 3 days. No lasting effects on dissolved oxygen or conductivity were evident at most sites although dissolved oxygen declined over time at the downwelling site at Bowmans Crossing. At the downwelling zones at all sites, hyporheic nitrate concentrations declined initially following the release, but then rose or leveled off by Day 49. This initial drop in concentration was attributed to flushing of nitrate from the sediments. At two sites, nitrate concentrations had increased by Day 49 in the upwelling zones while at the third site, it fell significantly, associated with very low dissolved oxygen and likely reductive loss of nitrate. Electrical conductivity data indicate that potential inputs of agriculturally enriched groundwater may contribute to the nitrogen dynamics of the Hunter River. This study highlights the spatial heterogeneity that occurs in the hyporheic zone within and among sites of a regulated river, and emphasises the need for multiple-site surveys and an understanding of groundwater dynamics to assess physicochemical responses of the hyporheic zone to environmental flow releases.     

Mesocosm experiments reveal the direction of groundwater–surface water exchange alters the hyporheic refuge capacity under warming scenarios

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Vertical migration in streams: seasonal use of the hyporheic zone by the spinous loach <Emphasis Type="Italic">Cobitis shikokuensis</Emphasis>

Vertical hydrological connectivity between the surface stream and benthic and hyporheic zones plays a key ecological role in the biodiversity of lotic ecosystems because it allows surface and benthic organisms to use the hyporheic zone as a seasonal habitat and refuge. Use of the hyporheic zone by surface/benthic organisms has been well studied in invertebrates, but little is known about the importance of this connectivity for fishes. We investigated streambed surface and hyporheic densities (5–10, 15–20 and 20–25 cm below the streambed surface) of a stream fish, Cobitis shikokuensis, over a 20-month period in the Shigenobu River, southwestern Japan, to test the hypothesis that it uses the hyporheic zone for spawning and overwintering. In total, 1,804 individuals (13–58 mm total length) were captured from 33 streambed surface samplings and 102 individuals (10–46 mm total length) were present in 1,147 samples of 57 hyporheic samplings. Population densities in both zones peaked in late summer–early autumn due to the recruitment of age 0+ fish and a female with eggs was found in the hyporheic zone during the reproductive season. Both 0+ and older fish were absent from the streambed surface during winter, and fish densities were also lower in the hyporheic zone at this time. However, the vertical distribution of the fish tended to be skewed towards the deeper hyporheic layers from autumn to spring. These findings indicate that C. shikokuensis vertically migrates between the streambed surface and the hyporheic zone for spawning, rearing and overwintering, suggesting that the integrity of vertical hydrological connectivity in lotic systems is crucial for certain fish species.     

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.