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.     

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.     

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.     

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.     

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.     

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.     

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.     

Factors controlling hyporheic respiration in a desert stream

1. Experimental manipulations were performed to determine the biological, chemical and physical attributes that govern sediment respiration in the hyporheic zone of Sycamore Creek, a Sonoran Desert stream. 2. Hyporheic respiration per unit volume of sediment was inversely related to diameter of sediment particles, indicating that respiration is affected by availability of substrate for microbial colonization (i.e. sediment surfaces). Respiration rate per unit surface area on sediments was positively correlated with particle diameter, indicating greater metabolic activity of microbes on larger sediments. 3. Hyporheic respiration was more than twice as high in water collected from the surface flow than from subsurface flow. Further, hyporheic respiration was highest immediately following exposure of sediments to surface water and declined over time, presumably due to exhaustion of labile organic matter. 4. Microbial activity was stimulated by addition of algal leachate; however, amendments of leaf leachate had little effect. Respiration was also elevated with dextrose and leucine amendments, but not with inorganic nitrogen additions, indicating hyporheic respiration is carbon limited. 5. Water from the stream surface is probably enriched in labile organic matter derived from algae and stimulates respiration at points of hydrologic downwelling where surface water enters hyporheic sediments. The physical structure of sediments further affects metabolism by affecting the area available for microbial attachment.     

Interactions between fauna and sediment control the breakdown of plant matter in river sediments

1. A substantial portion of particulate organic matter (POM) is stored in the sediment of rivers and streams. Leaf litter breakdown as an ecosystem process mediated by microorganisms and invertebrates is well documented in surface waters. In contrast, this process and especially the implication for invertebrates in subsurface environments remain poorly studied. 2. In the hyporheic zone, sediment grain size distribution exerts a strong influence on hydrodynamics and habitability for invertebrates. We expected that the influence of shredders on organic matter breakdown in river sediments would be influenced strongly by the physical structure of the interstitial habitat. 3. To test this hypothesis, the influence of gammarids (shredders commonly encountered in the hyporheos) on degradation of buried leaf litter was measured in experimental systems (slow filtration columns). We manipulated the structure of the sedimentary habitat by addition of sand to a gravel‐based sediment column to reproduce three conditions of accessible pore volume. Ten gammarids were introduced in columns together with litter bags containing alder leaves at a depth of 8 cm in sediment. Leaves were collected after 28 days to determine leaf mass loss and associated microbial activity (fungal biomass, bacterial abundance and glucosidase, xylosidase and aminopeptidase activities). 4. As predicted, the consumption of buried leaf litter by shredders was strongly influenced by the sediment structure. Effective porosity of 35% and 25% allowed the access to buried leaf litter for gammarids, whereas a lower porosity (12%) did not. As a consequence, leaf litter breakdown rates in columns with 35% and 25% effective porosity were twice as high as in the 12% condition. Microbial activity was poorly stimulated by gammarids, suggesting a low microbial contribution to leaf mass loss and a direct effect of gammarids through feeding activity. 5. Our results show that breakdown of POM in subsurface waters depends on the accessibility of food patches to shredders.     

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.     

Effects of burial on leaf litter quality,microbial conditioning and palatability to three shredder taxa

Summary 1. Heterotrophic microorganisms are crucial for mineralising leaf litter and rendering it more palatable to leaf‐shredding invertebrates. A substantial part of leaf litter entering running waters may be buried in the streambed and thus be exposed to the constraining conditions prevailing in the hyporheic zone. The fate of this buried organic matter and particularly the role of microbial conditioning in this habitat remain largely unexplored. 2. The aim of this study was to determine how the location of leaf litter within the streambed (i.e. at the surface or buried), as well as the leaf litter burial history, may affect the leaf‐associated aquatic hyphomycete communities and therefore leaf consumption by invertebrate detritivores. We tested the hypotheses that (i) burial of leaf litter would result in lower decomposition rates associated with changes in microbial assemblages compared with leaf litter at the surface and (ii) altered microbial conditioning of buried leaf litter would lead to decreased quality and palatability to their consumers, translating into lower growth rates of detritivores. 3. These hypotheses were tested experimentally in a second‐order stream where leaf‐associated microbial communities, as well as leaf litter decomposition rates, elemental composition and toughness, were compared across controlled treatments differing by their location within the streambed. We examined the effects of the diverse conditioning treatments on decaying leaf palatability to consumers through feeding trials on three shredder taxa including a freshwater amphipod, of which we also determined the growth rate. 4. Microbial leaf litter decomposition, fungal biomass and sporulation rates were reduced when leaf litter was buried in the hyporheic zone. While the total species richness of fungal assemblages was similar among treatments, the composition of fungal assemblages was affected by leaf litter burial in sediment. 5. Leaf litter burial markedly affected the food quality (especially P content) of leaf material, probably due to the changes in microbial conditioning. Leaf litter palatability to shredders was highest for leaves exposed at the sediment surface and tended to be negatively related to leaf litter toughness and C/P ratio. In addition, burial of leaf litter led to lower amphipod growth rates, which were positively correlated with leaf litter P content. 6. These results emphasise the importance of leaf colonisation by aquatic fungi in the hyporheic zone of headwater streams, where fungal conditioning of leaf litter appears particularly critical for nutrient and energy transfer to higher trophic levels.     

Early stages of leaf decomposition are mediated by aquatic fungi in the hyporheic zone of woodland streams

1. Leaf litter constitutes the major source of organic matter and energy in woodland stream ecosystems. A substantial part of leaf litter entering running waters may be buried in the streambed as a consequence of flooding and sediment movement. While decomposition of leaf litter in surface waters is relatively well understood, its fate when incorporated into river sediments, as well as the involvement of invertebrate and fungal decomposers in such conditions, remain poorly documented. 2. We tested experimentally the hypotheses that the small interstices of the sediment restrict the access of the largest shredders to buried organic matter without compromising that of aquatic hyphomycetes and that fungal decomposers in the hyporheic zone, at least partly, compensate for the role of invertebrate detritivores in the benthic zone. 3. Alder leaves were introduced in a stream either buried in the sediment (hyporheic), buried after 2 weeks of exposure at the sediment surface (benthic‐hyporheic), or exposed at the sediment surface for the entire experiment (benthic). Leaf decomposition was markedly faster on the streambed surface than in the two other treatments (2.1‐ and 2.8‐fold faster than in the benthic‐hyporheic and hyporheic treatments, respectively). 4. Fungal assemblages were generally less diverse in the hyporheic habitat with a few species tending to be relatively favoured by such conditions. Both fungal biomass and sporulation rates were reduced in the hyporheic treatment, with the leaves subject to the benthic‐hyporheic treatment exhibiting an intermediate pattern. The initial 2‐week stage in the benthic habitat shaped the fungal assemblages, even for leaves later subjected to the hyporheic conditions. 5. The abundance and biomass of shredders drastically decreased with burial, except for Leuctra spp., which increased and was by far the most common leaf‐associated taxon in the hyporheic zone. Leuctra spp. was one of the rare shredder taxa displaying morphological characteristics that increased performance within the limited space of sediment interstices. 6. The carbon budgets indicated that the relative contributions of the two main decomposers, shredders and fungi, varied considerably depending on the location within the streambed. While the shredder biomass represented almost 50% of the initial carbon transformed after 80 days in the benthic treatment, its contribution was <0.3% in the hyporheic one and 2.0% in the combined benthic‐hyporheic treatment. In contrast, mycelial and conidial production in the permanently hyporheic environment accounted for 12% of leaf mass loss, i.e. 2–3 times more than in the two other conditions. These results suggest that the role of fungi is particularly important in the hyporheic zone. 7. Our findings indicate that burial within the substratum reduces the litter breakdown rate by limiting the access of both invertebrate and fungal decomposers to leaves. As a consequence, the hyporheic zone may be an important region of organic matter storage in woodland streams and serve as a fungal inoculum reservoir contributing to further dispersal. Through the temporary retention of litter by burial, the hyporheic zone must play a significant role in the carbon metabolism and overall functioning of headwater stream ecosystems.     

Biotic and abiotic interactions between surface and interstitial systems in rivers

Water chemistry and community assemblages of surface and interstitial invertebrates were studied at seven sues on the French Rivers Rhône and Am at surface and at 50 cm depth into the bed sediments Chemical factors allowed differentiation of surface water from groundwater and detection of water exchanges defining clear downwelling and upwelling zones At some sites, interstitial water showed both surface and phreatic conditions, characterizing the underflows of the Rhone or of the Am In the interstitial area, most taxa showed no significant correlations with water chemistry Some epigean and hypogean fauna showed correlations with certain factors Clear relationships appeared between the water exchanges and the distribution of surface and interstitial faunas In interstitial samples, epigean and some hypogean species characterized the downwelling zones while others seemed to be strictly linked to the upwelling zones In surface samples, the presence of hypogean species was associated with regions of groundwater upwelling     

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

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

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The breakdown of buried and surface-placed leaf litter in an upland stream

The breakdown of buried leaves (Eucalyptus viminalis) was investigated using surface-placed and buried leaf packs in a riffle of the Acheron River, Victoria. Leaf packs buried to a depth of 10 cm were rapidly colonized by invertebrates, with the total numbers of individuals and species exceeding those on surface leaf packs. A larger proportion of leaves in buried leaf packs was grazed in comparison with those on the surface, with the intensity of grazing also being higher for leaves in buried packs. Both surface and buried leaf packs broke down rapidly with no significant difference in weight loss with time. The high level of breakdown of buried E. viminalis leaf litter observed in this study suggests that the hyporheos of Australian headwater streams may significantly contribute to the decomposition of particulate organic matter.     

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.     

Nitrogen processing in the hyporheic zone of a pastoral stream

The distribution of nitrogen-transforming processes, and factors controlling their rates, were determined within the hyporheic zone of a lowland stream draining agricultural land. In the field, physicochemical parameters were measured along a 10m-long hyporheic flow line between downwelling and upwelling zones. Sediment cores were retrieved from the stream bed surface, and from 20, 40 and 60cm deep in each zone, and in the laboratory, water from the corresponding depth was percolated through each core at the natural flow rate. Concentrations of nitrogen species and oxygen were measured before and after flow through each core. Denitrification was measured using a ¹⁵N-nitrate tracer. Shallow and downwelling zone samples were clearly distinct from deeper and upwelling zone samples in terms of physicochemical conditions, microbial processes and factors controlling nitrogen processing. Denitrification was highest in surface and downwelling zone cores, despite high oxygen levels, probably due to high pore-water nitrate concentrations in these cores and isolation of the denitrifying bacteria from oxygen in the bulk water by the hyporheic biofilms. Denitrification was limited by oxygen inhibition in the downwelling group, and by nitrate availability in the upwelling group. Strong evidence indicated that dissimilatory nitrate reduction to ammonium, occurred in almost all cores, and outcompeted denitrification for nitrate. In contrast, nitrification was undetectable in all but two cores, probably because of intense competition for oxygen. Field patterns and lab experiments indicated that the hyporheic zone at this moderately N-rich site is a strong sink for nitrate, fitting current theories that predict where hyporheic zones are nitrate sinks or nitrate sources.     

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.