Surface and Hyporheic Oligochaete Assemblages in a French Suburban Stream

The Chaudanne stream received urban inputs discharged through combined sewer overflows (CSOs). The water quality was not severely impaired, with pollution mainly of organic origin. Oligochaete assemblages were studied in the coarse surface sediments and the hyporheic zone and sampled at four sites on seven occasions during 2000 and 2001. The seasonal distribution of oligochaete assemblages was analyzed by a PCA, the oligochaete species being assigned to functional traits (FTrs). Site 1, located upstream of the CSOs, was characterized by FTrs 1 and 2 (species indicating permeability and those intolerant to water pollution). Below the CSOs, high densities of oligochaetes occurred in the benthic layer of sites 3 and 4, with a predominance of FTr3 (species with tolerance to water pollution). At site 4, FTr4 species (indicative of sludge conditions) constantly predominated in the hyporheic system, but predominated in the benthos only during low stream discharges associated with peaks in CSOs. FTr3 was related to amounts of the oxidized forms of nitrogen, high stream discharges and probably to groundwater upwellings and the sludge tolerant species group (FTr4) was associated with high NH₊⁴ contents. We are now testing the relevance and generalization of this new approach.     

Oligochaete communities in the hyporheic zone of a glacial river,the Roseg River,Switzerland

We examined the composition and distribution of oligochaete communities in the hyporheic zone of the Roseg River, a glacial river in southeastern Switzerland. Ten oligochaete species were collected from 11 sites distributed along an 11-km reach of the river, downstream of the Tschierva Glacier. The most frequently encountered species were Dorydrilus michaelseni, Propappus volki, Cernosvitoviella atrata and Cernosvitoviella carpatica. Six other species were relatively scarce and exhibited a discontinuous distribution pattern only in the downstream sites. At the most downstream site, a population of eyeless Nais communis was discovered with anatomical traits apparently related to a subterranean life-adaptation of this surface-living species. The species richness and abundance of oligochaete communities were relatively constant over time, but increased at sites where groundwater entered the stream. The spatial distribution of several oligochaete species was linked to the longitudinal arrangement of groundwater upwelling areas. This observation suggested that groundwater was an important upstream migration pathway for oligochaetes during the glacial retreat.     

Superficial and hyporheic oligochaete communities as indicators of pollution and water exchange in the River Moselle, France

Benthic oligochaetes were sampled on three occasions (June, August and October 1992) in the upper (0–10 cm) and hyporheic (35–45 cm depths) sediments at five sites of the River Moselle, from upstream of the town of Epinal to Velle-sur-Moselle. The first site (upstream from Epinal) is considered as unpolluted and the four remaining sites are polluted by industrial effluents. The most polluted stations were generally dominated by the pollution tolerant taxon Limnodrilus. Numbers of individuals of this taxon decreased at the less polluted last site in recovery zone, and were also scarce in the first unpolluted site. It is noteworthy that these tendencies were observed in both superficial and hyporheic substrates and to the greatest degree in hyporheic ones. At the unpolluted site, the hyporheic habitat is dominated by the groundwater species Propappus volki, Pristina spp., Pristinella spp. At the less polluted site (last site), the deep sediments are dominated by groundwater species and the Tubificidae without hair setae decrease from June to October. As a result of water exchange between superficial and subterranean waters, superficial substrates of the first and the last stations tend to be colonised by a high proportion of hyporheic species that suggests that flow is primarily from subterranean to superficial waters. The contrary is the case at other polluted stations which are characterised by the invasion of hyporheic substrates by the pollution tolerant superficial taxa Limnodrilus. This suggests that water flows from the river to the deeper groundwater. These two stations are located near drinking water plants which utilise groundwater, thus increasing the vulnerability of groundwater to surface contaminants.     

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.     

Reappraisal and range extension of non-indigenous Mysidae (Crustacea, Mysida) in continental and coastal waters of eastern France

A survey of species belonging to the family Mysidae, conducted in June 2007 in fresh- to brackish waters of eastern France, revealed a recent range extension of the invasive Ponto-Caspian species Hemimysis anomala to the Moselle, Saône, and Rhône rivers. In the estuary of the Grand Rhône it reached for the first time the Mediterranean coast. The network of navigation canals in NE France was likely a key element of its north to south pathway starting from the Rhine River. Important range extensions were also noted for Limnomysis benedeni in this network and in the Moselle River. The euryhaline species Neomysis integer, endemic in coastal waters of the NE Atlantic, was found in the Rhône delta, thus confirming previous very rare records in the 1930–1950s from the Mediterranean coast of France. Invasion mechanisms and pathways, expansion potential, and establishment conditions of the species are discussed.     

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

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

Genetic analysis of potential postglacial watershed crossings in Central Europe by the bullhead (Cottus gobio L.)

Natural colonizations across watersheds have been frequently proposed to explain the present distributions of many freshwater fish species. However, detailed studies of such potential watershed crossings are still missing. Here, we investigated potential postglacial watershed crossings of the widely distributed European bullhead (Cottus gobio L.) in two different areas along the Rhine–Rhône watershed using detailed genetic analysis. The main advantage of studying bullheads vs. other freshwater fish species is that their distribution has been lightly influenced by human activities and as such, interpretations of colonization history are not confounded by artificial transplantations. The genetic analyses of eight microsatellite loci revealed strong genetic similarities between populations of both sides of the Rhine–Rhône watershed in the Lake Geneva area, giving strong evidence for a natural watershed crossing of bullheads from the upper Rhine drainage into the Rhône drainage in the Lake Geneva area likely facilitated by the retreat of the glaciers after the last glacial maximum some 20 000 years ago. Populations from the Lake Geneva basin were genetically more similar to populations from across the watershed in the upper Rhine drainage than to populations further downstream in the lower Rhône. In contrast, populations from Belfort, an area, which was not covered by ice during the last glacial maximum, showed strong genetic differentiation between populations of the upper Rhine and Rhône drainages. Based on our results on the bullhead, we propose that glacial retreat may have eased the dispersal of numerous European freshwater fish species across several geological boundaries.     

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.     

Spatial distribution patterns of the hyporheic invertebrate communities in a polluted river in Romania

The purpose of this study was to examine the sensitivity, in a field situation, of the hyporheic fauna to pollution by heavy metals and also to test the use of oxidative stress enzymes produced by this fauna as a sensitive indicator of oxidative stress generated by chemical contamination. This was done by surveying the patterns of distribution, structure, and composition of hyporheic invertebrate communities in one of the most polluted rivers in Romania. Twelve permanent sampling stations with differing water qualities were established along a 180 km transect of the Arie? River. Data on hyporheic invertebrate abundance and richness, chemistry of the surface and hyporheic water and interstitial suspended particles were analyzed via multifactorial analyses. In the downstream, more polluted stations, epigean species were less abundant and hyporheic communities, especially macrocrustaceans and oligochetes, became dominant. The higher levels of hyporheic invertebrate biodiversity in the moderately polluted stations compared to highly polluted, and the increase of the number of some hyporheos (especially macrocrustaceans) in the moderately polluted stations, suggested that the hyporheic fauna was more tolerant of heavy metal pollution than the surface water fauna of the area. However, the different richness and abundance of hyporheic fauna in sites of similar water chemistry suggested that additional factors, such as sediment structure are shaping the spatial distribution of hyporheic fauna. Strong correlations between superoxide dismutase (SOD) activity in pooled tissues extracts and some chemical parameters suggest that oxidative stress enzymes may prove to be sensitive indicators of chemical pollution in hyporheic zones.     

The role of water exchange between a stream channel and its hyporheic zone in nitrogen cycling at the terrestrial-aquatic interface

The subsurface riparian zone was examined as an ecotone with two interfaces. Inland is a terrestrial boundary, where transport of water and dissolved solutes is toward the channel and controlled by watershed hydrology. Streamside is an aquatic boundary, where exchange of surface water and dissolved solutes is bi-directional and flux is strongly influenced by channel hydraulics. Streamside, bi-directional exchange of water was qualitatively defined using biologically conservative tracers in a third order stream. In several experiments, penetration of surface water extended 18 m inland. Travel time of water from the channel to bankside sediments was highly variable. Subsurface chemical gradients were indirectly related to the travel time. Sites with long travel times tended to be low in nitrate and DO (dissolved oxygen) but high in ammonium and DOC (dissolved organic carbon). Sites with short travel times tended to be high in nitrate and DO but low in ammonium and DOC. Ammonium concentration of interstitial water also was influenced by sorption-desorption processes that involved clay minerals in hyporheic sediments. Denitrification potential in subsurface sediments increased with distance from the channel, and was limited by nitrate at inland sites and by DO in the channel sediments. Conversely, nitrification potential decreased with distance from the channel, and was limited by DO at inland sites and by ammonium at channel locations. Advection of water and dissolved oxygen away from the channel resulted in an oxidized subsurface habitat equivalent to that previously defined as the hyporheic zone. The hyporheic zone is viewed as stream habitat because of its high proportion of surface water and the occurrence of channel organisms. Beyond the channel's hydrologic exchange zone, interstitial water is often chemically reduced. Interstitial water that has not previously entered the channel, groundwater, is viewed as a terrestrial component of the riparian ecotone. Thus, surface water habitats may extend under riparian vegetation, and terrestrial groundwater habitats may be found beneath the stream channel.     

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

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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.     

Bioassessment of Selected Inland Water Bodies in Sri Lanka Using Benthic Oligochaetes with Consideration of Temporal Variations

The organic pollution status of three inland water bodies in Sri Lanka was evaluated using benthic oligochaete communities with the environmental characteristics considering temporal variation over one year period. Oligochaete species richness and abundance was consistently higher in the highly eutrophic and organically polluted Lake Beira in comparison with the less polluted water bodies. In a cluster analysis of oligochaete community compositions, the assemblages grouped by the water bodies rather than by months. Principal component analysis revealed that the structure of the oligochate community was mainly influenced by conductivity, nitrate, biochemical oxygen demand levels of water and organic carbon content of the sediment. In the ordination, the polluted Lake Beira separated from the other water bodies along the first axis. The results revealed that there could be a potential for using benthic oligochaetes for bioassessment of inland water bodies in Sri Lanka. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Characterizing seasonal changes in physicochemistry and bacterial community composition in hyporheic sediments

The hyporheic zone of stream ecosystems is a critical habitat for microbial communities. However, the factors influencing hyporheic bacterial communities along spatial and seasonal gradients remain poorly understood. We sought to characterize patterns in bacterial community composition among the sediments of a small stream in southern Ontario, Canada. We used sampling cores to collect monthly hyporheic water and sediment microbial communities in 2006 and 2007. We described bacterial communities terminal-restriction fragment length polymorphism (TRFLP) and tested for spatial and seasonal relationships with physicochemical parameters using multivariate statistics. Overall, the hyporheic zone appears to be a DOC, oxygen, and nitrogen sink. Microbial communities were distinct from those at the streambed surface and from soil collected in the adjacent watershed. In the sediments, microbial communities were distinct between the fall, spring, and summer seasons, and bacterial communities were more diverse at streambed surface and near-surface sites compared with deeper sites. Moreover, bacterial communities were similar between consecutive fall seasons despite shifting throughout the year, suggesting recurring community assemblages associated with season and location in the hyporheic zone. Using canonical correspondence analysis, seasonal patterns in microbial community composition and environmental parameters were correlated in the following way: temperature was related to summer communities; DOC (likely from biofilm and allochthonous inputs) influenced most fall communities; and nitrogen associated strongly with winter and spring communities. Our results also suggest that labile DOC entering the hyporheic zone occurred in concert with shifts in the bacterial community. Generally, seasonal patterns in hyporheic physicochemistry and microbial biodiversity remain largely unexplored. Therefore, we highlight the importance of seasonal and spatial resolution when assessing surface- and groundwater interactions in stream ecosystems.     

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.     

Transport of Algal Cells in Hyporheic Sediments of the River Elbe (Germany)

The advective transport of algal cells into the interstices of the hyporheic zone of the River Elbe was spatially and temporally heterogenous. Even deep sediment layers were reached by large phytoplankton species. Therefore, it is suggested that (i) the advective interstitial transport patterns vary between different algal sizes and morphotypes and (ii) sediment characteristics, expressed by the permeability coefficient k_f of porous media, affect retention and retardation of surface water algae during subsurface transport. The transport behaviour of different green algae (Chlorella sp., Scenedesmus acuminatus, Desmodesmus communis, and Pediastrum duplex) and algal sized microspheres was tested in flow‐through column experiments with hyporheic sediments. The algal cell transport was directly related to the permeability of the column sediments. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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

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

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.     

Contribution to the study of the Rhyacodrilinae (Tubificidae,Oligochaeta), with description of two new stygobiont species from the alluvial plain of the French upper Rhone,Rhyacodrilus amphigenus,sp. n. and Rhizodriloides phreaticola,g. n., sp. n.

Two new stygobiont species of Rhyacodrilinae (Tubificidae, Oligochaeta), Rhyacodrilus amphigenus, sp. n., and Rhizodriloides phreaticola, g. n., sp. n. are described from the phreatic and hyporheic waters of the alluvial plain of the French upper Rhôone. Further information is given about the distribution of the genera Rhyacodrilus and Rhizodriloides in France and ecological features of several species according to their rate of endemicity and a typological classification of the epigeous and subterranean aquatic environment.