Influence of the mode of macrofauna-mediated bioturbation on the vertical distribution of living benthic foraminifera: First insight from axial tomodensitometry

We investigated the influence of bioturbation by macrofauna on the vertical distribution of living (stained) benthic foraminifera in marine intertidal sediments. We investigated the links between macrofaunal bioturbation and foraminiferal distribution, by sampling from stations situated on a gradient of perturbation by oyster-farming, which has a major effect on benthic faunal assemblages. Sediment cores were collected on the French Atlantic coast, from three intertidal stations: an oyster farm, an area without oysters but affected by oyster biodeposits, and a control station. Axial tomodensitometry (CT-scan) was used for three-dimensional visualization and two-dimensional analysis of the cores. Biogenic structure volumes were quantified and compared between cores. We collected the macrofauna, living foraminifera, shells and gravel from the cores after scanning, to validate image analysis. We did not investigate differences in the biogenic structure volume between cores. However, biogenic structure volume is not necessarily proportional to the extent of bioturbation in a core, given that many biodiffusive activities cannot be detected on CT-scans. Biodiffusors and larger gallery-diffusors were abundant in macrofaunal assemblage at the control station. By contrast, macrofaunal assemblages consisted principally of downward-conveyors at the two stations affected by oyster farming. At the control station, the vertical distribution of biogenic structures mainly built by the biodiffusor Scorbicularia plana and the large gallery-diffusor Hediste diversicolor was significantly correlated with the vertical profiles of living foraminifera in the sediment, whereas vertical distributions of foraminifera and downward-conveyors were not correlated at the station affected by oyster farming. This relationship was probably responsible for the collection of foraminifera in deep sediment layers (> 6 cm below the sediment surface) at the control station. As previously suggested for other species, oxygen diffusion may occur via the burrows built by S. plana and H. diversicolor, potentially increasing oxygen penetration and providing a favorable microhabitat for foraminifera in terms of oxygen levels. By contrast, the absence of living foraminifera below 6 cm at the stations affected by oyster farming was probably associated with a lack of biodiffusor and large gallery-diffusor bioturbation. Our findings suggest that the effect of macrofaunal bioturbation on the vertical distribution of foraminiferal assemblages in sediments depends on the effects of the macrofauna on bioirrigation and sediment oxidation, as deduced by Eh values, rather than on the biogenic structure volume produced by macrofauna. The loss of bioturbator functional diversity due to oyster farming may thus indirectly affect infaunal communities by suppressing favorable microhabitats produced by bioturbation.     

Increased Capacity for Polycyclic Aromatic Hydrocarbon Mineralization in Bioirrigated Coastal Marine Sediments

Bioirrigation of marine sediments by benthic infauna has the potential to increase both the rate and depth of bacterial mineralization of polycyclic aromatic hydrocarbons (PAHs) by recirculating oxygenated bottom water into sediment burrows. Rates of heterotrophic bacterial production and mineralization of PAHs (naphthalene, phenanthrene, and fluoranthene) were measured in sections of sediment cores sampled from stations in San Diego Bay. Data suggest that rates of PAH biodegradation and bacterial heterotrophy were influenced by bioirrigation by benthic infauna. PAH mineralization and heterotrophic production were higher in core sections where sulfide was not detected relative to core sections containing sulfide. Depth-integrated capacity of the upper 17 cm of sediment to mineralize PAHs was 4 to 10 times higher at the station with bioirrigation coefficients that increased with depth. Remedial dredging of sediments to remove contaminant mass (and presumable lower ecological risk) will also remove benthic infauna. Removal of infauna and the subsequent lowering of bioirrigation in surface sediments would be expected to lower the capacity of intrinsic PAH bioremediation. This could cause local increases in ambient PAH concentration and consequently increase the ecological risk at the site and potentially degrade the health of the ecosystem by removing a sink for PAHs.     

High abundance of small zoobenthos around biogenic structures in tidal sediments of the Wadden Sea

On the tidal flats of the island of Sylt (eastern part of the North Sea) the quantity of micro- and meiofauna associated with shoots of seagrass(Zostera noltii), with infaunal bivalves(Macoma balthica), and with tubes and burrows of polychaetes(Pygospio elegans, Pectinaria koreni, Nereis diversicolor, Nereis virens, Arenicola marina) was found to add up to 5 to 33 % of the overall abundance. These structures, taken together, account for 10 to 50 % of the faunal abundance on an average tidal flat at Sylt. The quantitative effect of biogenic structures at the sediment surface (casts and funnels) is small compared to that of tubes and burrows penetrating the anaerobic subsurface layer. In providing stable oxic microenvironments these elite structures frequently bring together more individuals than occur in the entire reducing sediment below surface. Faunal composition of irrigated dwellings of large infauna is different from that of the oxic surface sediment. The common denominator of all elite structures of the subsurface is an oxic halo. Burrows without such a halo are unattractive. There is no evidence that owners of burrows prey on their smaller inmates.     

Assessment of the spatial variability of intertidal benthic communities by axial tomodensitometry: importance of fine-scale heterogeneity

At the water-sediment interface of aquatic ecosystems, the presence of biogenic structures produced by benthic invertebrates strongly affects biogeochemical processes. The quantification of these structures and the assessment of the vertical distribution of fauna are essential for determining the impact of communities in sediments. In the present study, computer axial tomodensitometry (CAT-scan) was used to measure the space occupied by an intertidal community of the St. Lawrence estuary. Three cores were sampled at a site that was considered homogeneous according to surface sediments. The vertical distribution of biogenic structures and gravel were measured in the three cores using CAT-scan; the vertical distribution of fauna was also analysed for each core. The biogenic structures were highest at the water-sediment interface and decreased with depth in the three cores. The number of invertebrates also decreased with depth. We observed similar distributions of biogenic structures in cores 1 and 2. However, fewer biogenic structures were observed below 90 mm in core 3. This result was correlated with a high quantity of gravel from 90 to 140 mm in core 3 whereas the other cores had lower quantities of coarse material. We found relationships among the distributions of biogenic structures, fauna, and sediment characteristics (gravel quantity) that can affect species distribution. The vertical distributions of Macoma balthica, Mya arenaria, Nereis virens, and small-sized gallery-producing species (nematodes and oligochaetes) could also be recorded with the CAT-scan method. Thus, CAT-scan is an excellent tool to determine the fine-scale heterogeneity in the space occupied by benthic invertebrates in sediments.     

Oxygen measurements in the burrows of freshwater insects

1. Thin-tipped micro-electrodes were used to measure oxygen concentrations in the burrows of two common aquatic insects, the mayfly Hexagenia limbata and the alderfly Sialis velata . Both species maintain their surroundings oxygenated by drawing water from above the sediment surface into their tubes. 2. The temporal pattern of oxygen in the burrows differed between the species. The constant high oxygen concentration (>75% of air saturation) measured in the tubes of the mayfly suggest that this animal pumps water almost continuously, which is consistent with its high oxygen requirements. In contrast, oxygen concentration in burrows of the alderfly fluctuated widely over time, suggesting that this animal irrigates only irregularly, probably because it can tolerate short periods of low oxygen concentration in its burrow. 3. The interval between pumping episodes by the alderfly decreased with increasing temperature, a result of increased oxygen consumption by the animal and by sediment at high temperature. 4. Based on the tube dimensions, oxygen penetration depth and animal density in lakes, we estimate that Hexagenia could create an oxic micro-environment equivalent to 3–35% of the volume of the surface oxidized sediment layer created by molecular diffusion. The mosaic of oxic micro-environments created by the burrowing and irrigation of freshwater animals could influence chemical and biological processes in sediments, the fluxes of materials between the sediment and the overlying water column, and the exposure of benthic animals to sedimentary contaminants.     

Benthic–pelagic coupling: a comparison of the community structure of benthic and planktonic heterotrophic protists in shallow inlets of the southern Baltic

1. The taxonomic composition, abundance and biomass of heterotrophic protists (ciliates, heterotrophic flagellates (HF), rhizopods and actinopods) in the sediment and water column of shallow inlets of the Southern Baltic was studied under a variety of environmental conditions during 1996–1997. A shallow, highly eutrophic station and a deeper, less eutrophic station were compared.
2. Community biomass ranged from 0.12 to 0.34 μg C cm?3 in the water column and from 1.5 to 105 μg C cm?3 in the sediment. Heterotrophic protists dominated zooplankton biomass at both stations (73% and 84% mean contribution), while they were of minor importance within the zoobenthos. Expressed per unit area, benthic biomass contributed a significant part (44% and 49%) to the total heterotrophic protistan community at both stations.
3. Although the methodology for counting ciliates and HF was focussed on a high taxonomic resolution, the results reveal some general trends in the distribution of heterotrophic protists: protozooplankton biomass was dominated by flagellates (80% mean biomass contribution) at the shallow station and by ciliates (73% mean biomass contribution) at the deep station. In the benthos at both stations, ciliates were the dominant protozoans, followed by the hitherto little‐studied rhizopods (25% and 35% mean biomass contribution) and flagellates.
4. The degree of benthic–pelagic coupling differed between taxonomic groups. Benthic and pelagic communities of ciliates showed little taxonomic overlap. In contrast, many heterotrophic flagellate species were found both in the benthos and in the pelagic. These benthic–pelagic species contributed significantly to the biomass of HF in the water column. The planktonic rhizopod community consisted of a subset of those species found in the benthos.
5. The abundance of benthic and pelagic protists was positively correlated at the shallow station, but taxonomic data indicate that the direct exchange between benthic and pelagic communities was only partly responsible.     

Do biogenic sulphur compounds in phytodetritus act as cues for deposit-feeder activity? Field experiments and observations on the echiuran worm Maxmuelleria lankesteri

In temperate coastal seas, phytodetritus settling from the spring phytoplankton bloom is a potential food source for benthic deposit-feeders. The ability to exploit this seasonally variable resource could be enhanced by sensitivity to chemical cues signalling its arrival at the seabed. The biogenic sulphur compound dimethylsulphide (DMS), a breakdown product of dimethylsulphonioproprionate (DMSP) produced by some phytoplankton species, is a potential candidate for this role. We investigated the behavioural response of a sedentary surface deposit-feeder, the echiuran worm Maxmuelleria lankesteri, to DMS by observations and manipulative experimentation under natural conditions in a Scottish sea loch. Experimental addition of sediment enriched with DMSP-producing phytoplankton caused no significant increase in either the frequency of feeding by M. lankesteri or the rate of sediment ejection from observed burrows. Naturally occurring (DMSP+DMS) content of surface sediment was low during the winter, then peaked in April before declining in May. There was no consistent relationship between this parameter and rate of sediment ejection from M. lankesteri burrows. The results therefore provide no evidence that M. lankesteri uses DMSP or DMS as a stimulus to increased activity. An observed imbalance between the frequency of surface deposit-feeding and sediment ejection from individual burrows remains unexplained.     

Video-logging for examining biogenic structures in deep heterogeneous subsurface sediments

Video-logging through a transparent tube is described as an alternative method for examining in situ the distribution of biogenic structures within the deep layers of heterogeneous sediments in which undisturbed cores are hard to be obtained. The method comprises four successive phases: the installation of transparent tubes into the sediment, the lowering of a video-camera for scanning sediments through the tube walls, the numerical treatment of images for correcting deformations and obtaining panoramic views of discrete sediment layers, and the mapping of biogenic structures. The method was used successfully to determine the density and depth distribution of galleries made by the oligochaetae Tubifex tubifex (Müller, 1874) within the infiltration bed of a stormwater basin.     

Disturbance,emigration, and refugia: How the mud snail,Ilyanassa obsoleta (Say), affects the habitat distribution of an epifaunal amphipod,Microdeutopus gryllotalpa (Costa)

In the presence of the mud snail Ilyanassa obsoleta (Say), the tubicolous amphipod Microdeutopus gryllotalpa (Costa) emigrates to snail-free sediments, as demonstrated in laboratory and field experiments. Emigration occurs predominantly in the dark when the amphipod is most active. Unlike crevices, the thickness of sediments in which the amphipod is established offers no protection from snail disturbance. Emigration is shown to be caused by the disturbance generated by the snail's plowing and burrowing across the sediment surface, and not a response to a reduction in the shared microfloral food supply. The crawling and burrowing of the smaller mud snail, Hydrobia totteni Morrison, does not disturb Microdeutopus, supporting the hypothesis that relative body sizes affects the ability of bioturbators/burrowers to disturb tube-dwellers. As the burrows of Microdeutopus extend only ≈2 cm below the sediment surface, thick mud layers do not offer any refuge from Ilyanassa. However, very small solid surfaces (≈1–2 mm in relief) to which the amphipods build tubes do provide some protection from Ilyanassa. In soft-sediment benthic communities, such small structures may provide significant refuge for small epifauna and shallow burrowing infauna escaping from small-scale, biogenic disturbance.     

Effects of the burrowing mud shrimp,Upogebia yokoyai,on carbon flow and microbial activity on a tidal flat

Mud shrimps, Upogebia spp., are major constituents of macrobenthic communities in tidal flats in Japan. The impact of Upogebia yokoyai on carbon flow on tidal flats was examined by comparing CO₂ emission rates from plots with and without burrows in the Kurose River estuary, Japan. In situ CO₂ emission rates from plots with burrows were significantly higher than from those without. Laboratory measurements using sediment core samples that excluded respiration of macrobenthic organisms showed similar trends. Although there were no significant differences in grain size distribution, water content, or ignition loss between the sediment cores with and without burrows, oxidation–reduction potential was significantly higher in sediment cores with burrows. Analysis of phospholipid fatty acids (PLFA) indicated that microbial biomass and community structure did not differ significantly between cores with and without burrows. However, microbial respiration activity, as indicated by CO₂ emission rates per total PLFA content, was significantly higher in sediment cores with burrows than in those without. Our results indicate that burrows of U. yokoyai change the physicochemical conditions and increase microbial activity in the sediment, significantly affecting carbon flow in the tidal flat.     

Distribution of sewage indicated by Clostridium perfringens at a deep-water disposal site after cessation of sewage disposal.

Clostridium perfringens, a marker of domestic sewage contamination, was enumerated in sediment samples obtained from the vicinity of the 106-Mile Site 1 month and 1 year after cessation of sewage disposal at this site. C. perfringens counts in sediments collected at the disposal site and from stations 26 nautical miles (ca. 48 km) and 50 nautical miles (ca. 92 km) to the southwest of the site were, in general, more than 10-fold higher than counts from an uncontaminated reference site. C. perfringens counts at the disposal site were not significantly different between 1992 and 1993, suggesting that sewage sludge had remained in the benthic environment at this site. At stations where C. perfringens counts were elevated (i.e., stations other than the reference station), counts were generally higher in the top 1 cm and decreased down to 5 cm. In some cases, C. perfringens counts in the bottom 4 or 5 cm showed a trend of higher counts in 1993 than in 1992, suggesting bioturbation. We conclude that widespread sludge contamination of the benthic environment has persisted for at least 1 year after cessation of ocean sewage disposal at the 106-Mile Site.     

The effect of benthic algae on phosphorus exchange between sediment and overlying water in shallow lakes: a microcosm study using 32P as a tracer

Sediments are of key importance in determining the nutrient levels of water in shallow lakes as they can act as either source or sink for phosphorus (P) depending on environmental conditions, sediment characteristics, and external nutrient loading. We examined the role of benthic algae in the P cycling between sediment and overlying water in experiments using ³²P as a tracer. Sediment and water samples were collected from Huizhou West Lake, a shallow, eutrophic waterbody located in Huizhou City, South China. Laboratory cultured benthic algae were transferred to cover the sediment core in tubes. When ³²P was added to the water in experimental tubes containing sediment cores with and without benthic algae, ³²P activity after 48 h was significantly lower in the tubes with algae, indicating that benthic algae removed P from the overlying water. When the tracer was injected into the sediment, ³²P activity in the water overlying sediment with benthic algae was substantially lower than in tubes with naked sediment, suggesting that benthic algae reduce the release of sediment P. Oxygen levels were significantly higher in the upper 3 mm of the sediments covered by benthic algae; thus, we hypothesized that oxygen produced by the algae helps inhibit the release of P from the sediment. Our study demonstrates that benthic algae are capable of reducing P levels in water overlying the sediment, suggesting that loss of benthic algae during eutrophication triggered by impoverished light conditions may accelerate the shift in shallow lakes from a clear water to a turbid state.     

The distribution of juvenile benthic invertebrates in an arctic glacial fjord

The distribution of juvenile benthic invertebrates with respect to their abiotic environment was investigated at four locations along a gradient in Kongsfjorden (79°N, 12°E), Spitsbergen, in September 1997. Heavy discharge of inorganic sediments released by the three glaciers was expected to be one of the main structuring factors of benthic communities in the fjord. Juveniles of 76 species/families are identified. Total abundance increases with distance from the fjord head due to the dominance of opportunistic polychaetes, while diversity is highest close to the glacier. Juvenile polychaetes are the most abundant taxon over all stations (92%), followed by bivalves, crustaceans and others. Polychaetes also provide most species (55%) and are the dominant taxon at every station. Their abundance is highly correlated with total organic carbon, indicating that most of them are deposit feeders. Young crustaceans rapidly decrease towards the fjord mouth, probably due to stronger bottom currents. Even though young oligochaetes are exclusively found near the glacier, no species seem to be well adapted to the high sedimentation rate close to the glacier. Juvenile suspension-feeding bivalves seem to be less disturbed by glacial discharge but appear to be more vulnerable to currents on the more exposed sites. The importance of deposit feeders and carnivores increases towards the outer stations. Non-metric multidimensional scaling confirms the distinct gradient in community composition along the fjord. Coupling the biotic data to abiotic factors (depth, bottom-water salinity, bottom-water temperature, sediment grain size and sedimentation rate) using canonical correspondence analysis revealed that hydrographic factors are more responsible for the structuring of the benthic juvenile community at the shallow stations close to the glacier (except the station directly at the glacier). At the outer deeper stations, sediment grain size and related properties may play a more important role.     

Response of invertebrates to lotic disturbance: is the hyporheic zone a patchy refugium?

1. Natural experiments, in the form of disturbance from spates, were used to study the resistance and resilience of interstitial communities. Investigations were conducted in a by-passed section of the Rhône River characterized by an artificial hydrology with frequent spates separated by regular minimum discharge of 30 m³ s^–1. 2. Three areas of a bar were studied, upwellings at the head of the bar (stations 1 and 2), and downwelling at the tail of the bar (station 3). In the head of the bar the substratum was characterized by stable cobbles, while mobile gravels dominated in the tail of the bar. At each station, samples were derived from four depths (0.5, 1.0, 1.5 and 2.0 m below the surface of the substratum). Fifteen spates occurred during the study period whose peak discharge ranged from 50 to 1640 m³ s^–1. Temporal variations of the fauna were studied by comparing the spate effect observed 1 day (resistance), 7 days (resilience) and 17 days after the spate. Within-class correspondence analysis was used to compare the temporal variability of the fauna within each class {station/depth}. 3. The fauna differed markedly between the three stations, and the relative density of stygobionts (i.e. hypogean fauna) decreased from 55% at station 1 to 4% at station 3. The spatio-temporal variability increased dramatically from station 1 to station 3. 4. The results suggest that the hyporheic zone acts as a patchy refugium: the stations were more or less active refugial zones, depending on hydrology (upwelling or downwelling), substratum stability and spate amplitude. 5. The downwelling station was the main refugium area for benthic taxa. Important migrations of benthic groups (e.g. Gammarus, Cladocera) or hyporheic taxa (e.g. Cyclopoida and Harpacticoida) were observed deep into the sediment (2 m). Vertical movements of stygobionts (Niphargus, Niphargopsis) were also observed at high amplitude spates. These movements were very important (great numbers of individuals migrated) at low and medium magnitude spates, but were unimportant at high discharge, when the threshold of sediment instability was exceeded. In this case the substratum became mobile and induced drift of benthic organisms. 6. Conversely, in the upwelling stable stations, accumulation was less important (lower number of species and lower densities) but more constant with increasing discharge, suggesting that substratum stability is also a key factor. 7. Generally recovery was rapid at all stations (within 7 days) but no relationships were found between resilience (rate of recovery) and the amplitude of spates.     

Short-term variations in the fluxes and composition of seston in near-bottom traps in the southern North Sea

In this study we attempted to measure the potential food availability for macrobenthic interface feeders using a new device (sediment recorder) which traps bed load particles at a height of 10 cm above the sea floor. The recorder is positioned flat on the seabed and alternately exposes a shallow collector mounted in a smooth surface mimicking the sediment surface. The samples were analyzed for their content of chlorophyll a (Chl) and dry weight (DM). We compared the sediment recorder data with the results obtained with a sediment trap suspended 3.2 m above the sea floor. Measurements were carried out in spring and fall at two stations in the southern North Sea with different sediment type, maximum current velocities and biomass (sand, station B vs silty sand, station F). The recorder deployments in November yielded higher Chl fluxes and a higher quality of particles (Chl:DM ratio) than the trap at both stations. Unlike the trap samples, in which Chl and DM were always positively and strongly correlated, maximum Chl and DM fluxes in the recorder samples were out of phase, particularly at station B, pointing to a highly variable quality during the tidal cycle. Unexpectedly, Chl fluxes and Chl:DM ratios in the recorder samples at station B were higher than at station F in both seasons, although the benthic biomass at the latter station was five times higher. The highest Chl fluxes and particle quality (Chl:DM) at station B were measured during slack tides when interface feeding would be least hampered by drag forces. Station B is nevertheless characterized by a low macrobenthic biomass and a scarcity of interface feeders, specifically of suspension-feeding bivalves. Possible reasons for this apparent discrepancy are discussed. Electronic Publication     

Diagenetic stoichiometry and benthic nutrient fluxes at the sediment–water interface of Lake Illawarra, Australia

Benthic flux measurements of O₂, TCO₂ and inorganic nutrients were made at three stations (seagrass beds, shallow bare sand and deep mud) in Lake Illawarra (Australia) to compare the characteristics of diagenesis and benthic biogeochemical processes for different primary producers (seagrass or microphytobenthos, (MPB)) and/or sediment types (sand or mud).Seagrass beds exhibited the highest gross primary productivity while the lowest rates occurred at the deep mud station. At the shallow bare sand station only, the gross primary production (GPP) and respiration (R) were balanced, while at the other two stations, R exceeded GPP by as much as 2 fold, indicating more organic carbon was decomposed than produced at the time of sampling. In general, nutrient fluxes displayed typical diurnal variation.Organic carbon oxidation scenarios, evaluated by either calcium carbonate dissolution or sulfate reduction models, indicated that both models can represent organic matter mineralization. The difference of estimated total carbon oxidized in this lake using the two models was small, ranging from 0.2% at deep mud station to maximum of 21% at seagrass station. In addition, N₂ flux rates (net denitrification), estimated using carbon and nitrogen stoichiometry, were of similar magnitude as the rates estimated using LOICZ budget modeling or measured using the N₂/Ar technique.Finally, a comparison of calculated diffusive fluxes and measured fluxes using incubation cores indicated that the results were of similar magnitude at the deep mud station, but the incubation cores fluxes were much higher than the calculated diffusive fluxes at the other two stations. This may have been caused by bioturbation or bioirrigation.     

In situ biochemical and bacterial variation of sediments enriched with mussel biodeposits

In order to estimate the qualitative variation of sedimented mussel deposits, biochemical and microbial measurements were undertaken after a sediment enrichment with fresh faeces and pseudo-faeces collected in a mussel farming area (Carteau, Gulf of Fos).During two months, cores sampled were investigated at three stations: a first station enriched with mussel deposits, a second considered as a reference station without mussel deposits and occasionally a third corresponding to a continuously enriched sediment under a rope culture.Carbohydrate, organic carbon and nitrogen contents in the sediments gave evidence of a short term variation after sedimentation.Bacterial production increased rapidly at the enriched station and returned to its initial level 8 days later. At this station, exoglucosidasic activity of bacteria, low at the beginning, presented a maximum two weeks after enrichment whereas exoproteolytic activity, which was high in the biodeposits, decreased in the course of the first week. These exoenzymatic activities were significantly higher at the enriched station than at the reference stations. Carbohydrate measurements were in agreement with these results.Degradation rate of biodeposits is discussed on the basis of exoenzymatic activity, organic carbon, nitrogen and free amino acid content at the three stations.     

Major contribution of sulfide-derived sulfur to the benthic food web in a large freshwater lake

In freshwater systems, contributions of chemosynthetic products by sulfur-oxidizing bacteria in sediments as nutritional resources in benthic food webs remain unclear, even though chemosynthetic products might be an important nutritional resource for benthic food webs in deep-sea hydrothermal vents and shallow marine systems. To study geochemical aspects of this trophic pathway, we sampled sediment cores and benthic animals at two sites (90 and 50 m water depths) in the largest freshwater (mesotrophic) lake in Japan: Lake Biwa. Stable carbon, nitrogen, and sulfur isotopes of the sediments and animals were measured to elucidate the sulfur nutritional resources for the benthic food web precisely by calculating the contributions of the incorporation of sulfide-derived sulfur to the biomass and of the biogeochemical sulfur cycle supporting the sulfur nutritional resource. The recovered sediment cores showed increases in ³⁴S-depleted sulfide at 5 cm sediment depth and showed low sulfide concentration with high δ³⁴S in deeper layers, suggesting an association of microbial activities with sulfate reduction and sulfide oxidation in the sediments. The sulfur-oxidizing bacteria may contribute to benthic animal biomass. Calculations based on the biomass, sulfur content, and contribution to sulfide-derived sulfur of each animal comprising the benthic food web revealed that 58%–67% of the total biomass sulfur in the benthic food web of Lake Biwa is occupied by sulfide-derived sulfur. Such a large contribution implies that the chemosynthetic products of sulfur-oxidizing bacteria are important nutritional resources supporting benthic food webs in the lake ecosystems, at least in terms of sulfur. The results present a new trophic pathway for sulfur that has been overlooked in lake ecosystems with low-sulfate concentrations.     

Seasonal Vertical Migration and Aestivation of <Emphasis Type="Italic">Rhyacodrilus hiemalis</Emphasis> (Tubificidae,Clitellata) in the Sediment of Lake Biwa,Japan

The vertical distribution of the tubificid worm Rhyacodrilus hiemalis Ohtaka, the numerically dominant species of oligochaete in the littoral of Lake Biwa, was studied with special reference to seasonal vertical migration in the lake sediment. Monthly collections of lake sediment cores were made using PVC tubes. Core sections of sample sediments ranged from 76 to 117 cm. The vertical distribution of the worms showed no diurnal variation; therefore diel vertical migration was not evident. Seasonal downward migration started in April, and upward migration started in October. From December to March, almost all worms remained in the near-surface sediment layer (surface to 30 cm deep), while from July to September almost all worms remained deeper than 30 cm. However, few individuals migrated deeper than 90 cm. No discontinuous layers were found in grain size composition, water content, loss on ignition, particulate carbon, nitrogen or phosphorus. In deep sediment there was no free oxygen, as evidenced by negative ORP values. For 4 months in summer, R. hiemalis aestivated, probably utilizing anaerobic respiration. It appeared that R. hiemalis moved deeper in the sediment in response to sediment temperature, because sediment temperatures in the deep layers seemed to converge at around 20–21 °C in the summer months. The life history traits of seasonal vertical migration and summer aestivation perhaps arose as an adaptation to the climatic conditions accompanying the geographical origin of R. hiemalis, and they also serve to minimize predation risk during summer when most invertebrate predatory fishes are active.     

Living benthic foraminifera of the Okhotsk Sea: Faunal composition,standing stocks and microhabitats

Live (Rose Bengal stained) benthic foraminifera were investigated in surface sediment samples from the Okhotsk Sea to reveal the relationship between faunal characteristics and environmental parameters. Live benthic foraminifera were quantified in the size fraction > 125 µm in the upper 8 cm of replicate sediment cores, recovered with a multicorer at five stations along the Sakhalin margin, and at three stations on the southwestern Kamchatka slope. The stations are from water depths between 625 to 1752 m, located close or within the present Okhotsk Sea oxygen minimum zone, with oxygen levels between 0.3 and 1.5 ml l^- 1. At the high-productivity and ice-free Kamchatka stations, live benthic foraminifera are characterized by maximal standing stocks (about 1700-3700 individuals per 50 cm²), strong dominance of calcareous species (up to 87-91% of total live faunas), and maximal habitat depths (down to 5.2-6.7 cm depth). Vertical distributions of total faunal abundances exhibit a clear subsurface maximum in sediments. At the Sakhalin stations, which are seasonally ice-covered and less productive, live benthic foraminifera show lower standing stocks (about 200-1100 individuals per 50 cm²), lower abundance of calcareous species (10-64% of total live faunas), and shallower habitat depths (down to 2.5-5.4 cm depth). Faunal vertical distributions are characterized by maximum in the uppermost surface sediments. It is suggested that 1) lower and strongly seasonal organic matter flux, caused by the seasonal sea ice cover and seasonal upwelling, 2) lower bottom water oxygenation (0.3-1.1 ml l^- 1), and 3) more pronounced influence of carbonate undersaturated bottom water along the Sakhalin margin are the main factors responsible for the observed faunal differences. According to species downcore distributions and average living depths, common calcareous species were identified as preferentially shallow, intermediate and deep infaunal. Foraminiferal microhabitat occupation correlates with the organic matter flux and the depth of the oxygenated layer in sediments.