Intercalibration of benthic flux chambers I. Accuracy of flux measurements and influence of chamber hydrodynamics

The hydrodynamic properties and the capability to measure sediment-water solute fluxes, at assumed steady state conditions, were compared for three radically different benthic chamber designs: the “Microcosm”, the “Mississippi” and the “Göteborg” chambers. The hydrodynamic properties were characterized by mounting a PVC bottom in each chamber and measuring mixing time, diffusive boundary layer thickness (DBL thickness) shear velocity (u∗), and total pressure created by the water mixing. The Microcosm had the most even distribution of DBL thickness and u∗, but the highest differential pressure at high water mixing rates. The Mississippi chamber had low differential pressures at high u∗. The Göteborg chamber was in between the two others regarding these properties. DBL thickness and u∗ were found to correlate according to the following empirical formula: DBL=76.18(u∗)−0.933. Multiple flux incubations with replicates of each of the chamber types were carried out on homogenized, macrofauna-free sediments in four tanks. The degree of homogeneity was determined by calculating solute fluxes (of oxygen, silicate, phosphate and ammonium) from porewater profiles and by sampling for porosity, organic carbon and meiofauna. All these results, except meiofauna, indicated that there were no significant horizontal variations within the sediment in any of the parallel incubation experiments. The statistical evaluations also suggested that the occasional variations in meiofauna abundance did not have any influence on the measured solute fluxes. Forty-three microelectrode profiles of oxygen in the DBL and porewater were evaluated with four different procedures to calculate diffusive fluxes. The procedure presented by Berg, Risgaard-Petersen and Rysgaard, 1989 [Limnol. Oceanogr. 43, 1500] was found to be superior because of its ability to fit measured profiles accurately, and because it takes into consideration vertical zonation with different oxygen consumption rates in the sediment. During the flux incubations, the mixing in the chambers was replicated ranging from slow mixing to just noticeable sediment resuspension. In the “hydrodynamic characterizations” these mixing rates corresponded to average DBL thickness from 120 to 550 μm, to u∗ from 0.12 to 0.68 cm/s, and to differential pressures from 0-3 Pa. Although not directly transferable, since the incubations were done on a “real” sediment with a rougher surface while in the characterizations a PVC plate simulated the sediments surface, these data give ideas about the prevailing hydrodynamic condition in the chambers during the incubations. The variations in water mixing did not generate statistically significant differences between the chamber types for any of the measured fluxes of oxygen or nutrients. Consequently it can be concluded that, for these non-permeable sediments and so long as appropriate water mixing (within the ranges given above) is maintained, the type of stirring mechanism and chamber design used were not critical for the magnitude of the measured fluxes. The average measured oxygen flux was 11.2 ± 2.7 (from 40 incubations), while the diffusive flux calculated (from 43 profiles using the Berg et al., 1989 [Limnol. Oceanogr. 43, 1500] procedure) was 11.1 ± 3.0 mmol m⁻² day⁻¹. This strongly suggests that accurate oxygen flux measurements were obtained with the three types of benthic chambers used and that the oxygen uptake is diffusive.     

Diffusive and total oxygen uptake of deep-sea sediments in the eastern South Atlantic Ocean:in situ and laboratory measurements

Total O₂ uptake rates were measured by the benthic flux chamber lander ELINOR, and O₂ microprofiles were measured by the profiling lander PROFILUR in the eastern South Atlantic. Diffusive O₂ fluxes through the diffusive boundary layer and the depth distribution of O₂ consumption rates within the sediment were calculated from the obtained microprofiles. The depth integrated O₂ consumption rate agreed closely with the diffusive O₂ uptake at all stations. Total O₂ uptake was 1.2–4.2 times the diffusive O₂ uptake, and the difference correlated with the abundance of macrofauna in the sediment. Diffusive O₂ uptake and O₂-penetration depths correlated with the organic content of the sediments and exhibited an inverse correlation with water depth. Total and diffusive rates of in situ O₂ uptake were higher than previously published data for shelf and abyssal sediments in the Atlantic, but were comparable to rates from upwelling areas in the eastern Pacific. Laboratory measurements on recovered sediment cores showed lower O₂ penetration depths and higher diffusive uptake rates than in situ measurements. The differences increased with increasing water depth. We primarily ascribe this compression of O₂ profiles to a transiently increased temperature during recovery and enhanced microbial activity in decompressed sediment cores. Total O₂ uptake rates measured in the laboratory on macrofauna-rich stations were, in contrast, lower than those measured in situ because of underrepresentation and disturbance of the macrofauna.     

Benthic Nutrient Recycling in Port Phillip Bay, Australia

Benthic chamber measurements of the reactants and products involved with biogenic matter remineralization (oxygen, ammonium, nitrate, nitrite, phosphate, silicate, TCO₂and alkalinity) were used to define solute exchange rates between the sediment and overlying water column of Port Phillip Bay, Australia. Measurements at various sites throughout the bay, conducted during the summers of 1994 and 1995, indicate that the variability in flux values within a site is comparable to year-to-year variability (±50%). Four regions of the bay were distinguished by sediment properties and the northern region was identified as having 3–30 times greater nutrient regeneration rates than the other regions. Benthic recycling accounted for 63 and 72% of the annualized N and P input, respectively, to the entire bay as determined by summing benthic, dissolved riverine, atmospheric and dissolved effluent sources. However, bay-wide sedimentary denitrification accounted for a loss of 63% of the potentially recyclable N. This fraction is higher than many other coastal regions with comparable carbon loading. Denitrification efficiency is apparently not enhanced by benthic productivity nor by bio-irrigation. The rate of bio-irrigation is negatively correlated with denitrification efficiency. Bio-irrigation was studied using radon-222 and CsCl spike injection chamber measurements. Radon fluxes from sediments in Port Phillip Bay were enhanced over the diffusive flux by 3–16 times. The modelled rate of loss of Cs from chamber water was positively correlated with radon flux enhancement results. Both methods identify regions within Port Phillip Bay that have particularly high rates of non-diffusive pore-water overlying water solute exchange.     

Influence of Particle Mixing on Vertical Profiles of Chlorophyll a and Bacterial Biomass in Sediments of the German Bight, Oyster Ground and Dogger Bank (North Sea)

In May and September 1999 11 stations were sampled in the southern and central North Sea, located in the German Bight, eastern Oyster Ground and Dogger Bank. The study focused on the influence of particle mixing on transport of chlorophyll a to deeper sediment layers and vertical bacterial distribution (max. DEPTH=10 cm). The sampling stations were chosen to reflect a gradient in environmental conditions in the North Sea. The sampling stations differed in respect to redox potential (eH up to −243 mV in the German Bight and up to 274 mV in the offshore regions), silt content (up to 54% in the German Bight and 0·34% at the northern Dogger Bank) and different proportion of fresh organic material on total organic matter content (C/N ratios ranging from 9·27 in the German Bight up to 1·72 in the offshore sediments). Although bacterial densities (8·55×10⁹ g⁻¹in the German Bight up to 0·35×10⁹ g⁻¹in offshore sediments) were significantly correlated to chlorophyll a content in the sediment (P<0·01), inconsistencies in the temporal pattern of both variables in the surficial sediment layer suggested, that the dynamics of bacterial densities is generally controlled by food supply but also by other variables. The chlorophyll a content in the surficial sediments of the German Bight (up to 1·84 μg g⁻¹) was significantly higher than in the Oyster Ground (up to 0·58 μg g⁻¹) and the Dogger Bank area (up to 0·68 μg g⁻¹). With increasing chlorophyll a input to the benthic realm a subsequent enhanced burial of this compound into deeper sediment layers was expected either by biological (bioturbation) or by physical sediment mixing. However, the vertical profile of chlorophyll a decreased steeply in the sediments of the German Bight. Contrary, subsurface peaks were measured in the offshore areas. It was concluded from these results, that the vertical distribution of organic matter in sediments is less limited by the quantitative input from the water column but concomitant with particle mixing itself. The extent and possible mechanisms of particle mixing in the different study areas in relation to specific environmental factors is discussed.     

Evaluation of the N2 flux approach for measuring sediment denitrification

Direct gas chromatographic measurement of denitrification rates via N₂ fluxes from aquatic sediments can avoid some of the artifacts and complexities associated with indirect approaches and tracer techniques. However, measurement protocols have typically been determined based upon initial results or previous studies. We present a process-level study and simulation model for evaluating and optimizing N₂ gas flux approaches in closed chamber incubations. Experimental manipulations and simulations of both artificial and natural sediments were used to conduct sensitivity analyses of key design parameters in N₂ flux measurements. Experimental results indicated that depletion of labile organic matter during the long incubations required by common protocols (for diffusive off-gassing of porewater N₂) may result in underestimates of denitrification rates in some systems. Simulations showed that the required incubation time was primarily a function of sediment thickness. The best approach found to minimize incubation time and reduce errors was to select the minimum sediment thickness necessary to include the entire depth distribution of nitrification–denitrification for a particular sediment system. Attempts to increase measurement sensitivity and shorten incubation times by reducing the headspace thickness to 1–2 cm generally cause denitrification to be underestimated by 3–13% for gas headspaces, and up to 80% for water headspaces. However, errors were negligible with gas and water headspace thicknesses of 10 cm and 15 cm, respectively. Anaerobic cores to control for non-denitrification N₂ fluxes shortened incubation time, but introduced artifacts in sediments with extensive macrofaunal irrigation.     

Validation of the three-dimensional ECOHAM model in the German Bight for 2004 including population dynamics of Pseudocalanus elongatus

A three-dimensional ecosystem model for the North Sea which includes competition between Pseudocalanus elongatus and the rest of the zooplankton biomass was applied to describe the seasonal cycle of zooplankton in 2003–2004. The paper presents the comparison of simulated stage-resolved abundances with copepod counts at several stations in the German Bight during the GLOBEC-Germany project from February to October 2004. A validation of influential state variables gives confidence that the model is able to calculate reliably the stage development and abundances of P. elongatus as well as the range of bulk zooplankton biomass, and thus the ratio of population biomass to total biomass. In the German Bight, the population is below 20% in spring. The ratio increases up to 50% during summer. The number of generations was estimated from peaks in egg abundance to about 4–8 generations of P. elongatus in the southern North Sea. A mean of four generations per year were estimated in the central North Sea, six to eight generations northwest of the Dogger Bank (tails end) and five generations in the German Bight.     

Oxygenation episodes on the continental shelf of central Peru: Remote forcing and benthic ecosystem response

The interplay between the oxygen minimum zone and remotely-forced oxygenation episodes determines the fate of the benthic subsystem off the Central Peruvian coast. We analyzed a 12 year monthly time-series of oceanographic and benthic parameters at 94 m depth off Callao, Central Peru (12°S), to analyze: (i) near-bottom oxygen level on the continental shelf in relation to dynamic height on the equator (095°W); and (ii) benthic ecosystem responses to oxygen change (macrobiotic infauna, meiofauna, and sulphide-oxidizing bacteria, Thioploca spp.). Shelf oxygenation episodes occurred after equatorial dynamic height increases one month before, consistent with the propagation of coastal trapped waves. Several but not all of these episodes occurred during El Niños. The benthic biota responded to oxygenation episodes by undergoing succession through three major ecological states. Under strong oxygen deficiency or anoxia, the sediments were nearly defaunated of macro-invertebrates and Thioploca was scarce, such that nematode biomass dominated the macro- and meiobiotas. When frequency of oxygenation events reduced the periods of anoxia, but the prevailing oxygen range was 10–20 μmol L⁻¹, mats of Thioploca formed and dominated the biomass. Finally, with frequent and intense (>40 μmol L⁻¹) oxygenation, the sediments were colonized by macrofauna, which then dominated biomass. The Thioploca state evolved during the 2002–2003 weak EN, while the macrofauna state was developed during the onset of the strong1997–1998 EN. Repeated episodes of strong oxygen deficiency during the summer of 2004, in parallel with the occurrence of red tides in surface waters, resulted in the collapse of Thioploca mats and development of the Nematode state. Ecological interactions may affect persistence or the transition between benthic ecosystem states.     

Biogeochemistry in highly reduced mussel farm sediments during macrofaunal recolonization by Amphiura filiformis and Nephtys sp.

Mussel farming is considered a viable means for reducing coastal eutrophication. This study assessed the importance of bioturbation by recolonizing fauna for benthic solute fluxes and porewater distributions in manipulated mussel farm sediments. Three consecutive time-series flux incubations were performed during an experimental period of three weeks in sieved farm sediment treated with the brittle star Amphiura filiformis and the polychaete Nephtys sp. The functional behavior of Nephtys sp. and interactions between Nephtys sp. and the spontaneously colonizing spionid Malacoceros fuliginosus determined the biogeochemical response in the Nephtys sp. treatment. For example, the oxic zone was restricted and benthic nitrate and silicate fluxes were reduced compared to the brittle star treatment. A. filiformis seemed to enhance the bioadvective solute transport, although an increased supply of oxygen was due to the highly reducing conditions of the sediment mainly seen as secondary effects related to porewater distributions and benthic nutrient fluxes.     

Effect of emersion and immersion on the porewater nutrient dynamics of an intertidal sandflat in Tokyo Bay

Porewater nutrient dynamics during emersion and immersion were investigated during different seasons in a eutrophic intertidal sandflat of Tokyo Bay, Japan, to elucidate the role of emersion and immersion in solute transport and microbial processes. The water content in the surface sediment did not change significantly following emersion, suggesting that advective solute transport caused by water table fluctuation was negligible. The rate of change in nitrate concentration in the top 10 mm of sediments ranged from −6.6 to 4.8 μmol N l⁻¹ bulk sed. h⁻¹ during the whole period of emersion. Steep nutrient concentration gradients in the surface sediment generated diffusive flux of nutrients directed downwards into deeper sediments, which greatly contributed to the observed rates of change in porewater nutrient concentration for several cases. Microbial nitrate reduction within the subsurface sediment appeared to be strongly supported by the downward diffusive flux of nitrate from the surface sediment. The stimulation of estimated nitrate production rate in the subsurface layer in proportion to the emersion time indicates that oxygenation due to emersion caused changes in the sediment redox environment and affected the nitrification and/or nitrate reduction rates. The nitrate and soluble reactive phosphorus pools in the top 10 mm of sediment decreased markedly during immersion (up to 68% for nitrate and up to 44% for soluble reactive phosphorus), however, this result could not be solely explained by molecular diffusion.     

Nutrient fluxes from deep sediment support nutrient budget in the oligotrophic waters of the Gulf of Aqaba

The role of deep sediment in supporting nutrient budget in the Gulf of Aqaba has been investigated by estimating the flux of inorganic nitrogen, phosphate and silicate. Fluxes were calculated directly by pore water profiles and indirectly by chamber incubations carried out onboard the RV Meteor cruise. The results showed that maximum potential fluxes calculated by chamber incubations were higher than those calculated by porewater profiles for all nutrients (6.4–28.5 fold). This has been attributed to the additional flux due to bioturbation and flux from advective porewater exchange in the case of chamber incubation, while porewater fluxes represent diffusive ones. Using a rough estimation considering flux results in addition to the sediment area and water mass of the Gulf of Aqaba, we estimate that 3.3 × 10⁵, 6.4 × 10⁴ and 6.5 × 10⁶ kg year⁻¹ of inorganic nitrogen, phosphate and silicate respectively are effused from deep sediments to the water column. This quanitity would certainly support the primary productivity in the oligotrophic water in the Gulf of Aqaba.     

Phytopigments and fatty acids as molecular markers for the quality of near-bottom particulate organic matter in the North Sea

In February, May and August 1994, four stations in the North Sea (viz. at the Broad Fourteens, Frisian Front, German Bight and Skagerrak) were visited to sample near-bottom particulate organic matter. Samples, taken by means of a pump, a sediment trap and a sediment recorder, were analysed on organic carbon, total nitrogen, phytopigments and fatty acids. These molecular markers were used to describe the nature and quality of the organic particles in the near-bottom water. Principal component analysis showed chlorophyll a, phaeopigments and fatty acids to be useful markers for the quality of organic matter and yield complementary information.The quality of the near-bottom particles appeared to be related to the local hydrography and depositional circumstances. The Broad Fourteens station, a non-depositional sandy site along the Dutch coast, showed organic particles to be relatively fresh, little influenced by resuspended sedimentary material. Near-bottom organic particles on this site contained relatively high shares of chlorophyll a and polyunsaturated fatty acids, characteristic of algal matter. On the other hand the particulate organic material on the two depositional locations, the Frisian Front and the German Bight stations, was influenced by resuspension of sedimentary organic particles poor in pigments and fatty acids. Amounts of carbon trapped in the near-bottom environment at the Skagerrak station were lower than expected from the literature.     

Deep penetrating benthic oxygen profiles measured in situ by oxygen optodes

For the first time in situ, deep penetrating O₂ profiles were measured in abyssal sediments in the western South Atlantic. Construction of deep penetrating O₂ optodes and adaptation to a benthic profiling lander are described. The opto-chemical oxygen sensors allow measurements to a depth of 55 cm in marine sediments. A vertical resolution of 0.5 cm was used to determine the O₂ dynamics in those oligotrophic deep sea sediments; the oxygen concentration across the sediment water interface was measured with a resolution of 100 μm. Oxygen penetration depth (OPD), diffusive oxygen uptake (DOU) and oxygen consumption rates were determined at four stations north of the Amazon fan and one at the Mid-Atlantic Ridge. Diffusive oxygen uptake rates ranged from 0.1 to 0.9 mmol m⁻² d⁻¹; the oxygen penetration depth ranged from 8 to 26 cm. Carbon consumption rates calculated from the diffusive oxygen uptake rates were in the range of 0.3–3.0 g C m⁻² a⁻¹. Comparison between in situ and laboratory DOU and OPD measurements confirmed previous findings that core recovery and warming have strong effects on the oxygen dynamics in deep sea sediments. Laboratory measurements yielded a decrease of 50–75% in OPD and consequently an increase in DOU by 1.5 and 18-times. Deep penetrating oxygen optodes provide a new tool to accurately determine oxygen dynamics (and thereby calculate carbon mineralization rates) in oligotrophic sediments. However, oxygen optodes as used in this study do not resolve the diffusive boundary layer (DBL). The data show that deep penetrating O₂ optodes in combination with high-resolution O₂ microelectrodes give a complete picture of the oxygen dynamics, including the DBL, in deep sea sediments.     

Distribution of green algal mats throughout shallow soft bottoms of the Swedish Skagerrak archipelago in relation to nutrient sources and wave exposure

Distribution and biomass of green algal mats were studied in marine shallow (0–1 m) soft-bottom areas on the Swedish west coast from 1994 to 1996, by combining aerial photography surveys with ground truth sampling. Filamentous green algae, dominated by species of the genera Cladophora and Enteromorpha, were generally present throughout the study area during July and August, and largely absent in late April and early May. These algae occurred at 60 to 90% of the locations investigated during the summer, and were estimated to cover between 30 and 50% of the total area of shallow soft bottoms of the Swedish Skagerrak archipelago. The distributional patterns were similar during the three years of the investigation and appeared unrelated to annual local nutrient inputs from point sources and river discharge. We postulate that the apparent lack of such a relationship is due to an altered state of nutrient dynamics throughout the archipelago. Mechanisms are likely to involve long-term, diffuse elevations in nutrient levels in coastal waters of the Skagerrak and the Kattegat over several decades leading to current eutrophic conditions, exceeding nutrient requirements for abundant filamentous algal growth. Patterns of algal abundance in our study were largely related to physical factors such as exposure to wind, waves and water exchange under conditions where nutrient loads among embayments seemed to be unlimited. Further, our results show that sediments covered by algal mats had higher carbon and nitrogen contents than unvegetated sediments. We hypothesise that sustained high nutrient loads, manifested in extensive biomass of filamentous algae during summer months, are re-mineralised via decay and sedimentation in the benthic realm. Hence, accumulated carbon and nutrients in the sediment could, in turn, constitute the basic pool for future algal mat production overlying soft bottoms in areas where tidal exchange is limited.     

Nutrient exchange across the sediment-water interface in the Potomac River estuary

The flux of ammonia, phosphate, silica and radon-222 from Potomac tidal river and estuary sediments is controlled by processes occurring at the sediment-water interface and within surficial sediment. Calculated diffusive fluxes range between 0·6 and 6·5 mmol m^?2 day^?1 for ammonia, 0·020 and 0·30 mmol m^?2 day^?1 for phosphate, and 1·3 and 3·8 mmol m^?2 day^?1 for silica. Measured in situ fluxes range between 1 and 21 mmol m^?2 day^?1 for ammonia, 0·1 and 2·0 mmol m^?2 day^?1 for phosphate, and 2 and 19 mmol m^?2 day^?1 for silica. The ratio of in situ fluxes to diffusive fluxes (flux enhancement) varied between 1·6 and 5·2 in the tidal river, between 2·0 and 20 in the transition zone, and from 1·3 to 5·1 in the lower estuary. The large flux enhancements from transition zone sediments are attributed to macrofaunal irrigation. Nutrient flux enhancements are correlated with radon flux enhancements, suggesting that fluxes may originate from a common region and that nutrients are regenerated within the upper 10–20 cm of the sediment column.The low fluxes of phosphate from tidal viver sediments reflect the control benthic sediment exerts on phosphorus through sorption by sedimentary iron oxyhydroxides. In the tidal river, benthic fluxes of ammonia and phosphate equal one-half and one-third of the nutrient input of the Blue Plains sewage treatment plant. In the tidal Potomac River, benthic sediment regeneration supplies a significant fraction of the nutrients utilized by primary producers in the water column during the summer months.     

High occurrence of the elasipodid holothurian Penilpidia ludwigi (von Marenzeller, 1893) in bathyal sediment traps moored in a western Mediterranean submarine canyon

One hundred and fifty specimens of the elasipod holothurian Penilpidia ludwigi (von Marenzeller, 1893) were collected in sediment traps moored near the seabed in the La Fonera Canyon (Catalan Sea, north-western Mediterranean) and on the adjacent continental slope. These provide the third record of this apparently endemic Mediterranean species and the first record from the western Mediterranean. This was the only holothurian species trapped and the most abundant macroscopic organism found in the traps between 1200 and 1700 m depth over the whole sampling period (March–November 2001). It was particularly abundant in spring during the main flux of organic particles in the canyon. This coupling suggests that Penilpidia may aggregate at the seafloor during these events, making food availability a plausible explanation of the seasonal occurrence. Lateral transport of material re-suspended locally or up-canyon by near bottom currents appear to be the mechanism behind this uncommon occurrence, although in situ observations have recently been made on the swimming capability of this holothurian. The occurrence of benthic organisms in sediment traps set close to the seabed can provide information on bathyal benthic and benthopelagic populations.     

Benthic trophic status and nutrient fluxes in shallow-water sediments

Proliferation of fast-growing ephemeral macroalgae in shallow-water embayments constitutes a large-scale environmental change of coastal marine ecosystems. Since inorganic nutrients essential for the initiation and maintenance of macroalgal growth may be supplied from the underlying sediment, we investigated the coupling between benthic inorganic nutrient (mainly N and P) fluxes and sediment properties in 6 bays representing a wide gradient of sediment characteristics (grain size, organic matter content, solid phase C and N). The initial characterization of bays was made in June and also included measurements of oxygen flux and microphytobenthic and macrofaunal biomass. In September, still within the growth season of the macroalgae, complementary experiments with sediment-water incubations for benthic flux measurements of oxygen and nutrients focused on trophic status (balance between auto- and heterotrophy) as a controlling factor for rates of measured benthic nutrient fluxes. Generally, sediments rendered autotrophic by microphytobenthic photosynthesis removed nutrients from the overlying water, while heterotrophic sediments supplied nutrients to the overlying bottom water. Estimations of the green-algal nutrient demand suggested that late in the growth season, net heterotrophic sediments could cover 20% of the N-demand and 70% of the P demand. As the benthic trophic status is a functional variable more closely coupled to nutrient fluxes than the comparably conservative structural parameter organic matter content, we suggest that the trophic status is a more viable parameter to classify sediments and predict benthic nutrient fluxes in shallow-water environments.     

Nitrogen cycling in deep-sea sediments of the Porcupine Abyssal Plain, NE Atlantic

Rates of transformation, recycling and burial of nitrogen and their temporal and spatial variability were investigated in deep-sea sediments of the Porcupine Abyssal Plain (PAP), NE Atlantic during eight cruises from 1996 to 2000. Benthic fluxes of ammonium (NH₄) and nitrate (NO₃) were measured in situ using a benthic lander. Fluxes of dissolved organic nitrogen (DON) and denitrification rates were calculated from pore water profiles of DON and NO₃, respectively. Burial of nitrogen was calculated from down core profiles of nitrogen in the solid phase together with ¹⁴C-based sediment accumulation rates and dry bulk density. Average NH₄ and NO₃-effluxes were 7.4 ± 19 μmol m⁻² d⁻¹ (n = 7) and 52 ± 30 μmol m⁻² d⁻¹ (n = 14), respectively, during the period 1996–2000. During the same period, the DON-flux was 11 ± 5.6 μmol m⁻² d⁻¹ (n = 5) and the denitrification rate was 5.1 ± 3.0 μmol m⁻² d⁻¹ (n = 22). Temporal and spatial variations were only found in the benthic NO₃ fluxes. The average burial rate was 4.6 ± 0.9 μmol m⁻² d⁻¹. On average over the sampling period, the recycling efficiency of the PON input to the sediment was 94% and the burial efficiency hence 6%. The DON flux constituted 14% of the nitrogen recycled, and it was of similar magnitude as the sum of burial and denitrification. By assuming the PAP is representative of all deep-sea areas, rates of denitrification, burial and DON efflux were extrapolated to the total area of the deep-sea floor (>2000 m) and integrated values of denitrification and burial of 8 ± 5 and 7 ± 1 Tg N year⁻¹, respectively, were obtained. This value of total deep-sea sediment denitrification corresponds to 3–12% of the global ocean benthic denitrification. Burial in deep-sea sediments makes up at least 25% of the global ocean nitrogen burial. The integrated DON flux from the deep-sea floor is comparable in magnitude to a reported global riverine input of DON suggesting that deep-sea sediments constitute an important source of DON to the world ocean.     

Population structure, density and food sources of Terebralia palustris (Potamididae: Gastropoda) in a low intertidal Avicennia marina mangrove stand (Inhaca Island, Mozambique)

Population structure and distribution of Terebralia palustris were compared with the environmental parameters within microhabitats in a monospecific stand of Avicennia marina in southern Mozambique. Stable carbon and nitrogen isotope analyses of T. palustris and potential food sources (leaves, pneumatophore epiphytes, and surface sediments) were examined to establish the feeding preferences of T. palustris. Stable isotope signatures of individuals of different size classes and from different microhabitats were compared with local food sources. Samples of surface sediments 2.5–10 m apart showed some variation (−21.2‰ to −23.0‰) in δ¹³C, probably due to different contributions from seagrasses, microalgae and mangrove leaves, while δ¹⁵N values varied between 8.7‰ and 15.8‰, indicating that there is a very high variability within a small-scale microcosm. Stable isotope signatures differed significantly between the T. palustris size classes and between individuals of the same size class, collected in different microhabitats. Results also suggested that smaller individuals feed on sediment, selecting mainly benthic microalgae, while larger individuals feed on sediment, epiphytes and mangrove leaves. Correlations were found between environmental parameters and gastropod population structure and distribution vs. the feeding preferences of individuals of different size classes and in different microhabitats. While organic content and the abundance of leaves were parameters that correlated best with the total density of gastropods (>85%), the abundance of pneumatophores and leaves, as well as grain size, correlated better with the gastropod size distribution (>65%). Young individuals (height < 3 cm) occur predominantly in microhabitats characterized by a low density of leaf litter and pneumatophores, reduced organic matter and larger grain size, these being characteristic of lower intertidal open areas that favour benthic microalgal growth. With increasing shell height, T. palustris individuals start occupying microhabitats nearer the mangrove trees characterized by large densities of pneumatophores and litter, as well as sediments of smaller grain size, leading to higher organic matter availability in the sediment.     

Predicting oxygen in small estuaries of the Baltic Sea: a comparative approach

Coastal eutrophication, manifested as hypoxia and anoxia, is a global problem. Only a few empirical models, however, exist to predict bottom oxygen concentration and percentage saturation from nutrient load or morphometry in coastal waters, which are successfully used to predict phytoplankton biomass both in lakes and in estuaries. Furthermore, hardly any empirical models exist to predict bottom oxygen from land-use. A data set was compiled for 19 estuaries in the northern Baltic Sea, which included oxygen concentration and percentage saturation, water chemistry, estuary morphometry, and land-use characteristics. In regression analyses, bottom oxygen was predicted both as a function of the percentage of watershed under agriculture and of mean depth. These models accounted for ca. 55% of the variation in oxygen. Additionally, oxygen was linked to fetch (diameter of the area in the direction of the prevailing wind), which accounted for 30% of the variation in oxygen. This suggests that shallow Finnish estuaries are wind-sensitive. In ‘pits’ (sub-thermocline waters of deep basins), near-bottom total nitrogen strongly correlated with oxygen percentage saturation (R²=0.81). Neither chlorophyll a, total phosphorus nor nutrient loading explained oxygen variation in entire estuaries or in ‘pits’, probably mainly due to annual sedimentation/sediment–water flux dynamics. On the basis of the results of cross-validation, the models have general applicability among Finnish estuaries.     

Dissolved titanium distributions in the Mid-Atlantic Bight

Although titanium is abundant in Earth's crust, its sources and distribution in the ocean are poorly understood. To elucidate its behavior, distributions of dissolved (< 0.2 μm) Ti were determined in surface waters and vertical profiles from the Mid-Atlantic Bight (MAB). Concentrations of Ti decreased from 390 pM at the Delaware Bay mouth to < 100 pM across the Delaware continental shelf. In vertical profiles, small increases in bottom waters suggest a possible flux of Ti from shelf sediments, consistent with previous reports of pore water enrichments of dissolved Ti in MAB sediments. Concentrations in surface waters of the outer shelf and slope ranged between 30 and 140 pM, with most values below 90 pM. Concentrations in a 1000 m vertical profile in the eastern Gulf Stream ranged between 110 and 280 pM, and showed a variable distribution attributed to the mixing of water masses in the outer MAB. A simple model of Ti sources to the MAB suggests that atmospheric deposition of dissolved Ti is comparable to net riverine contributions and therefore must be considered in applications of Ti as a tracer of oceanographic processes.