Trophic ecology and vertical patterns of carbon and nitrogen stable isotopes in zooplankton from oxygen minimum zone regions

The unique physical and biogeochemical characteristics of oxygen minimum zones (OMZs) influence plankton ecology, including zooplankton trophic webs. Using carbon and nitrogen stable isotopes, this study examined zooplankton trophic webs in the Eastern Tropical North Pacific (ETNP) OMZ. δ¹³C values were used to indicate zooplankton food sources, and δ¹⁵N values were used to indicate zooplankton trophic position and nitrogen cycle pathways. Vertically stratified MOCNESS net tows collected zooplankton from 0 to 1000 m at two stations along a north-south transect in the ETNP during 2007 and 2008, the Tehuantepec Bowl and the Costa Rica Dome. Zooplankton samples were separated into four size fractions for stable isotope analyses. Particulate organic matter (POM), assumed to represent a primary food source for zooplankton, was collected with McLane large volume in situ pumps.The isotopic composition and trophic ecology of the ETNP zooplankton community had distinct spatial and vertical patterns influenced by OMZ structure. The most pronounced vertical isotope gradients occurred near the upper and lower OMZ oxyclines. Material with lower δ¹³C values was apparently produced in the upper oxycline, possibly by chemoautotrophic microbes, and was subsequently consumed by zooplankton. Between-station differences in δ¹⁵N values suggested that different nitrogen cycle processes were dominant at the two locations, which influenced the isotopic characteristics of the zooplankton community. A strong depth gradient in zooplankton δ¹⁵N values in the lower oxycline suggested an increase in trophic cycling just below the core of the OMZ. Shallow POM (0–110 m) was likely the most important food source for mixed layer, upper oxycline, and OMZ core zooplankton, while deep POM was an important food source for most lower oxycline zooplankton (except for samples dominated by the seasonally migrating copepod Eucalanus inermis). There was no consistent isotopic progression among the four zooplankton size classes for these bulk mixed assemblage samples, implying overlapping trophic webs within the total size range considered.     

Diel variations of the bathymetric distribution of zooplankton groups and biomass in Cap-Ferret Canyon,France

The bathymetric distribution, abundance and diel vertical migrations (DVM) of zooplankton were investigated along the axis of the Cap-Ferret Canyon (Bay of Biscay, French Atlantic coast) by a consecutive series of synchronous net hauls that sampled the whole water column (0–2000 m in depth) during a diel cycle. The distribution of appendicularians (maximum 189 individuals m⁻³), cladocerans (maximum 287 individuals m⁻³), copepods (copepods<4 mm, maximum 773 individuals m⁻³, copepods>4 mm, maximum 13 individuals m⁻³), ostracods (maximum 8 individuals m⁻³), siphonophores (maximum >2 individuals m⁻³) and peracarids (maximum >600 individuals 1000 m⁻³) were analysed and represented by isoline diagrams. The biomass of total zooplankton (maximum 18419 μg C m⁻³, 3780 μg N m⁻³) and large copepods (>4 mm maximum 2256 μg C m⁻³, 425 μg N m⁻³) also were determined. Vertical migration was absent or affected only the epipelagic zone for appendicularians, cladocerans, small copepods and siphonophores. Average amplitude of vertical migration was about 400–500 m for ostracods, some hyperiids and mysids, and large copepods, which were often present in the epipelagic, mesopelagic, and bathypelagic zones. Large copepods can constitute more than 80% of the biomass corresponding to total zooplankton. They may play an important role in the active vertical transfer of carbon and nitrogen.     

Aerobic ammonium oxidation in the oxycline and oxygen minimum zone of the eastern tropical South Pacific off northern Chile (∼20°S)

Aerobic NH₄⁺ oxidation rates were measured along the strong oxygen gradient associated with the oxygen minimum zone (OMZ) of the eastern tropical South Pacific off northern Chile (∼20°S) during 2000, 2003, and 2004. This process was examined by comparing NH₄⁺ rates of change during dark incubations, with and without the addition of allylthiourea, a classical inhibitor of the ammonia monooxygenase enzyme of ammonium-oxidizing bacteria. The contribution of aerobic NH₄⁺ oxidation in dark carbon fixation and NO₂⁻ rates of change were also explored. Thirteen samples were retrieved from the oxycline (252 to ⩽5 μM O₂; 15 to ∼65 m depth) and three from the oxygen minimum core (⩽5 μM O₂; 100–200 m depth). Aerobic NH₄⁺ oxidation rates were mainly detected in the upper part (15–30 m depth) of the oxycline, with rates ranging from 0.16 to 0.79 μM d⁻¹, but not towards the oxycline base (40–65 m depth). In the oxygen minimum core, aerobic NH₄⁺ oxidation was in the upper range and higher than in the upper part of the oxycline (0.70 and 1.0 μM d⁻¹). Carbon fixation rates through aerobic NH₄⁺ oxidation ranged from 0.18 to 0.43 μg C L⁻¹ d⁻¹ and contributed between 33% and 57% of the total dark carbon fixation, mainly towards the oxycline base and, in a single experiment, in the upper part of the oxycline. NO₂⁻ consumption was high (up to 10 μM d⁻¹) towards the oxycline base and OMZ core, but was significantly reduced in experiments amended with allylthiourea, indicating that aerobic NH₄⁺ oxidation could contribute between 8% and 76% of NO₂⁻ production, which in turn could be available for denitrifiers. Overall, these results support the important role of aerobic NH₄⁺ oxidizers in the nitrogen and carbon cycling in the OMZ and at its upper boundary.     

Nitrous oxide and N-nutrient cycling in the oxygen minimum zone off northern Chile

Measurements of dissolved gases (O₂, N₂O), nutrients (NO₃⁻, NO₂⁻, PO₄³⁻), and oceanographic variables were performed off northern Chile (∼21°S) between March 2000 and July 2004, in order to characterize the existing oxygen minimum zone (OMZ) and identify processes involved in N₂O cycling. Both N₂O and NO₃⁻ displayed sharp, shallow peaks with concentrations of up to 124 nM (1370% saturation) and 26 μM, respectively, in association with a strong oxycline that impinges on the euphotic zone. NO₂⁻ accumulation below the oxycline's base reached up to 9 μM. The vertical distribution of physical and chemical parameters and the existing relationships between apparent oxygen utilization (AOU), apparent N₂O production (ΔN₂O), and NO₃⁻ revealed three main layers within the upper OMZ. The first layer, or the upper part of the oxycline, is located between the base of the mixed layer and the mid-point of the oxycline (around σ_t=25.5 kg m⁻³). There the O₂ declines from ∼250 to ∼50 μM, and strong (but opposing) O₂ and NO₃⁻ gradients and their associated AOU–ΔN₂O and AOU–NO₃⁻ relationships indicate that nitrification produces N₂O and NO₃⁻ in the presence of light. The second layer, or lower part of the oxycline, represents the upper OMZ boundary and is located between the middle and the base of the oxycline (25.9<σ_t<26.1 kg m⁻³). In this layer NO₃⁻ reduction begins at O₂ levels ranging from ∼50 to ∼11 μM and accumulation of 41–68% of the ΔN₂O pool occurs. The accumulation of N₂O (but not of NO₂⁻ or NH₄⁺) and the observed AOU–ΔN₂O and AOU–NO₃⁻ relationships (which are opposite to those of the overlying first layer) suggest that a coupling between nitrification and NO₃⁻ reduction is involved in N₂O cycling in this second layer. The third layer is the OMZ core, where the O₂ concentration remains constant (O₂<11 μM). It coincides with σ_t>26.2 kg m⁻³, which is typical of Equatorial Subsurface Water (ESSW). In this layer, N₂O and NO₃⁻ continue to decrease, but a large NO₂⁻ accumulation is observed. Considering all the data, a biogeochemical model for the upper OMZ off northern of Chile is proposed, in which nitrification and denitrification differentially mediate N₂O cycling in each layer.     

Distribution of zooplankton biomass at three seamounts in the NE Atlantic

During different seasons of the years 2003–2005 in the NE Atlantic, zooplankton were sampled with a MOCNESS (multiple opening/closing net and environmental sensing system, mesh size 333 μm) above the slopes and summits of Seine, Sedlo and Ampère seamounts and at remote reference sites outside the influence of the seamounts (far field). Wet weights of different zooplankton size classes (<0.5, 0.5–2, >2 cm) were measured. Night and day hauls were analysed in order to detect diel vertical migrations of the zooplankton, as well as a possible trapping effect due to the shallow topography.Biomass concentrations, independent of daytime, season and summit height, were reduced above the summits at all three seamounts compared to the slope and far-field sites. No trapping effect or retention of biomass was apparent above the seamounts. The vertical distribution patterns of the size class <0.5 cm did not differ between night and day hauls at most sites, but indications of diel vertical migrations were found in the larger size fractions. With the exception of gelatinous organisms, zooplankton >0.5 cm were nearly absent above the summits of Seine and Ampère seamounts, but considerable numbers were found above the slopes and at the far-field sites. Possible explanations for the observed distribution patterns of zooplankton biomass and size classes are discussed, including retention and lateral advection due to the hydrography at the seamounts, as well as predation by resident seamount fish.     

Polychaete community structure in the South Eastern Arabian Sea continental margin (200–1000 m)

Macrofaunal polychaete communities (>500 µm) in the South Eastern Arabian Sea (SEAS) continental margin (200–1000 m) are described, based on three systematic surveys carried out in 9 transects (at ~200 m, 500 m and 1000 m) between 7°00′and 14°30′N latitudes. A total of 7938 polychaetes belonging to 195 species were obtained in 136 grab samples collected at 27 sites. Three distinct assemblages were identified in the northern part of the SEAS margin (10–14°30′N), occupying the three sampled depth strata (shelf edge, upper and mid-slope) and two assemblages (shelf edge and slope) in the south (7–10°N). Highest density of polychaetes and dominance of a few species were observed in the shelf edge, where the Arabian Sea oxygen minimum zone (OMZ) impinged on the seafloor, particularly in the northern transects. The resident fauna in this region (Cossura coasta, Paraonis gracilis, Prionospio spp. and Tharyx spp.) were characteristically of smaller size, and well suited to thrive in the sandy sediments in OMZ settings. Densities were lowest along the most northerly transect (T9), where dissolved oxygen (DO) concentrations were extremely low (<0.15 ml l⁻¹, i.e.<6.7 μmol l⁻¹). Beyond the realm of influence of the OMZ (i.e. mid-slope, ~1000 m), the faunal density decreased while species diversity increased. The relative proportion of silt increased with depth, and the dominance of the aforementioned species decreased, giving way to forms such as Paraprionospio pinnata, Notomastus sp., Eunoe sp. and lumbrinerids. Relatively high species richness and diversity were observed in the sandy sediments of the southern sector (7–9°N), where influence of the OMZ was less intense. The area was also characterized by certain species (e.g. Aionidella cirrobranchiata, Isolda pulchella) that were nearly absent in the northern region. The gradients in DO concentration across the core and lower boundary of the OMZ, along with bathymetric and latitudinal variation in sediment texture, were responsible for differences in polychaete size and community structure on the SEAS margin. Spatial and temporal variations were observed in organic matter (OM) content of the sediment, but these were not reflected in the density, diversity or distribution pattern of the polychaetes.     

Diel variation in the vertical distribution of particulate matter (>0.15 mm) in the NW Mediterranean Sea investigated with the Underwater Video Profiler

Day/night variations in the size distribution of the particulate matter >0.15 mm (PM) were studied in May 1995 during the DYNAPROC time-series cruise in the northwestern Mediterranean Sea. Data on vertical distributions of PM (>0.15 mm) and zooplankton were collected with the Underwater Video Profiler (UVP). The comparisons of the UVP data with plankton net data and POC data from water bottles indicated that more than 97% of the particles detected by the UVP were non-living particles (0.15 mm) and that the PM contributed 4–34% of the total dry weight measured on GF/F filters. Comparison of seven pairs of day and night vertical profiles performed during the cruise showed that in the upper 800 m, the mean size and the volume of particles was higher at night than during the day. During the night, the integrated volume of the PM increased on average by 32±20%. This increase corresponded to a shift of smaller size classes (<0.5 mm) towards the larger ones (>0.5 mm). During the day, the pattern was reversed, and the quantity of PM >0.5 mm decreased. During the study period, the standing stock of PM (60–800 m) decreased from 7.5 to less than 2 g m⁻² but the diel variations persisted, except for two short periods in the superficial layer following a wind event. The cyclic feeding activity induced by the diel vertical migration of zooplankton could be the best candidate to explain the observed diel fluctuations in the size classes of PM in the water column. However, our results also suggest that in the upper layer additional driving forces such as the increase of the level of turbulence after a wind event or the modification of the zoo- and phytoplankton community can influence the PM temporal evolution.     

The swarm dynamics of northern krill (Meganyctiphanes norvegica) and pteropods (Cavolinia inflexa) during vertical migration in the Ligurian Sea observed by an acoustic Doppler current profiler

A ship-mounted 153 kHz narrow-band ADCP and 1 m² MOCNESS were deployed between 16 and 24 Sept. 1997 in the Ligurian central zone (∼43°20′N 7°48′E). Results from both instruments showed that the zooplankton community performed vertical migrations that conformed to the classical pattern of ascent at dusk (∼18:30 h) and descent at dawn (∼06:30 h). Depth-discrete net samples between 0 and 500 m showed that the community was dominated by two species, the euphausiid Meganyctiphanes norvegica (Northern krill) and the pteropod Cavolinia inflexa, which migrated in separate discrete bands that were detectable by the ADCP. Information from the ADCP was used to estimate vertical migration speed in two ways: (i) from the trajectory of the back-scattering bands over time and (ii) from the Doppler-shift vertical velocity measured within depth zones at the corresponding time and depth of these bands. Estimates of the migration speed of C. inflexa were between 2 and 7 cm s⁻¹ upwards and between 4 and 7 cm s⁻¹ downwards. M. norvegica was estimated to migrate at speeds between 7 and 8 cm s⁻¹ upwards and over 11 cm s⁻¹ downwards. The consistently lower migration speeds estimated from Doppler measurements as compared with estimates obtained from measuring trajectories of back-scattering bands over time was believed to result from a methodological artefact. The Doppler measurements were nevertheless useful in a relative sense in revealing the relative speed of individuals within swarms. It was shown that individuals at the front of the upwardly migrating band of M. norvegica moved more slowly than those at the rear. These results illustrate the extra biological information that can be obtained by ADCPs compared with conventional echo-sounders.     

Vertical zonation and distributions of calanoid copepods through the lower oxycline of the Arabian Sea oxygen minimum zone

This paper provides the first comprehensive analysis of calanoid copepod vertical zonation and community structure at midwater depths (300–1000 m) through the lower oxygen gradient (oxycline) (0.02 to 0.3 ml/L) of an oxygen minimum zone (OMZ). Feeding ecology was also analyzed. Zooplankton were collected with a double 1 m² MOCNESS plankton net in day and night vertically-stratified oblique tows from 1000 m to the surface at six stations during four seasons as part of the 1995 US Joint Global Ocean Flux Study (JGOFS) Arabian Sea project. The geographic comparison between a eutrophic more oxygenated onshore station and an offshore station with a strong OMZ served as a natural experiment to elucidate the influence of depth, oxygen concentration, season, food resources, and predators on the copepod distributions.Copepod species and species assemblages of the Arabian Sea OMZ differed in their spatial and vertical distributions relative to environmental and ecological characteristics of the water column and region. The extent and intensity of the oxycline at the lower boundary of the OMZ, and its spatial and temporal variability over the year of sampling, was an important factor affecting distributional patterns. Calanoid copepod species showed vertical zonation through the lower OMZ oxycline. Clustering analyses defined sample groups with similar copepod assemblages and species groups with similar distributions. No apparent diel vertical migration for either calanoid or non-calanoid copepods at these midwater depths was observed, but some species had age-related differences in vertical distributions. Subzones of the OMZ, termed the OMZ Core, the Lower Oxycline, and the Sub-Oxycline, had different copepod communities and ecological interactions. Major distributional and ecological changes were associated with surprisingly small oxygen gradients at low oxygen concentrations. The calanoid copepod community was most diverse in the most oxygenated environments (oxygen >0.14 ml/L), but the rank order of abundance of species was similar in the Lower Oxycline and Sub-Oxycline. Some species were absent or much scarcer in the OMZ Core. Two copepod species common in the Lower Oxycline were primarily detritivorous but showed dietary differences suggesting feeding specialization. The copepod Spinocalanus antarcticus fed primarily on components of the vertical particulate flux and suspended material, a less versatile diet than the co-occurring copepod Lucicutia grandis. Vertical zonation of copepod species through the lower OMZ oxycline is probably a complex interplay between physiological limitation by low oxygen, potential predator control, and potential food resources. Pelagic OMZ and oxycline communities, and their ecological interactions in the water column and with the benthos, may become even more widespread and significant in the future ocean, if global warming increases the extent and intensity of OMZs as predicted.     

Vertical distribution of zooplankton and active flux across an anticyclonic eddy in the Canary Island waters

The vertical distribution (0–900 m) of zooplankton biomass and indices of feeding (gut fluorescence, GF) and metabolism (electron transfer system, ETS) were studied across an anticyclonic eddy south of Gran Canaria Island (Canary Islands). Two dense layers of organisms were clearly observed during the day, one above 200 m and the other at about 500 m, coincident with the deep scattering layer (DSL). The biomass displacement due to interzonal migrants in the euphotic zone was more than 2-fold higher than that previously reported for the southern area of this archipelago. The gut flux estimated (0.14–0.44 mgC m⁻² d⁻¹) was similar to the values previously found in the Canaries. The respiratory flux outside the eddy (1.85 mgC m⁻² d⁻¹) was in the lower range of values reported for this area. Inside the eddy, migrant biomass and respiration rates were 2- and 4- fold higher than in the surrounding waters. Active flux mediated by diel vertical migrants inside the eddy (8.28 mgC m⁻² d⁻¹) was up to 53% of the passive carbon flux to the mesopelagic zone (15.8 mgC m⁻² d⁻¹). It is, therefore, suggested that the anticyclonic eddy enhanced both migration from deep waters and active flux.     

Seasonal variation of diel vertical migration of zooplankton from ADCP backscatter time series data in the Lazarev Sea,Antarctica

Ten-month time series of mean volume backscattering strength (MVBS) and vertical velocity obtained from three moored acoustic Doppler current profilers (ADCPs) deployed from February until December 2005 at 64°S, 66.5°S and 69°S along the Greenwich Meridian were used to analyse the diel vertical zooplankton migration (DVM) and its seasonality and regional variability in the Lazarev Sea. The estimated MVBS exhibited distinct patterns of DVM at all three mooring sites. Between February and October, the timing of the DVM and the residence time of zooplankton at depth were clearly governed by the day–night rhythm. Mean daily cycles of the ADCP-derived vertical velocity were calculated for successive months and showed maximum ascent and descent velocities of 16 and –15 mm s^?1. However, a change of the MVBS pattern occurred in late spring/early austral summer (October/November), when the zooplankton communities ceased their synchronous vertical migration at all three mooring sites. Elevated MVBS values were then concentrated in the uppermost layers (<50 m) at 66.5°S. This period coincided with the decay of sea ice coverage at 64°S and 66.5°S between early November and mid-December. Elevated chlorophyll concentrations, which were measured at the end of the deployment, extended from 67°S to 65°S and indicated a phytoplankton bloom in the upper 50 m. Thus, we propose that the increased food supply associated with an ice edge bloom caused the zooplankton communities to cease their DVM in favour of feeding.     

Zooplankton metabolism and carbon demand at two seamounts in the NE Atlantic

Zooplankton metabolic rates, determined from electron transfer system (ETS) activity, were studied at two seamounts (Seine: 34°N, 14°W, summit depth ∼170 m; Sedlo: 40°N, 27°W, summit depth ∼750 m) in the northeast (NE) Atlantic during three cruises in November 2003, April 2004 and July 2004. ETS activity and respiratory carbon demand were measured for samples taken at seamount and open-ocean locations in order to probe the hypothesis of locally enhanced seamount productivity. ETS activity and biomass revealed no consistent diel patterns of feeding activity and vertical migration at Seine and Sedlo Seamounts. Spatial differences of biomass-specific ETS activity were observed at both seamounts and coincided with differences in food abundance and quality. At Seine Seamount in April 2004, biomass-specific ETS activity was on average higher at the seamount locations compared to the open ocean, though the enhancement was of a lower magnitude than spatial and temporal variability and had no apparent influence on zooplankton respiratory carbon demand or biomass. A persistent pattern of reduced zooplankton biomass above the summit location at Seine Seamount in April 2004 and July 2004 resulted in a local reduction of respiratory carbon demand. At Sedlo Seamount in November 2003, large spatial differences in biomass-specific ETS activity observed at the seamount locations resulted in a large range of respiratory carbon demand at the seamount, but were not reflected in zooplankton biomass. The depth-integrated (0–150 m) median respiratory carbon demand of the zooplankton community estimated from day and night hauls was 2.1 mg C m⁻² d⁻¹ at Seine Seamount (range: 0.3–6.3) and 2.9 mg C m⁻² d⁻¹ at Sedlo Seamount (range: 1.6–12.0). The sporadic nature and low magnitude of locally higher zooplankton respiration rates at the seamounts, which did not result in locally higher zooplankton standing stock biomass, lead us to reject the hypothesis that locally enhanced seamount productivity provides an autochthonous food supply to the resident faunas at Seine and Sedlo Seamounts. Instead, we conclude that the faunas at both seamounts are more likely supported by advection of food from the surrounding ocean.     

Seasonal changes in zooplankton abundance,biomass, size structure and dominant copepods in the Oyashio region analysed by an optical plankton counter

To identify seasonal patterns of change in zooplankton communities, an optical plankton counter (OPC) and microscopic analysis were utilised to characterise zooplankton samples collected from 0 to 150 m and 0 to 500 m in the Oyashio region every one to three months from 2002 to 2007. Based on the OPC measurements, the abundance and biomass of zooplankton peaked in June (0–150 m) or August (150–500 m), depending on the depth stratum. The peak periods of the copepod species that were dominant in terms of abundance and biomass indicated species-specific patterns. Three Neocalanus species (Neocalanus cristatus, Neocalanus flemingeri and Neocalanus plumchrus) exhibited abundance peaks that occurred before their biomass peaks, whereas Eucalanus bungii and Metridia pacifica experienced biomass peaks before their abundance peaks. The abundance peaks corresponded to the recruitment periods of early copepodid stages, whereas the biomass peaks corresponded to the periods when the dominant populations reached the late copepodid stages (C5 or C6). Because the reproduction of Neocalanus spp. occurred in the deep layer (>500 m), their biomass peaks were observed when the major populations reached stage C5 after the abundance peaks of the early copepodid stages. The reproduction of E. bungii and M. pacifica occurred near the surface layer. These species first formed biomass peaks of C6 and later developed abundance peaks of newly recruited early copepodid stages. From the comparison between OPC measurements and microscopic analyses, seasonal changes in zooplankton biomass at depths of 0–150 m were governed primarily by E. bungii and M. pacifica, whereas those at depths of 150–500 m were primarily caused by the three Neocalanus species.     

Meiofaunal distributions on the Peru margin:: relationship to oxygen and organic matter availability

A quantitative study of metazoan meiofauna was carried out on bathyal sediments (305, 562, 830 and 1210 m) along a transect within and beneath the oxygen minimum zone (OMZ) in the southeastern Pacific off Callao, Peru (12°S). Meiobenthos densities ranged from 1517 (upper slope, middle of OMZ) to 440–548 ind. 10 cm⁻² (lower slope stations, beneath the OMZ). Nematodes were the numerically dominant meiofaunal taxon at every station, followed by copepods and nauplii. Increasing bottom-water oxygen concentration and decreasing organic matter availability downslope were correlated with observed changes in meiofaunal abundance. The 300-m site, located in the middle of the OMZ, differed significantly in meiofaunal abundance, dominance, and in vertical distribution pattern from the deeper sites. At 305 m, nematodes amounted to over 99% of total meiofauna; about 70% of nematodes were found in the 2–5 cm interval. At the deeper sites, about 50% were restricted to the top 1 cm. The importance of copepods and nauplii increased consistently with depth, reaching ∼12% of the total meiofauna at the deepest site. The observation of high nematode abundances at oxygen concentrations <0.02 ml l⁻¹ supports the hypothesis that densities are enhanced by an indirect positive effect of low oxygen involving (a) reduction of predators and competitors and (b) preservation of organic matter leading to high food availability and quality. Food input and quality, represented here by chloroplastic pigment equivalents (CPE) and sedimentary labile organic compounds (protein, carbohydrates and lipids), were strongly, positively correlated with nematode abundance. By way of contrast, oxygen exhibited a strong negative correlation, overriding food availability, with abundance of other meiofauna such as copepods and nauplii. These taxa were absent at the 300-m site. The high correlation of labile organic matter (C-LOM, sum of carbon contents in lipids, proteins and carbohydrates) with CPE (Pearson's r=0.99, p<0.01) suggests that most of the sedimentary organic material sampled was of phytodetrital origin. The fraction of sediment organic carbon potentially available to benthic heterotrophs, measured as C-LOM/Total organic carbon, was on average 17% at all stations. Thus, a residual, refractory fraction, constitutes the major portion of organic matter at the studied bathyal sites.     

Anammox bacteria and the anaerobic oxidation of ammonium in the oxygen minimum zone off northern Chile

Anammox is the anaerobic oxidation of ammonium by nitrite or nitrate to yield N₂. This process, along with conventional denitrification, contributes to nitrogen loss in oxygen-deficient systems. Anammox is performed by a special group of bacteria belonging to the Planctomycetes phylum. However, information about the distribution, activity, and controlling factors of these anammox bacteria is still limited. Herein, we examine the phylogenetic diversity, vertical distribution, and activity of anammox bacteria in the coastal upwelling region and oxygen minimum zone off northern Chile. The phylogeny of anammox bacteria was studied using primers designed to specifically target 16S rRNA genes from Planctomycetes in samples taken during a cruise in 2004. Anammox bacteria-like sequences affiliated with Candidatus “Scalindua spp.” dominated the 16S rRNA gene clone library. However, 62% of the sequences subgrouped separately within this cluster and together with a single sequence retrieved from the suboxic zone of the freshwater Lake Tanganyika. The vertical distribution and activity of anammox bacteria were explored through CARD-FISH (fluorescence in situ hybridization with catalyzed reporter deposition) and ¹⁵N labeling incubations, respectively, at two different open-ocean stations during a second cruise in 2005. Anammox bacterial CARD-FISH counts (up to 3000 cells ml⁻¹) and activity (up to 5.75 nmol N₂ L⁻¹ d⁻¹) were only detected at the station subjected directly to the upwelling influence. Anammox cell abundance and activity were highest at 50 m depth, which is the upper part of the OMZ. In this layer, a high abundance of cyanobacteria and a marked nitrogen deficit were also observed. Thus, our results show the presence of a new subcluster within the marine anammox phylogeny and indicate high vertical variability in the abundance and activity of anammox bacteria that could be related to an intensification of carbon and nitrogen cycling in the upper part of the OMZ.     

Biomass size-spectra of macrobenthic communities in the oxygen minimum zone off Chile

Estimates of macrofaunal secondary production and normalized biomass size-spectra (NBSS) were constructed for macrobenthic communities associated with the oxygen minimum zone (OMZ) in four areas of the continental margin off Chile. The presence of low oxygen conditions in the Humboldt Current System (HCS) off Chile was shown to have important effects on the size structure and secondary production of the benthic communities living in this ecosystem. The distribution of normalized biomass by size was linear (log₂–log₂ scale) at all stations. The slope of the NBSS ranged from −0.481 to −0.908. There were significant differences between the slopes of the NBS-spectra from the stations located in the OMZ (slope = −0.837) and those located outside the OMZ (slope = −0.463) (p < 0.05). The results of this study suggest that low oxygen conditions (<0.5 ml L⁻¹) appear to influence biomass size-spectra, because small organisms are better able to satisfy their metabolic demands. The annual secondary production was higher off central Chile (6.8 g C m⁻² y⁻¹) than off northern Chile (2.02 g C m⁻² y⁻¹) and off southern Chile (0.83 g C m⁻² y⁻¹). A comparison with other studies suggests that secondary production in terms of carbon equivalents was higher than in other upwelling regions.     

High-frequency fluxes of labile compounds in the central Ligurian Sea,northwestern Mediterranean

Sinking particles were collected every 4 h with drifting sediment traps deployed at 200 m depth in May 1995 in a 1-D vertical system during the DYNAPROC observations in the northwestern Mediterranean sea. POC, proteins, glucosamine and lipid classes were used as indicators of the intensity and quality of the particle flux. The roles of day/night cycle and wind on the particle flux were examined. The transient regime of production from late spring bloom to pre-oligotrophy determined the flux intensity and quality. POC fluxes decreased from, on average, 34 to 11 mg m⁻² d⁻¹, representing 6–14% of the primary production under late spring bloom conditions to 1–2% under pre-oligotrophic conditions. Total protein and chloroplast lipid fluxes correlated with POC and reflected the input of algal biomass into the traps. As the season proceeded, changes in the biochemical composition of the exported material were observed. The C/N ratio rose from 7.8 to 12. Increases of serine (10–28% of total proteins), total lipids (7–9 to 14–28% of POC) and reserve lipids (1–5 to 5–22% of total lipids) were noticeable, whereas total protein content in POC decreased (20–27 to 18–7%). N-acetyl glucosamine, a tracer of fecal pellet flux, showed that zooplankton grazing was a major vector of downward export during the decaying bloom. Against this background pattern, episodic events specifically increased the flux, modifying the quality and the settling velocity of particles. Day/night signals in biotracers (POC, N-acetyl glucosamine, protein and chloroplast lipids) showed that zooplankton migrations were responsible for sedimentation of fresh material through fast sinking particles (V=170–180 m d⁻¹) at night. Periodic signatures of re-processed material (high lipolysis and bacterial biomass indices) suggested that other zooplankton fecal pellets or small aggregates, probably of lower settling velocities (V<170 m d⁻¹), contributed to the flux during calm periods. At the beginning of the experiment, during the development of a prymnesiophyte bloom in the upper layers, the sterol signal with no periodicity enabled us to estimate high particle settling velocities (⩾600 m d⁻¹) likely related to large aggregate formation. A wind event increased biotracer fluxes (POC, protein, chloroplast lipids). The rapid transmission of surface signals through extremely fast sinking particles could be a general feature of particle fluxes in marine areas unaffected by horizontal advection.     

Impacts of the wintertime mesozooplankton community to downward carbon flux in the subarctic and subtropical Pacific Oceans

We compared wintertime depth distributions of the mesozooplankton community and dominant copepods between the subtropical (S1) and subarctic (K2) Pacific Oceans to evaluate the relative importance of actively transported carbon by vertical migrants to sinking particulate organic carbon flux. Primary production was higher and the ratio of sinking particulate organic carbon flux to primary production was lower at S1 compared with those at K2. The mesozooplankton community was lower in abundance and biomass at S1 compared to K2. Copepods were the dominant group among both mesozooplankton abundance and biomass throughout the water column down to 1000 m at both sites. The depth distribution showed that diel vertical migration was obvious for the mesozooplankton abundance and biomass at S1 but was not apparent for the abundance at K2, because the dominant component was diurnally migrating species at S1 and overwintering species residing at mesopelagic depths at K2. The major components of diel migrants were copepods and euphausiids at S1 and only euphausiids at K2. Respiratory flux by the diurnally migrating mesozooplankton was estimated to be 2 mgC m⁻² day⁻¹ at S1 and 7 mgC m⁻² day⁻¹ at K2. The respiratory flux was equivalent to 131% of sedimentary fecal pellet flux at S1 and 136% of that at K2. Because pathways of downward carbon flux are facilitated by the mesozooplankton community, the actively transported carbon (respiration of dissolved inorganic carbon, excretion of dissolved organic carbon and egestion of fecal pellets at depth) might be larger during winter than the flux of sinking fecal pellets.     

The oxygen minimum zone in the Arabian Sea during 1995

This paper focuses on the characteristics of the oxygen minimum zone (OMZ) as observed in the Arabian Sea over the complete monsoon cycle of 1995. Dissolved oxygen, nitrite, nitrate and density values are used to delineate the OMZ, as well as identify regions where denitrification is observed. The suboxic conditions within the northern Arabian Sea are documented, as well as biological and chemical consequences of this phenomenon. Overall, the conditions found in the suboxic portion of the water column in the Arabian Sea were not greatly different from what has been reported in the literature with respect to oxygen, nitrate and nitrite distributions. Within the main thermocline, portions of the OMZ were found that were suboxic (oxygen less than ∼4.5 μM) and contained secondary nitrite maxima with concentrations that sometimes exceeded 6.0 μM, suggesting active nitrate reduction and denitrification. Although there may have been a reduction in the degree of suboxia during the Southwest monsoon, a dramatic seasonality was not observed, as has been suggested by some previous work. In particular, there was not much evidence for the occurrence of secondary nitrite maxima in waters with oxygen concentrations greater than 4.5 μM. Waters in the northern Arabian Sea appear to accumulate larger nitrate deficits due to longer residence times even though the denitrification rate might be lower, as evident in the reduced nitrite concentrations in the northern part of the basin. Organism distributions showed string relationships to the oxygen profiles, especially in locations where the OMZ was pronounced, but the biological responses to the OMZ varied with type of organism. The regional extent of intermediate nepheloid layers in our data corresponds well with the region of the secondary nitrite maximum. This is a region of denitrification, and the presence and activities of bacteria are assumed to cause the increase in particles. ADCP acoustic backscatter measurements show diel vertical migration of plankton or nekton and movement into the OMZ. Daytime acoustic returns from depth were strong, and the dawn sinking and dusk rise of the fauna were obvious. However, at night the biomass remaining in the suboxic zone was so low that no ADCP signal was detectable at these depths. There are at least two groups of organisms, one that stays in the upper mixed layer and another that makes daily excursions. A subsurface zooplankton peak in the lower OMZ (near the lower 4.5 μM oxycline) was also typically present; these animals occurred day and night and did not vertically migrate.     

Short-term environmental variability in cold-water coral habitat at Viosca Knoll,Gulf of Mexico

The Lophelia pertusa community at Viosca Knoll (VK826) is the most extensive found to date in the Gulf of Mexico. As part of a multi-disciplinary study, the physical setting of this area was described using benthic landers, CTD transects and remotely operated vehicle observations. The site was broadly characterised into three main habitats: (1) dense coral cover that resembles biogenic reef complexes, (2) areas of sediment, and (3) authigenic carbonate blocks with sparse coral and chemosynthetic communities. The coral communities were dominated by L. pertusa but also contained numerous solitary coral species. Over areas that contained L. pertusa, the environmental conditions recorded were similar to those associated with communities in the north-eastern Atlantic, with temperature (8.5–10.6 °C) and salinity (～35) falling within the known species niche for L. pertusa. However, dissolved oxygen concentrations (2.7–2.8 ml l^?1) and density (σ_Θ, 27.1–27.2 kg m^?3) were lower and mass fluxes from sediment trap data appeared much higher (4002–4192 mg m^?2 d^?1). Yet, this species still appears to thrive in this region, suggesting that L. pertusa may not be as limited by lower dissolved oxygen concentrations as previously thought. The VK826 site experienced sustained eastward water flow of 10–30 cm s^?1 over the 5-day measurement period but was also subjected to significant short-term variability in current velocity and direction. In addition, two processes were observed that caused variability in salinity and temperature; the first was consistent with internal waves that caused temperature variations of 0.8 °C over 5–11 h periods. The second was high-frequency variability (20–30 min periods) in temperature recorded only at the ALBEX site. A further pattern observed over the coral habitat was the presence of a 24 h diel vertical migration of zooplankton that may form part of a food chain that eventually reaches the corals. The majority of detailed studies concerning local environmental conditions in L. pertusa habitats have been conducted within the north-eastern Atlantic, limiting most knowledge of the niche of this species to a single part of an ocean basin. Data presented here show that the corals at VK826 are subjected to similar conditions in temperature, salinity, and flow velocity as their counterparts in the north-east Atlantic, although values for dissolved oxygen and density (sigma-theta: σ_Θ) are different. Our data also highlight novel observations of short-term environmental variability in cold-water coral habitat.