Zooplankton associated with the oxygen minimum zone system in the northern upwelling region of Chile during March 2000

Zooplankton in the coastal upwelling region off northern Chile may play a significant biogeochemical role by promoting carbon flux into the subsurface OMZ (oxygen minimum zone). This work identifies the dominant zooplankton species inhabiting the area influenced by the OMZ in March 2000 off Iquique (20°S, northern Chile). Abundance and vertical distribution studies revealed 17 copepod and 9 euphausiid species distributed between the surface and 600 m at four stations sampled both by day and by night. Some abundant species remained in the well-oxygenated upper layer (30 m), with no evidence of diel vertical migration, apparently restricted by a shallow (40–60 m) oxycline. Other species, however, were found closely associated with the OMZ. The large-sized copepod Eucalanus inermis was found below the oxycline and performed diel vertical migrations into the OMZ, whereas the very abundant Euphausia mucronata performed extensive diel vertical migrations between the surface waters and the core of the OMZ (200 m), even crossing it. A complete assessment of copepods and euphausiids revealed that the whole sampled water column (0–600 m) is occupied by distinct species having well-defined habitats, some of them within the OMZ. Ontogenetic migrations were evident in Eucalanidae and E. mucronata. Estimates of species biomass showed a substantial (>75% of total zooplankton biomass) daily exchange of C between the photic layer and the OMZ. Both E. inermis and E. mucronata can actively exchange about 37.8 g C m⁻² d⁻¹ between the upper well-oxygenated (0–60 m) layer and the deeper (60–600 m) OMZ layer. This migrant biomass may contribute about 7.2 g C m⁻² d⁻¹ to the OMZ system through respiration, mortality, and production of fecal pellets within the OMZ. This movement of zooplankton in and out of the OMZ, mainly as a result of the migratory behavior of E. mucronata, suggests a very efficient mechanism for introducing large amounts of freshly produced carbon into the OMZ system and should, therefore, be considered when establishing C budgets for coastal upwelling systems.     

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.     

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.     

Mesozooplankton of the Arabian Sea: Patterns influenced by seasons, upwelling, and oxygen concentrations

The intensive study of the Arabian Sea during the 1990s included mesozooplankton investigations by the Netherlands, United Kingdom, Pakistan, India, Germany and the United States. Several major discoveries resulted. First, the high biomass of mesozooplankton observed during the Northeast Monsoon season is sustained by primary productivity stimulated by convective mixing and by an active microbial loop. The apparent ‘paradox’ of high standing stocks of mesozooplankton coinciding with low standing stocks of phytoplankton thus was resolved. Second, the Southwest Monsoon (upwelling) season supports a burst of mesozooplankton growth, much of which is exported to the interior of the Arabian Sea by strong currents and eddy activity and to depth at the end of the season when diapause causes at least one very abundant copepod to leave the epipelagic zone. Third, the oxygen minimum zone severely restricts the vertical distribution of mesozooplankton in the eastern region of the Arabian Sea. The copepod that withstands conditions in the OMZ most readily, Pleuromamma indica, has increased in abundance over the past thirty years suggesting the OMZ may have grown in size and/or intensity in that time. Fourth, the Fall Intermonsoon and Northeast Monsoon seasons are characterized everywhere by increased abundance of the cyclopoid copepod genus, Oithona. Abundances of Oithona measured in the 1990s are much higher than those of the 1930s, suggesting food web alterations over the past half-century.     

Fine-scale vertical distribution of coastal and offshore copepods in the Golfo de Arauco, central Chile, during the upwelling season

In order to understand the mechanism by which zooplankters from different origins co-occur during the upwelling season within Golfo de Arauco, one of the most productive areas in central Chile, we assessed short term variations in the vertical distribution of the most abundant copepod species. Fine-scale, day and night vertical zooplankton sampling was done with a pump over 12 days in summer. The water column in the gulf consisted of three layers: Equatorial Subsurface Water of low dissolved oxygen content in the deeper part of the water column, strong temperature and oxygen gradients at mid-depth (15-25 m), and a layer of warmer, more oxygenated, less saline water at the surface. Copepods within the gulf originated from offshore, from the continental shelf, and from the coastal area. Most taxa showed distinctive vertical distributions. Three copepod groups were identified by their mean weighted depths of residence. One group included shallow residents found above the thermocline/oxycline (Acartia tonsa, Centropages brachiatus, Corycaeus sp., Paracalanus parvus, Oncaea sp.). A second group was comprised by species distributed at or below the thermocline/oxycline (Oithona sp., Oncaea conifera, Lucicutia sp., Metridia sp., Heterorhabdus papilliger). The third group was composed of vertical migrators that crossed the thermocline/oxycline (Calanus chilensis, Calanoides patagoniensis, Aetideus armatus, Pleuromamma piseki). In spite of their different vertical distribution ranges, the most abundant and frequent copepod species (P. parvus, C. chilensis, C. patagoniensis, C. brachiatus) share a common capacity to withstand wide ranges of oxygen concentration and temperature. This characteristic, along with the capacity to vary their life strategies under different environmental conditions, seems to facilitate the maintenance of large numbers of copepods in coastal waters along the Humboldt Current.     

Oxygen minimum zones in the eastern tropical Atlantic and Pacific oceans

Within the eastern tropical oceans of the Atlantic and Pacific basin vast oxygen minimum zones (OMZ) exist in the depth range between 100 and 900 m. Minimum oxygen values are reached at 300–500 m depth which in the eastern Pacific become suboxic (dissolved oxygen content <4.5 μmol kg⁻¹) with dissolved oxygen concentration of less than 1 μmol kg⁻¹. The OMZ of the eastern Atlantic is not suboxic and has relatively high oxygen minimum values of about 17 μmol kg⁻¹ in the South Atlantic and more than 40 μmol kg⁻¹ in the North Atlantic. About 20 (40%) of the North Pacific volume is occupied by an OMZ when using 45 μmol kg⁻¹ (or 90 μmol kg⁻¹, respectively) as an upper bound for OMZ oxygen concentration for ocean densities lighter than σ_θ < 27.2 kg m⁻³. The relative volumes reduce to less than half for the South Pacific (7% and 13%, respectively). The abundance of OMZs are considerably smaller (1% and 7%) for the South Atlantic and only 0% and 5% for the North Atlantic. Thermal domes characterized by upward displacements of isotherms located in the northeastern Pacific and Atlantic and in the southeastern Atlantic are co-located with the centres of the OMZs. They seem not to be directly involved in the generation of the OMZs.OMZs are a consequence of a combination of weak ocean ventilation, which supplies oxygen, and respiration, which consumes oxygen. Oxygen consumption can be approximated by the apparent oxygen utilization (AOU). However, AOU scaled with an appropriate consumption rate (aOUR) gives a time, the oxygen age. Here we derive oxygen ages using climatological AOU data and an empirical estimate of aOUR. Averaging oxygen ages for main thermocline isopycnals of the Atlantic and Pacific Ocean exhibit an exponential increase with density without an obvious signature of the OMZs. Oxygen supply originates from a surface outcrop area and can also be approximated by the turn-over time, the ratio of ocean volume to ventilating flux. The turn-over time corresponds well to the average oxygen ages for the well ventilated waters. However, in the density ranges of the suboxic OMZs the turn-over time substantially increases. This indicates that reduced ventilation in the outcrop is directly related to the existence of suboxic OMZs, but they are not obviously related to enhanced consumption indicated by the oxygen ages. The turn-over time suggests that the lower thermocline of the North Atlantic would be suboxic but at present this is compensated by the import of water from the well ventilated South Atlantic. The turn-over time approach itself is independent of details of ocean transport pathways. Instead the geographical location of the OMZ is to first order determined by: (i) the patterns of upwelling, either through Ekman or equatorial divergence, (ii) the regions of general sluggish horizontal transport at the eastern boundaries, and (iii) to a lesser extent to regions with high productivity as indicated through ocean colour data.     

Trophic structure of zooplankton in the Fram Strait in spring and autumn 2003

The trophic structure of zooplankton was investigated in Fram Strait (north western Svalbard) in spring and autumn of 2003. Depth-stratified zooplankton samples were collected at 12 stations on the shelf (200 m), across the shelf-slope (500 m) and over deep water (>750 m), using a Multiple Plankton Sampler equipped with 0.180-mm mesh size nets.Higher zooplankton abundance and estimated biomass were found in the shelf area. Abundance and biomass were two times higher in August, when sea-surface temperature was higher than in May. Herbivores dominated numerically in May, and omnivores in August, suggesting a seasonal sequence of domination by different trophic groups. Cirripedia nauplii and Fritillaria borealis prevailed in spring, whereas copepod nauplii and Calanus finmarchicus were numerically the most important herbivores in autumn. Small copepods, Oithona similis and Triconia borealis, were the most numerous omnivorous species in both seasons, but their abundances increased in autumn. Chaetognatha (mainly Eukrohnia hamata) accounted for the highest abundance and biomass among predatory taxa at all deep-water stations and during both seasons. Regarding vertical distribution, herbivores dominated numerically in the surface layer (0–20 m), and omnivores were concentrated somewhat deeper (20–50 m) during both seasons. Maximum abundance of predators was found in the surface layer (0–20 m) in spring, and generally in the 20–50 m layer in autumn. This paper presents the first comprehensive summary of the zooplankton trophic structure in the Fram Strait area. Our goals are to improve understanding of energy transfer through this ecosystem, and of potential climate-induced changes in Arctic marine food webs.     

Infaunal macrobenthos of the oxygen minimum zone on the Indian western continental shelf

The Arabian Sea is characterized by a mid‐depth layer of reduced dissolved oxygen (DO) concentration or oxygen minimum zone (OMZ ‐DO concentration <0.5 ml·l^?1) at ~150–1000 m depth. This OMZ results from the flux of labile organic matter coupled with limited intermediate depth water ventilation. Generally, benthic animals in the OMZ have morphological and physiological adaptations that maximize oxygen uptake in the limited oxygen availability. Characteristics of OMZ benthos have been described from only a few localities in the Arabian Sea. We measured the bottom water DO and studied the characteristics of infaunal macrobenthos of the Indian western continental shelf by collecting samples at 50, 100 and 200 m in depth from 7° to 22° N. The DO values observed at 200 m (0.0005–0.24 ml·l^?1) indicated that this area is lying within an OMZ. Five major taxa, namely Platyhelminthes, Sipunculoidea, Echiuroidea, Echinodermata and Cephalochordata were absent from the samples collected from this OMZ. In general, declines in total macrobenthic density and biomass and polychaete species richness and diversity were observed in this OMZ compared with the shallower depths above it. Community analyses of polychaetes revealed the dominance of species belonging to families Spionidae, Cirratulidae and Paraonidae in this OMZ. Low oxygen condition was more pronounced in the northern continental shelf edge (≤0.03 ml·l^?1), where the majority of spionids including Prionospio pinnata and cirratulids were absent; whereas amphipod, isopod and bivalve communities were not impacted.     

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.     

Calanoid copepod resting egg abundance and hatching success in the sediment of the Seine estuary (France)

Benthic, viable resting eggs of calanoid copepods were found for the first time in the Seine estuary (France) during July 2008. Vertical distribution of the resting eggs in the sediment was determined up to 10 cm depth. Hatching success of the eggs extracted from different 1-cm thick sediment layers was experimentally tested immediately after extraction and after a long refractory phase (i.e. 11 months) of storage at low temperature (4–5 °C). The hatching success of resting eggs obtained immediately after sediment incubation was lower (0.72%) than the value observed after 11 months (4.50%) with an overall hatching success of 2.37%. The marine, calanoid copepod Temora longicornis was the primary species to hatch from the eggs; however, the estuarine calanoid copepod Eurytemora affinis also hatched from resting eggs. The mean abundance of eggs found in sediment (1.42 × 10⁶ eggs m⁻²) was comparable to that reported for other marine and estuarine calanoid copepods. The Seine estuary sediment had a high variability of egg abundance (between 0.14 and 8.10 × 10⁷ eggs m⁻³) suggesting that the hydrodynamics of this macrotidal estuary are likely responsible for this variability. Significant sediment resuspension occurs in the Seine estuary during flood periods and spring tides leading to resting eggs to contribute along the year to the nauplii recruitment of calanoid copepods. On average, around 400,000 nauplii m⁻³ month⁻¹ of the main calanoid copepods can emerge from the surface layer sediment in the Seine estuary, suggesting that resting eggs could play an important role in the population dynamics of key calanoid copepods in the Seine estuary.     

Enhanced preservation of organic matter in sediments deposited within the oxygen minimum zone in the northeastern Arabian Sea

The presence of a strongly developed oxygen minimum zone (OMZ; [O₂]<2 μM) in the northeastern Arabian Sea affords the opportunity to investigate whether oxygen deficiency in bottom waters enhances the preservation of organic matter in the underlying sediments. We explored if the observed patterns of organic matter accumulation could be explained by differences in productivity, sedimentation rate, water depth, and mineral texture. The differences in the burial rates of organic matter in sediments deposited within or below the OMZ could not be explained on the basis of these factors. All collected evidence points to a coupling of low oxygen concentrations and enhanced organic matter preservation. Under more oxygenated conditions bioturbation as well as the presence of labile manganese and iron oxides are probably important factors for a more efficient microbially mediated degradation of organic matter. Pore water profiles of dissolved Mn²⁺ and Fe²⁺ show that reduction of manganese and iron oxides plays a minor role in sediments lying within the OMZ and a larger role in sediments lying below the OMZ.     

Parameterization of iron and manganese cycling in the Black Sea suboxic and anoxic environment

New and published data on the distribution and speciation of manganese and iron in seawater are analyzed to identify and parameterize major biogeochemical processes of their cycling within the suboxic (15.6σ_t16.2) and anoxic layers (σ_t16.2) of the Black Sea. A steady-state transport-reaction model is applied to reveal layering and parameterize kinetics of redox and dissolution/precipitation processes. Previously published data on speciation of these elements in seawater are used to specify the nature of the transformations. Two particulate species of iron (Fe(III) hydroxide and Fe(II) sulfide) are necessary to adequately parameterize the vertical profile of suspended iron, while three particulate species (hydrous Mn(IV) oxide, Mn(II) sulfide, and Mn(II) carbonate) are necessary to describe the profile of suspended manganese. In addition to such processes as mixing and advection, precipitation, sinking, and dissolution of manganese carbonate are found to be essential in maintaining the observed vertical distribution of dissolved Mn(II). These results are used to interpret the observed difference in the form of vertical distribution for dissolved Mn(II) and Fe(II). Redox transformations of iron and manganese are coupled via oxidation of dissolved iron by sinking suspended manganese at σ_t16.2±0.2 kg m⁻³. The particulate manganese, necessary for this reaction, is supplied through oxidation of dissolved Mn(II). The best agreement with observations is achieved when nitrate, rather than oxygen, is set to oxidize dissolved Mn(II) in the lower part of the suboxic layer (15.90σ_t16.2). The results support the idea that, after sulfides of these metals are formed, they sink with particulate organic matter. The sinking rates of the particles and specific rates of individual redox and dissolved-particulate transformations have been estimated by fitting the vertical profile of the net rate.     

The spatial arrangement of reefs alters the ecological patterns of fauna between interspersed algal habitats

Reef landscapes dominated by canopy-forming species are often irregular mosaics of habitats, with important influences on associated fauna. This study tested if differences in the ecological patterns of mobile fauna inhabiting interspersed (morphologically distinct) algal habitats were altered by the spatial arrangement of reefs of varying proximity to the shoreline. Specifically, prosobranch gastropods were used as models to test that: (1) there were differences in the ecological patterns (species composition and abundances) between three algal habitats (the kelp Ecklonia radiata, fucalean macroalgae, and erect red algae); (2) the magnitude of these differences depended on the position of reef lines (‘in-shore’ vs. ‘off-shore’); and (3) these effects were regionally consistent across a 4° latitudinal gradient (600 km of coastline) in Western Australia. The ecological patterns of algal-associated gastropods responded strongly to the presence of algal habitats with different physical structure at small spatial scales. Importantly, differences in assemblage structure (e.g. differences in total abundances) between habitats across the latitudinal gradient were especially accentuated on the in-shore reefs compared with the off-shore reefs, where a general amelioration of differences between habitats was observed, probably associated with a more widespread effect of stronger wave forces across habitats. Overall, red algae supported higher total abundances and species richness (per algal weight) compared to the other algal habitats, particularly on in-shore reefs. Patterns for individual species were considerably location-dependent, reflecting the natural variability of species across geographical gradients. In contrast, patterns at the assemblage-level were consistent, providing evidence for the existence of general rules underlying the assemblage-level organization of mobile invertebrates on subtidal reefs across this geographical gradient.     

Physical dynamics and biological implications of Cyclone Noah in the lee of Hawaii during E-Flux I

Quasi-synoptic observations of the horizontal and vertical structure of a cold-core cyclonic mesoscale eddy feature (Cyclone Noah) were conducted in the lee of Hawai’i from November 4–22, 2004 as part of the E-Flux interdisciplinary collaborative research program. Cyclone Noah appears to have spun up to the southwest of the ‘Alenuihaha Channel (between Maui and Hawai’i) as a result of strong and persistent northeasterly trade winds through the channel. Shipboard hydrographic surveys 2.5 months later suggest that Noah weakened and was in a hypothesized spin-down phase of its life cycle. Although the initial surface expression of Noah was limited in scale to 40 km in diameter and, as evidenced by surface temperatures, 2–3 °C cooler than the surrounding waters, depth profiles revealed a fully developed semi-elliptical shallow feature (200 m), 144 km long and 90 km wide (based on sigma-t=23 kg m⁻³) with tangential speeds of 40–80 cm s⁻¹, and substantial isopycnal doming. Potential vorticity distribution of Noah suggests that radial horizontal flow of the core water was inhibited from the surface to depths of 75 m, with high vorticity confined above the sigma-t=23.5 kg m⁻³ isopycnal surface. Upward displacements of isopycnal surfaces in the eddy's center (50 m) were congruent with enhanced pigment concentrations (0.50 mg m⁻³). Comparisons of the results obtained for E-Flux I (Noah) and E-Flux III (Opal) suggest that translation characteristics of cyclonic Hawaiian lee eddies may be important in establishing the biogeochemical and biological responses of the oligotrophic ocean to cyclonic eddies.     

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.     

Benthic foraminifera living in Gulf of Mexico bathyal and abyssal sediments: Community analysis and comparison to metazoan meiofaunal biomass and density

Benthic foraminiferal biomass, density, and species composition were determined at 10 sites in the Gulf of Mexico. During June 2001 and 2002, sediment samples were collected with a GoMex box corer. A 7.5-cm diameter subcore was taken from a box core collected at each site and sliced into 1-cm or 2-cm sections to a depth of 2 or 3 cm; the >63-μm fraction was examined shipboard for benthic foraminifera. Individual foraminifers were extracted for adenosine triphosphate (ATP) using a luciferin–luciferase assay, which indicated the total ATP content per specimen; that data was converted to organic carbon. Foraminiferal biomass and density varied substantially (2–53 mg C m⁻²; 3600–44,500 individuals m⁻², respectively) and inconsistently with water depth: although two 1000-m deep sites were geographically separated by only 75 km, the foraminiferal biomass at one site was relatively low (9 mg C m⁻²) while the other site had the highest foraminiferal biomass (53 mg C m⁻²). Although most samples from Sigsbee Plain (>3000 m) had low biomass, one Sigsbee site had >20 mg foraminiferal C m⁻². The foraminiferal community from all sites (i.e. bathyal and abyssal locales) was dominated by agglutinated, rather than calcareous or tectinous, species. Foraminiferal density never exceeded that of metazoan meiofauna at any site. Foraminiferal biomass, however, exceeded metazoan meiofaunal biomass at 5 of the 10 sites, indicating that foraminifera constitute a major component of the Gulf's deep-water meiofaunal biomass.     

Vertical and horizontal distribution of fluorescent dissolved organic matter in the Southern Ocean

The vertical and horizontal distribution of fluorescent dissolved organic matter (FDOM), determined by fluorescence intensity at 320 nm excitation and 420 nm emission, were clarified in nine stations on two transects at the Southern Ocean, including a subtropical, subantarctic, polar frontal and Antarctic zone. All vertical profiles of fluorescence intensity showed that levels were lowest in the surface waters, increased with increasing the depth in mid-depth waters ( 2000 m), and then stayed within a relatively narrow range from there to the bottom. Such vertical profiles of FDOM were similar to those of nutrients, but were adverse to dissolved oxygen. In water columns below the temperature-minimum subsurface water (dichothermal waters) in the Antarctic zone and below the winter mixed layer in the other zones, we determined the relationships of fluorescence intensity to concentrations of nutrients and apparent oxygen utilization (AOU) over the entire area of the present study, and found significant linear correlations between the levels of fluorescence intensity and nutrient concentrations (r =  0.70 and 0.71 for phosphate and nitrate + nitrite, respectively) and AOU (r = 0.91). From the strong correlation coefficient between fluorescence intensity and AOU, we concluded that FDOM in the Southern Ocean is formed in situ via the biological oxidation of organic matter. The regeneration of the nutrients/consumption of the oxygen/formation of FDOM was active in mid-depth waters. However, the correlations between fluorescence intensities and nutrients and AOU were different in the mid-depth water masses, Subantarctic Mode Water (SAMW), and Antarctic Intermediate Water (AAIW), indicating that the sources of organic matter responsible for FDOM formation were different. A considerable amount of FDOM in the SAMW is thought to be produced by the remineralization of DOM in addition to sinking particulate organic matter, while DOM is less responsible for FDOM formation in the AAIW.     

Dissolved manganese and silicic acid in the Columbia River plume: A major source to the California current and coastal waters off Washington and Oregon

The spatial distributions of dissolved manganese and nutrients were examined in the Columbia River plume off Oregon and Washington during the summer of 2004 and 2005 as part of the River Influence on Shelf Ecosystems (RISE) program. Factors influencing the hydrochemical characteristics of the freshly formed and aged Columbia River plume were investigated. Hydrographic data and nutrient concentrations were used to delineate three distinct water sources for the Columbia River Plume: California Current surface water, coastal upwelled water, and Columbia River water. The warm, intermediate salinity, nutrient poor California Current water contains low levels of dissolved manganese (< 5 nM) and silicic acid (< 5 μM), and is depleted in nitrate. The cold, high salinity, nutrient rich, freshly upwelled water is highly variable (2–20 nM) in dissolved manganese and can be as high as  45 μM in silicic acid and  30 μM nitrate. The variable Columbia River has summer temperatures ranging from  13 to 24 °C, high silicic acid concentrations (ranging from  120 to 200 μM), and lower nitrate concentrations (ranging from  2 to 20 μM). During the summer, the concentrations of silicic acid and dissolved manganese can exceed 100 μM and 200 nM, respectively, in near-field Columbia River plumes. These values are markedly greater than those of surface coastal waters (even during upwelling conditions). As the plume advects and mixes, the concentrations of these two constituents remain relatively high within plume waters. The concentrations of dissolved manganese in the near-field plume vary with tidal amplitude, exhibiting much higher concentrations for a given salinity during spring tides than during neap tides. For example, the Columbia River plume at a salinity of 20 has a concentration of dissolved manganese of  240 nM during spring tides, as compared to only  60 nM during low amplitude tides. Silicic acid concentrations in the near-field plume remain relatively constant throughout the tidal month. Calculations indicate there is roughly an equivalent yearly delivery of dissolved manganese and silicic acid to the coastal waters off Oregon and Washington by upwelled waters and by the Columbia River plume.     

Assessment of storage lipid accumulation patterns in eucalanoid copepods from the eastern tropical Pacific Ocean

Members of the copepod family Eucalanidae are widely distributed throughout the world׳s oceans and have been noted for their accumulation of storage lipids in high- and low-latitude environments. However, little is known about the lipid composition of eucalanoid copepods in low-latitude environments. The purpose of this study was to examine fatty acid and alcohol profiles in the storage lipids (wax esters and triacylglycerols) of Eucalanus inermis, Rhincalanus rostrifrons, R. nasutus, Pareucalanus attenuatus, and Subeucalanus subtenuis, collected primarily in the eastern tropical north Pacific near the Tehuantepec Bowl and Costa Rica Dome regions, noted for its oxygen minimum zone, during fall 2007 and winter 2008/2009. Adult copepods and particulate material were collected in the upper 50 m and from 200 to 300 m in the upper oxycline. Lipid profiles of particulate matter were generated to help ascertain information on ecological strategies of these species and on differential accumulation of dietary and modified fatty acids in the wax ester and triacylglycerol storage lipid components of these copepods in relation to their vertical distributions around the oxygen minimum zone. Additional data on phospholipid fatty acid and sterol/fatty alcohol fractions were also generated to obtain a comprehensive lipid data set for each sample. Rhincalanus spp. accumulated relatively large amounts of storage lipids (31–80% of dry mass (DM)), while E. inermis had moderate amounts (2–9% DM), and P. attenuatus and S. subtenuis had low quantities of storage lipid (0–1% DM). E. inermis and S. subtenuis primarily accumulated triacylglycerols (>90% of storage lipids), while P. attenuatus and Rhincalanus spp. primarily accumulated wax esters (>84% of storage lipids). Based on previously generated molecular phylogenies of the Eucalanidae family, these results appear to support genetic predisposition as a major factor explaining why a given species accumulates primarily triacylglycerols or wax esters, and also potentially dictating major fatty acid and alcohol accumulation patterns within the more highly modified wax ester fraction. Comparisons of fatty acid profiles between triacylglycerol and wax ester components in copepods with that in available prey suggested that copepod triacylglycerols were more reflective of dietary fatty acids, while wax esters contained a higher proportion of modified or de novo synthesized forms. Sterols and phospholipid fatty acids were similar between species, confirming high levels of regulation within these components. Similarities between triacylglycerol fatty acid profiles of E. inermis collected in surface waters and at >200 m depth indicate little to no feeding during their ontogenetic migration to deeper, low-oxygen waters.     

Temporal variability in oxygen and nutrient concentrations in the southern Aegean Sea and the Straits of the Cretan Arc

Nutrient and oxygen data collected in the southern Aegean Sea (Cretan Sea) and the straits of the Cretan Arc, during the four seasonal PELAGOS cruises in 1994–1995, are investigated and compared with data collected from 1987 to 1992 within the same area. During the cruises of the PELAGOS Project, nutrient enrichment of the intermediate layers of the Cretan Sea was observed, as a result of intrusion of ‘nutrient-rich, oxygen-poor’ Transition Mediterranean Water (TMW) compensating the Cretan Deep Water (CDW) outflow. TMW occupied the intermediate layers of the entire Cretan Sea. The concentrations of nutrients within this layer were often two times higher than those observed in the same area during previous studies undertaken before 1992 (increase 2.5 μmol/l of nitrate, 0.05 μmol/l of phosphate and 2.5μmol/l of silicate). The decrease of oxygen in this layer is about 0.8ml/l (35 μmol/l). Outflow of CDW occurs principally through the Antikithira and Kassos Straits (the two deeper straits of the Cretan Arc); it results in an increase of oxygen content but a decrease in the nutrient content of water in the deep and bottom layers outside the Cretan Sea. The major mesoscale features in the area have a major influence of the distributions and exchanges of nutrients and oxygen through the straits of the Cretan Arc. The surface and the intermediate layers were richer in nutrients and poorer in oxygen in spring (March 1994), than in autumn (September 1994).