Zooplankton distribution and migration in low-oxygen modewater eddies

The eastern tropical North Atlantic (ETNA) features a mesopelagic oxygen minimum zone (OMZ) at approximately 300-600 m depth. Here, oxygen concentrations rarely fall below 40 µmol O2 kg-1, but are expected to decline under future projections of global warming. The recent discovery of mesoscale eddies that harbour a shallow suboxic (<5 µmol O2 kg-1) OMZ just below the mixed layer could serve to identify zooplankton groups that may be negatively or positively affected by on-going ocean deoxygenation. In spring 2014, a detailed survey of a suboxic anticyclonic modewater eddy (ACME) was carried out near the Cape Verde Ocean Observatory (CVOO), combining acoustic and optical profiling methods with stratified multinet hauls and hydrography. The multinet data revealed that the eddy was characterized by an approximately 1.5-fold increase in total area-integrated zooplankton abundance. At nighttime, when a large proportion of acoustic scatterers is ascending into the upper 150 m, a drastic reduction in mean volume backscattering (Sv, shipboard ADCP, 75kHz) within the shallow OMZ of the eddy was evident compared to the nighttime distribution outside the eddy. Acoustic scatterers were avoiding the depth range between about 85 to 120 m, where oxygen concentrations were lower than approximately 20 µmol O2 kg-1, indicating habitat compression to the oxygenated surface layer. This observation is confirmed by time-series observations of a moored ADCP (upward looking, 300kHz) during an ACME transit at the CVOO mooring in 2010. Nevertheless, part of the diurnal vertical migration (DVM) from the surface layer to the mesopelagic continued through the shallow OMZ. Based upon vertically stratified multinet hauls, Underwater Vision Profiler (UVP5) and ADCP data, four strategies have been identified to be followed by zooplankton in response to the eddy OMZ: i) shallow OMZ avoidance and compression at the surface (e.g. most calanoid copepods, euphausiids), ii) migration to the shallow OMZ core during daytime, but paying O2 debt at the surface at nighttime (e.g. siphonophores, Oncaea spp., eucalanoid copepods), iii) residing in the shallow OMZ day and night (e.g. ostracods, polychaetes), and iv) DVM through the shallow OMZ from deeper oxygenated depths to the surface and back. For strategy i), ii) and iv), compression of the habitable volume in the surface may increase prey-predator encounter rates, rendering zooplankton and micronekton more vulnerable to predation and potentially making the eddy surface a foraging hotspot for higher trophic levels. With respect to long-term effects of ocean deoxygenation, we expect avoidance of the mesopelagic OMZ to set in if oxygen levels decline below approximately 20 µmol O2 kg-1. This may result in a positive feedback on the OMZ oxygen consumption rates, since zooplankton and micronekton respiration within the OMZ as well as active flux of dissolved and particulate organic matter into the OMZ will decline.

Combined effects of low pH and low oxygen on the early-life stages of the barnacle Balanus amphitrite

Ocean acidification (OA) is anticipated to interact with the more frequently occurring hypoxic conditions in shallow coastal environments. These could exert extreme stress on the barnacle-dominated fouling communities. However, the interactive effect of these two emerging stressors on early-life stages of fouling organisms remains poorly studied. We investigated both the independent and interactive effect of low pH (7.6 vs. ambient 8.2) and low oxygen (LO; 3 mg/l vs. ambient 5 mg/l) from larval development through settlement (attachment and metamorphosis) and juvenile growth of the widespread fouling barnacle, Balanus amphitrite. In particular, we focused on the critical transition between planktonic and benthic phases to examine potential limiting factors (i.e. larval energy storage and the ability to perceive cues) that may restrain barnacle recruitment under the interactive stressors. LO significantly slowed naupliar development, while the interaction with low pH (LO-LP) seemed to alleviate the negative effect. However, 20-50% of the larvae became cyprid within 4 d post-hatching, regardless of treatment. Under the two stressors interaction (LO-LP), the barnacle larvae increased their feeding rate, which may explain why their energy reserves at competency were not different from any other treatment. In the absence of a settlement-inducing cue, a significantly lower percentage of cyprids (15% lower) settled in LO and LO-LP. The presence of an inducing cue, however, elevated attachment up to 50-70% equally across all treatments. Post-metamorphic growth was not altered, although the condition index was different between LO and LO-LP treatments, potentially indicating that less and/or weaker calcified structures were developed when the two stressors were experienced simultaneously. LO was the major driver for the responses observed and its interaction with low pH should be considered in future studies to avoid underestimating the sensitivity of biofouling species to OA and associated climate change stressors.

Response of A. aurita ephyrae to three stressors: high-temperature, high-CO2 and low-oxygen

Global change is affecting marine ecosystems through a combination of different stressors such as warming, ocean acidification and oxygen depletion. Very little is known about the interactions among these factors, especially with respect to gelatinous zooplankton. Therefore, in this study we investigated the direct effects of pH, temperature and oxygen availability on the moon jellyfish Aurelia aurita, concentrating on the ephyral life stage. Starved one-day-old ephyrae were exposed to a range of pCO2 (400-4000 ppm) and three different dissolved oxygen levels (from saturated to hypoxic conditions), in two different temperatures (5 and 15 °C) for 7 days. Carbon content and swimming activity were analysed at the end of the incubation period, and mortality noted. General linearized models were fitted through the data, with the best fitting models including two- and three-way interactions between pCO2, temperature and oxygen concentration. The combined effect of the stressors was small but significant, with the clearest negative effect on growth caused by the combination of all three stressors present (high temperature, high CO2, low oxygen). We conclude that A. aurita ephyrae are robust and that they are not likely to suffer from these environmental stressors in a near future.

Stable oxygen isotope record and relative abundances of planktonic foraminifera of ODP Hole 117-728A

High resolution stratigraphy based on oxygen isotope ratios of the planktonic foraminifers Neogloboquadrina dutertrei (d'Orbigny), Globigeriniodes ruber (d'Orbigny), and Globigerina bulloides (d'Orbigny), magnetic susceptibility, and calcium carbonate content covers the sedimentary record of ODP Hole 728A drilled on the Oman Margin from approximately 10 k.y. to 525 k.y., comprising isotopic stages 1-13. Below stage 13 isotopic stage boundaries cannot be defined with certainty in our data. Sediment accumulation rates were calculated from the isotopic record of N. dutertrei by matching it with the age model SPECMAP curve. During the glacial periods sediment accumulation rates were higher than during the interglacial periods, reflecting increased input from the shelf during low-stands of sea level and increased eolian input. Periodograms for the past 524 k.y. on oxygen isotope records of N. dutertrei, G. ruber, and G. bulloides, on calcium carbonate content, magnetic susceptibility, and on a foraminiferal fragmentation record show powers matching the Milankovitch periodicities. High powers are concentrated around 103 k.y. In the spectra of oxygen isotope ratios of N. dutertrei, magnetic susceptibility, and foraminiferal fragmentation these are significant at the 80% confidence level with respect to a first order autoregressive model. Power concentrations near 43 k.y., matching obliquity, are present but subdued in all spectra. Power concentrations near 23 k.y., matching precession, are significant in the spectra of the oxygen isotope record of N. dutertrei, magnetic susceptibility, and calcium carbonate content record. Fragmentation of planktonic foraminifers increased during the interglacial periods. This is attributed to dissolution of the tests in an expanded oxygen minimum zone (OMZ), where undersaturation of calcium carbonate is caused by enhanced production in the euphotic zone, which would suggest stronger monsoonal induced upwelling during interglacial periods. Extension of the OMZ could also be increased by outflow of low oxygen marginal basin bottom water.

Upwelling and isolation in oxygen-depleted anticyclonic modewater eddies and implications for nitrate cycling

The physical (temperature, salinity, velocity) and biogeochemical (oxygen, nitrate) structure of an oxygen depleted coherent, baroclinic, anticyclonic mode-water eddy (ACME) is investigated using high-resolution autonomous glider and ship data. A distinct core with a diameter of about 70 km is found in the eddy, extending from about 60 to 200 m depth and. The core is occupied by fresh and cold water with low oxygen and high nitrate concentrations, and bordered by local maxima in buoyancy frequency. Velocity and property gradient sections show vertical layering at the flanks and underneath the eddy characteristic for vertical propagation (to several hundred-meters depth) of near inertial internal waves (NIW) and confirmed by direct current measurements. A narrow region exists at the outer edge of the eddy where NIW can propagate downward. NIW phase speed and mean flow are of similar magnitude and critical layer formation is expected to occur. An asymmetry in the NIW pattern is seen that possible relates to the large-scale Ekman transport interacting with ACME dynamics. NIW/mean flow induced mixing occurs close to the euphotic zone/mixed layer and upward nutrient flux is expected and supported by the observations. Combing high resolution nitrate (NO3-) data with the apparent oxygen utilization (AOU) reveals AOU:NO3- ratios of 16 which are much higher than in the surrounding waters (8.1). A maximum NO3- deficit of 4 to 6 µmol kg-1 is estimated for the low oxygen core. Denitrification would be a possible explanation. This study provides evidence that the recycling of NO3-, extracted from the eddy core and replenished into the core via the particle export, may quantitatively be more important. In this case, the particulate phase is of keys importance in decoupling the nitrogen from the oxygen cycling.

Records of past mid-depth ventilation: Cretaceous ocean anoxic event 2 vs. recent oxygen minimum zones

Present day oceans are well ventilated, with the exception of mid-depth oxygen minimum zones (OMZs) under high surface water productivity, regions of sluggish circulation, and restricted marginal basins. In the Mesozoic, however, entire oceanic basins transiently became dysoxic or anoxic. The Cretaceous ocean anoxic events (OAEs) were characterised by laminated organic-carbon rich shales and low-oxygen indicating trace fossils preserved in the sedimentary record. Yet assessments of the intensity and extent of Cretaceous near-bottom water oxygenation have been hampered by deep or long-term diagenesis and the evolution of marine biota serving as oxygen indicators in today's ocean. Sedimentary features similar to those found in Cretaceous strata were observed in deposits underlying Recent OMZs, where bottom-water oxygen levels, the flux of organic matter, and benthic life have been studied thoroughly. Their implications for constraining past bottom-water oxygenation are addressed in this review. We compared OMZ sediments from the Peruvian upwelling with deposits of the late Cenomanian OAE 2 from the north-west African shelf. Holocene laminated sediments are encountered at bottom-water oxygen levels of < 7 µmol/kg under the Peruvian upwelling and < 5 µmol/kg in California Borderland basins and the Pakistan Margin. Seasonal to decadal changes of sediment input are necessary to create laminae of different composition. However, bottom currents may shape similar textures that are difficult to discern from primary seasonal laminae. The millimetre-sized trace fossil Chondrites was commonly found in Cretaceous strata and Recent oxygen-depleted environments where its diameter increased with oxygen levels from 5 to 45 µmol/kg. Chondrites has not been reported in Peruvian sediments but centimetre-sized crab burrows appeared around 10 µmol/kg, which may indicate a minimum oxygen value for bioturbated Cretaceous strata. Organic carbon accumulation rates ranged from 0.7 and 2.8 g C /cm2 /kyr in laminated OAE 2 sections in Tarfaya Basin, Morocco, matching late Holocene accumulation rates of laminated Peruvian sediments under Recent oxygen levels below 5 µmol/kg. Sediments deposited at > 10 µmol/kg showed an inverse exponential relationship of bottom-water oxygen levels and organic carbon accumulation depicting enhanced bioirrigation and decomposition of organic matter with increased oxygen supply. In the absence of seasonal laminations and under conditions of low burial diagenesis, this relationship may facilitate quantitative estimates of palaeo-oxygenation. Similarities and differences between Cretaceous OAEs and late Quaternary OMZs have to be further explored to improve our understanding of sedimentary systems under hypoxic conditions.

Nitrogen cycling driven by organic matter export in the South Pacific oxygen minimum zone

Oxygen minimum zones are expanding globally, and at present account for around 20-40% of oceanic nitrogen loss. Heterotrophic denitrification and anammox-anaerobic ammonium oxidation with nitrite-are responsible for most nitrogen loss in these low-oxygen waters. Anammox is particularly significant in the eastern tropical South Pacific, one of the largest oxygen minimum zones globally. However, the factors that regulate anammox-driven nitrogen loss have remained unclear. Here, we present a comprehensive nitrogen budget for the eastern tropical South Pacific oxygen minimum zone, using measurements of nutrient concentrations, experimentally determined rates of nitrogen transformation and a numerical model of export production. Anammox was the dominant mode of nitrogen loss at the time of sampling. Rates of anammox, and related nitrogen transformations, were greatest in the productive shelf waters, and tailed off with distance from the coast. Within the shelf region, anammox activity peaked in both upper and bottom waters. Overall, rates of nitrogen transformation, including anammox, were strongly correlated with the export of organic matter. We suggest that the sinking of organic matter, and thus the release of ammonium into the water column, together with benthic ammonium release, fuel nitrogen loss from oxygen minimum zones.

(Table 2) Oxygen isotopic of composition of Pliocene to late Pleistocene planktonic foraminifera of the Mediterranean Sea

Stable isotopic and micropaleontological studies were made of selected sapropels (organic-rich sediments) deposited in the Mediterranean Sea during the last 5.0 m.y. to determine the processes responsible for their formation. Distinct isotopic and faunal changes occur across sapropels of late Pleistocene, early Pleistocene and latest Pliocene age, while smaller isotopic changes and more stable faunal assemblages are associated with the early and mid-late Pliocene sapropels. The large d18O depletions and euryhaline fauna associated with latest Pliocene-Pleistocene sapropels supports a density stratification model with a low salinity surface layer. In contrast, early Pliocene and mid-late Pliocene sapropels appear to have been formed as the result of sluggish circulation and low oxygen contents in bottom waters of the eastern Mediterranean due to the stable, warm climatic conditions of that time period.

Benthic foraminifera in Neogene laminated diatom ooze of ODP Hole 138-844C

Several widely correlatable intervals of laminated Thalassiothrix diatom mat deposits occur in Neogene sediments recovered from the eastern equatorial Pacific Ocean. The presence of laminated sediments in extensive areas of the deep open ocean floor raises fundamental questions concerning the cause of preservation of the laminations and the nature of the benthic environment during episodes of mat deposition. Traditional explanations for the preservation of laminations have centered on restriction of dissolved oxygen. Studies of benthic foraminifers through the laminated intervals show no evidence for an increase in absolute or relative abundance of species characteristic of a low oxygen environment, but rather a decrease in relative abundance of infaunal forms attesting to the impenetrability of the diatom meshwork formed by the interlocking Thalassiothrix frustules. These results support evidence from coring of the high tensile strength of the Thalassiothrix laminations suggesting that the diatom meshwork was of sufficient tensile strength and impenetrability to suppress infaunal benthic activity. Comparison of the relative abundances of foraminifers in the enclosing ôbackgroundö sediment of foraminifer nannofossil ooze and the laminated diatom oozes shows that some epifaunal species (e.g., Cibicides spp.) increase in relative abundance within the laminated sediment, whereas others (e.g., Epistominella exigua) show a marked decrease in relative abundance. Other species show more complex changes in abundance related to the occurrence of the laminated sediments, which may indicate a combination of controls that include the physical nature of the substrate and the amount of organic flux.

(Table 2) Similarity Indexes (Pearson's correlation coefficient) of the silicoflagellate assemblages of IODP Exp302

The silicoflagellate and ebridian assemblages in early middle Eocene Arctic cores obtained by IODP Expedition 302 (ACEX) were studied in order to decipher the paleoceanography of the upper water column. The assemblages in Lithologic Unit 2 (49.7-45.1 Ma), one of the biosiliceous intervals, were usually endemic as compared to the assemblages that occurred outside of the Arctic Ocean. The presence of these endemic assemblages is probably due to a unique environmental setting, controlled by the degree of mixing between the low-salinity Arctic waters and relatively high salinity waters supplied from outside the Arctic Ocean, such as the Atlantic and possibly the Western Siberian Sea. Using the basin-to-basin fractionation model, the early middle Eocene Arctic Ocean corresponds to an estuarine circulation type, which includes the modern-day Black Sea. The abundant down-core occurrence of ebridians strongly suggests the past presence of low-salinity waters, and may indicate that low oxygen concentrations prevailed in the euphotic layer, on the basis of the ecology of the modern ebridian Hermesinum adriaticum.