Strong Oxygen Minimum Zone Research Articles

Invertebrate trophic structure on marine ferromanganese and phosphorite hardgrounds

AbstractThe Southern California Borderland hosts a variety of geologic and oceanographic features that allow for diverse habitats to occur in a restricted region with a strong oxygen minimum zone (OMZ) and hard substrates. These include ferromanganese (FeMn) crusts and phosphorites targeted for deep‐seabed mining in other regions. Baseline studies regarding hardground macro‐ (> 0.3 mm) and megafaunal (> 2 cm) invertebrates are lacking, although they contribute to understanding nutrient cycling and resilience of deep‐sea communities to ocean deoxygenation, fishing, or mineral extraction. With the goal of understanding how substrate type, depth, and dissolved oxygen concentration influence invertebrate trophic structure, we surveyed δ13C and δ15N values of invertebrates on hard substrates on the Southern California Borderland margin along a depth gradient (120–2400 m) through the OMZ at inshore (< 100 km from shore) and offshore (100–250 km from shore) sites, using generalized additive models and community‐level metrics. Macrofaunal isotopic values correlate with substrate type, exhibiting higher trophic diversity on FeMn crusts and specialized communities on phosphorites. Megafaunal isotopic values correlate with proximity to shore; animals offshore seem to depend more on phytoplanktonic production than animals inshore. In general, δ15N increased with decreasing dissolved oxygen and increasing depth, possibly due to remineralization processes within the OMZ and with depth. We discuss how feeding modes and community composition might influence the observed patterns. This study elucidates the importance of the environmental context in shaping invertebrate trophic structure on continental margins and provides baseline knowledge that may be useful in regions where these minerals are targeted for extraction.

Mid-Holocene intensification of the oxygen minimum zone in the northeastern Arabian Sea

• The Arabian Sea has an intense oxygen minimum zone at intermediate water depths. • Lower sea level and weak OMZ conditions in the study area existed during the early Holocene . • The rise in sea level and increased surface water productivity intensified OMZ during the middle Holocene . • The enhanced sub-surface denitrification in the OMZ increases the nitrous oxide emission. The Arabian Sea is characterized by a strong Oxygen Minimum Zone (OMZ) bearing sub-oxic conditions at the intermediate water depth. We analyzed a sediment core near the upper margin of OMZ (174 m water depth) from offshore Saurashtra, northeastern (NE) Arabian Sea to reconstruct multi-proxy biogeochemical response in the area during the Early-Middle Holocene (∼10–4 ka before present). The results indicate lower foraminiferal productivity (both benthic and planktic) and weak sub-surface denitrification causing mild OMZ conditions at the study site during the early Holocene (∼10–8 ka). Subsequently, an increased foraminiferal productivity and sub-surface (both the water column and sediment) denitrification in the area led to intensified OMZ conditions during the mid-Holocene (after ∼8 ka). Our results corroborate other denitrification records from the Arabian Sea, suggesting the large-scale changes in Arabian Sea OMZ during the mid-Holocene. We suggest that the increased nutrient supply with significant freshwater input may have enhanced the primary productivity and further resulted in the intensified OMZ and escalated outgassing of N 2 O from the area during the mid-Holocene.

Marine biodiversity from zero to a thousand meters at Clipperton Atoll (Île de La Passion), Tropical Eastern Pacific.

Clipperton Atoll (Île de La Passion) is the only atoll in the Tropical Eastern Pacific (TEP) ecoregion and, owing to its isolation, possesses several endemic species and is likely an important stepping stone between Oceania, the remainder of the TEP, including other oceanic islands and the west coast of Central America. We describe the biodiversity at this remote atoll from shallow water to depths greater than one thousand meters using a mixture of technologies (SCUBA, stereo baited remote underwater video stations, manned submersible, and deep-sea drop cameras). Seventy-four unique taxa of invertebrates were identified during our expedition. The majority (70%) of these taxa were confined to the top 400 m and consisted mostly of sessile organisms. Decapod crustaceans and black corals (Antipatharia) had the broadest depth ranges, 100–1,497 m and 58–967 m, respectively. Decapods were correlated with the deepest depths, while hard corals were correlated with the shallow depths. There were 96 different fish taxa from 41 families and 15 orders, of which 70% were restricted to depths <200 m. While there was a decreasing trend in richness for both fish and invertebrate taxa with depth, these declines were not linear across the depth gradient. Instead, peaks in richness at ∼200 m and ∼750 m coincided with high turnover due to the appearance of new taxa and disappearance of other taxa within the community and is likely associated with the strong oxygen minimum zone that occurs within the region. The overall depth effect was stronger for fishes compared with invertebrates, which may reflect ecological preferences or differences in taxonomic resolution among groups. The creation of a no-take marine reserve 12 nautical miles around the atoll in 2016 will help conserve this unique and relatively intact ecosystem, which possesses high predator abundance.

Fish debris in sediments from the last 25 kyr in the Humboldt Current reveal the role of productivity and oxygen on small pelagic fishes

Upwelling of cold, nutrient-rich water from the oxygen minimum zone (OMZ) off Peru sustains the world’s highest production of forage fish, mostly composed of anchovy (Engraulis ringens). However, the potential impacts of climate change on upwelling dynamics and thus fish productivity in the near future are uncertain. Here, we reconstruct past changes in fish populations during the last 25,000 years to unravel their response to changes in OMZ intensity and productivity. We quantified and identified fish scales and bones deposited in laminated sediments from Pisco (Peru) with an average sampling resolution of 20.4 years (±7.1). The records span the Last Glacial Maximum to the recent Holocene and thus encompass a variety of combinations of productivity, oxygen, and global temperature. Our results reveal that productivity appears to be the main factor controlling small pelagic fish abundance, while sub-surface oxygenation affects mainly anchovy and likely sardine populations. Lower productivity and higher oxygen concentrations during the glacial resulted in lower total fish productivity, whereas higher productivity and a stronger OMZ in some time intervals during the Holocene resulted in higher fish abundances. A variety of different conditions between these two oceanographic end members indicate preferred environmental conditions for a variety of small pelagic fishes. There is no evidence in our record for an out of phase relationship between anchovy and sardine at the timescales examined in the present study. Anchovy have been the predominant small pelagic fish throughout the record, at least over centennial to millennial timescales. Its abundance reached a maximum during the Current Warm Period, an era characterized by high productivity and intense OMZ conditions. Thus, industrial fisheries developed during a period of exceptional productivity in relation to that of the last 25 kyr. The records reveal that dramatic decreases in pelagic fish abundances have occurred in response to past large-scale climate changes than those observed in the instrumental period, which suggests that future climate change may result in substantial changes in ecosystem structure.

A 43 kyr record of protist communities and their response to oxygen minimum zone variability in the Northeastern Arabian Sea

An extensive oxygen minimum zone (OMZ) occurs in the northeastern (NE) Arabian Sea where sedimentary records show evidence of alternating strong and weak OMZs that correlate with North Atlantic climate variability during the last glacial–interglacial cycle. OMZs are expanding world-wide, but information on long-term OMZ-ecosystem interactions is mainly limited to fossilized species, notably foraminifera. Here, we provide a first comprehensive ancient sedimentary DNA record of both fossilizing and non-fossilizing protists and their response to OMZ variability in the NE Arabian Sea over the last 43 ka. Protist communities changed significantly during strong vs. weak OMZ conditions coincident with interstadials and stadials respectively. Dinoflagellates were identified as significant indicator taxa for strong OMZs during glacial as well as interglacial interstadials, whereas diatoms were significant indicators for strong OMZs only during glacial interstadials. The chlorophyte Chlorella was found to be the main phototrophic protist in nutrient-depleted surface waters during glacial stadials. Notably, strong OMZ conditions shaped past protist communities by creating isolated habitats for those capable of sustaining oxygen depletion either by adapting a parasitic life cycle (e.g. apicomplexans) or by establishing mutualistic connections with others (e.g. radiolarians and mixotrophic dinoflagellates) or by forming cysts (e.g. colpodeans). Notably, a long-term increase in eutrophication and a decrease in the diatom/dinoflagellate ratio was observed during the late Holocene favoring the pelagic component of the marine food web. A similar scenario could be expected in the context of predicted worldwide expansion of coastal OMZs associated with global climate change.

Changes in tropical Atlantic surface-water environments inferred from late Albian planktic foraminiferal assemblages (ODP Site 1258, Demerara Rise)

Abundance patterns of planktic and benthic foraminifera from the middle to upper Albian of tropical Atlantic Ocean Drilling Program Site 1258 (Demerara Rise) display a pronounced shift between two clearly distinguishable assemblages: (1) a foraminiferal fauna dominated by coiled taxa, which is inferred to represent a deep and/or weak oxygen minimum zone, and (2) abundant clavate and biserial taxa, inferred to record a shallow and/or strong oxygen minimum zone at Demerara Rise. The entire sequence is rich in organic matter (3–6 wt% TOC) of a marine phytoplankton origin (δ13C ∼ −28‰), consistent with a highly productive region. Both the high productivity and changes in foraminiferal assemblages are consistent with changes in the mean latitudinal position of the Intertropical Convergence Zone (ITCZ) over the South American continent. Movements of the ITCZ are proposed to reduce the amount of runoff during the 'clavate-dominated' cycles, leading to the intensification of upwelling and therefore a stronger oxygen minimum zone compared to the intervals dominated by coiled foraminifera. We have previously found a similar hydrographic cycle in Turonian to Coniacian sections from the same area suggesting that this climate cyclicity and palaeogeography was long-lived off NE South America (∼15 million years). Due to the stratigraphic position of the interval with high abundances of clavate planktic foraminifera and the inferred strong oxygen minimum zone, it is tentatively proposed that this interval represents a local expression of the OAE 1c event within the long-living Demerara Rise upwelling system.

Benthic foraminiferal shell weight: Deglacial species-specific responses from the Santa Barbara Basin

Abstract Here we present a record of size-normalized shell weight for four species of benthic foraminifera through a period of rapid environmental change during the most recent deglaciation (Santa Barbara Basin, CA). A strong Oxygen Minimum Zone (OMZ), the product of high surface productivity and poor ventilation, characterizes the eastern Pacific; this subsurface zone is mechanistically coupled with high concentrations of dissolved inorganic carbon. The OMZ migrated vertically during warming of the last deglaciation, leading to rapid shifts in the oxygenation and inorganic carbon system of the benthos. The size-normalized weight (SNW) of benthic foraminifers Uvigerina peregrina , Bolivina interjuncta , and Bolivina tumida reflects only the broad trends of the vertical migration of the OMZ, and inorganic carbon system, overshadowed by clear species-specific trends. The relative importance of OMZ migrations versus other environmental variables and optimal growth conditions differs across species of benthic foraminifera. In U. peregrina , SNW primarily peaks with foraminiferal density and increased abundance of that species, while B. interjuncta and B. tumida increase in SNW with a shrinking of the OMZ (and carbon maximum) in the late Holocene. Bolivina argentea shows no long-term trends in SNW potentially due to its ability to migrate through the sediment. Our results suggest that, while inorganic carbon and dissolved oxygen may play a role in determining shell weight across species of benthic foraminifera, neither parameter alone is responsible for changes in benthic foraminiferal shell weight in the fossil record.

Redox chemistry of West Antarctic Peninsula margin surface sediments

Continental margin sediments are commonly studied using trace metal enrichments as proxies for characterizing their modern redox conditions and tracking past changes in bottom water ventilation and marine primary productivity. Currently little is known about the sedimentary redox history of the continental shelf west of the Antarctic Peninsula. Yet, environmental conditions in the Antarctic Peninsula region have changed rapidly in recent decades in response to global warming. Paleoclimate archives including bulk sediment trace metal enrichments can provide insight into both past and future environmental changes in this climatically sensitive region, and characterization of redox conditions in surface (modern) sediments is essential for establishing a framework for paleoredox interpretations. In this study we measured concentrations of trace metals (Ag, Cd, Re, and Mo) and productivity proxies (total Ba, organic carbon, and biogenic silica) in surface sediments from the Marguerite Bay, Gerlache Strait, and Bransfield Strait areas. Proxy concentrations suggest that sediments are generally suboxic due to seasonally high export of organic carbon from surface waters. Comparison of West Antarctic Peninsula trace metal/Al ratios to trace metal data from continental margins outside the Antarctic region demonstrates that although the West Antarctic Peninsula water column differs from other locations (e.g., it lacks a strong oxygen minimum zone), sedimentary redox chemistry is similarly controlled by organic matter decomposition, through slow recycling of highly seasonal export production falling through a cold water column.

Environmental Physiology of the Jumbo Squid,Dosidicus gigas(d'Orbigny, 1835) (Cephalopoda: Ommastrephidae): Implications for Changing Climate*

Dosidicus gigas (d'Orbigny, 1835) is a large, active squid that undergoes a diel vertical migration in the Eastern Tropical and Temperate Pacific. It is a voracious predator on zooplankton and micronekton and supports a large fishery. It is further preyed upon by large vertebrate predators, including whales. Its horizontal distribution is closely tied to productive upwelling regions that are characterized by strong oxygen minimum zones (OMZs). The apparent association with extreme hypoxia is surprising given its large size and high oxygen demand. As part of its daily vertical migration, D. gigas experiences daily temperature changes of 15–20°C, oxygen partial pressures ranging from near anoxia (< 0.8 kPa) to air-saturation (21 kPa) and pH changes from ∼8.1 to < 7.6 at depth. Oxygen minimum zones are believed to be expanding due to climate change, with minimum oxygen levels in the core of the OMZ declining and the low oxygen horizon shoaling. Simultaneously, surface waters are becoming more acidic and temperatures are rising. Here I review the extensive studies of this species that have been conducted over the past decade. D. gigas has a high affinity respiratory protein in the blood that supports a low critical oxygen partial pressure (3.8 kPa at 20 °C) and aerobic survival at night in the upper 200 meters of the water column. A pronounced pH- and temperature sensitivity of oxygen binding promotes oxygen transport across a depth range and in support of high rates of oxygen utilization but may impose constraints on high-temperature and CO2 tolerance. At its deeper, colder daytime habitat depth, D. gigas undergoes a pronounced metabolic suppression. Reduced activity levels and an apparent suspension of transcription and translation contribute to a ∼80% reduction in oxygen demand under 1% oxygen (0.8 kPa at 10 °C). Anaerobic metabolic pathways contribute some energy under these conditions. This metabolic suppression likely limits feeding at depth. Sub-critical oxygen levels, rather than temperature, predator avoidance or prey availability, appear to set the daytime depth distribution. Thus, expanding oxygen minimum zones will alter the daytime depth of peak abundance for these squids while ocean acidification and warming may impose a shallow ceiling above which squid performance is limited. The role of climate change in setting the vertical and horizontal distribution of the species is discussed.

Ocean biogeochemistry in the warm climate of the late Paleocene

Abstract. The late Paleocene is characterized by warm and stable climatic conditions that served as the background climate for the Paleocene–Eocene Thermal Maximum (PETM, ~55 million years ago). With respect to feedback processes in the carbon cycle, the ocean biogeochemical background state is of major importance for projecting the climatic response to a carbon perturbation related to the PETM. Therefore, we use the Hamburg Ocean Carbon Cycle model (HAMOCC), embedded in the ocean general circulation model of the Max Planck Institute for Meteorology, MPIOM, to constrain the ocean biogeochemistry of the late Paleocene. We focus on the evaluation of modeled spatial and vertical distributions of the ocean carbon cycle parameters in a long-term warm steady-state ocean, based on a 560 ppm CO2 atmosphere. Model results are discussed in the context of available proxy data and simulations of pre-industrial conditions. Our results illustrate that ocean biogeochemistry is shaped by the warm and sluggish ocean state of the late Paleocene. Primary production is slightly reduced in comparison to the present day; it is intensified along the Equator, especially in the Atlantic. This enhances remineralization of organic matter, resulting in strong oxygen minimum zones and CaCO3 dissolution in intermediate waters. We show that an equilibrium CO2 exchange without increasing total alkalinity concentrations above today's values is achieved. However, consistent with the higher atmospheric CO2, the surface ocean pH and the saturation state with respect to CaCO3 are lower than today. Our results indicate that, under such conditions, the surface ocean carbonate chemistry is expected to be more sensitive to a carbon perturbation (i.e., the PETM) due to lower CO32− concentration, whereas the deep ocean calcite sediments would be less vulnerable to dissolution due to the vertically stratified ocean.

Benthic foraminiferal diversity response to the climate induced changes in the eastern Arabian Sea oxygen minimum zone during the last 30 ka BP

A high resolution record of deep sea benthic foraminiferal diversity variations in the eastern Arabian Sea for the last 30kaBP was obtained from two sediment cores (SK17 and MD131) retrieved from the present day Oxygen Minimum Zone (OMZ) in the Indian margin off Goa. The benthic foraminiferal diversity is represented in terms of Sander's rarefaction number (S100), Shannon Wiener Index [H(S)], Equitability (E′) and Alpha Index (α). Records of diversity indices exhibit millennial scale changes during the late glacial and deglacial periods, corresponding to the Northern Hemisphere climatic events. We compared the faunal diversity with proxy records of primary productivity (Corg %) and bottom water oxygen (low-O2 taxa %). We suggest that benthic foraminiferal diversity in the eastern Arabian Sea OMZ is largely controlled by the primary productivity induced organic carbon flux and strength of bottom water oxygenation. The less diverse fauna along with increased percentages of Corg and low- O2 taxa during the last glacial maximum (18–22.5kaBP) suggest eutrophic and oxygen-poor benthic environment, attributed mainly to a strong OMZ associated with intense winter monsoon wind induced productivity and a weak deep ocean circulation. The intervals of distinct increase in diversity closely correspond with North Atlantic cold Heinrich events, when eastern Arabian Sea experienced significant declines in monsoon driven productivity and better deep sea ventilation due to enhanced inflow of Antarctic Intermediate Water (AAIW).

Oxygen decreases and variability in the eastern equatorial Pacific

Observations indicate increasingly large and strong oxygen minimum zones (OMZs) in the tropical Pacific over recent decades. Here we report on oxygen decreases and variability within the eastern equatorial Pacific OMZ. We construct time series from historical and profiling float oxygen data and analyze data from repeat hydrographic sections at 110°W and 85°50′W. Historical data are quite sparse for constructing oxygen time series, but floats with oxygen sensors prove to be good tools to fill measurement gaps in later parts of these time series. In the region just south of the equator a time series over the last 34 years reveals that oxygen decreases from 200 to 700 m at a rate between 0.50 and 0.83 μmol kg−1 yr−1. This strong decrease seems to be related to changes in the Pacific Decadal Oscillation (PDO). Oscillations on shorter time scales (e.g., an El Niño signal in the upper 350 m) are superimposed upon this trend. In the section data, a general trend of decreasing oxygen is present below the surface layer. While velocity differences appear related to oxygen differences in the equatorial channel, there is less correlation elsewhere. Contrasting with long‐term trend computations, the trends derived from two repeat sections are obscured by the influence of seasonal and longer‐term variability. Multidecadal variability (e.g., PDO) has the strongest influence on long‐term trends, while El Niño, isopycnal heave, current variability, seasonal cycles, and temperature changes are less important.

Late Quaternary changes in surface productivity and oxygen minimum zone (OMZ) in the northwestern Arabian Sea: Micropaleontologic and sedimentary record at ODP site 728A

Changes in the abundance of selected planktic foraminiferal species and some sedimentological parameters at ODP site 728A were examined to understand the fluctuations in the surface productivity and deep sea oxygenation in the NW Arabian Sea during last ∼540 kyr. The increased relative abundances of high fertility taxa, i.e., Globigerinita glutinata and Globigerina bulloides mainly during interglacial intervals indicate intense upwelling. Strong SW summer monsoon probably increased the upwelling in the western Arabian Sea during interglacial intervals and caused high surface productivities due to the lateral transport of eutrophic waters. Most of the glacial periods (i.e., MIS 2, 4, 6, 8 and 12) are characterized by higher relative abundances of Neogloboquadrina pachyderma and Neogloboquadrina dutertrei associated with Globigerinoides ruber. The more stratified condition and deep mixed layer due to increased NE winter monsoon are mainly responsible for the higher relative abundances of N. pachyderma during glacial periods. Some of the glacial intervals (i.e., MIS 6 and 8) are also characterized by pteropod spikes reflecting deepening of aragonite compensation depth (ACD) and relatively less intense oxygen minimum zone (OMZ) in this region due to deep sea mixing and thermocline ventilation, and relatively less intense surface productivity during winter monsoon. The interglacial periods are largely devoid of pteropod shells indicating more aragonite dissolution due to increased intensity of OMZ in the northwestern Arabian Sea. In general, the interglacial periods are characterized by low sediment accumulation rates than the glacial intervals. On an average, the total biogenic carbonate percentages were higher during interglacial and during periods of higher surface productivity. Most terrigenous material was trapped on shelf during intervals of high sea level stands of interglacial, whereas more erosion of shelf increased the sedimentation rates during glacial periods. In addition, the fragmentation record may be the result of changes in intensity and vertical distribution of the OMZ with time. During glacial intervals, the lower boundary of the OMZ probably was in a shallower position than during interglacial periods, when dissolution increased as a result of higher organic production. The higher rates of sinking organic matter result into a stronger OMZ as oxygen is used to disintegrate the organic matter. This process lowers the p H of water which results into increased dissolution of calcium carbonate.

Faunal responses to oxygen gradients on the Pakistan margin: A comparison of foraminiferans, macrofauna and megafauna

The Pakistan Margin is characterised by a strong mid-water oxygen minimum zone (OMZ) that intercepts the seabed at bathyal depths (150–1300 m). We investigated whether faunal abundance and diversity trends were similar among protists (foraminiferans and gromiids), metazoan macrofauna and megafauna along a transect (140–1850 m water depth) across the OMZ during the 2003 intermonsoon (March–May) and late/post-monsoon (August–October) seasons. All groups exhibited some drop in abundance in the OMZ core (250–500 m water depth; O2: 0.10–0.13 mL L?1=4.46–5.80 ?M) but to differing degrees. Densities of foraminiferans >63 ?m were slightly depressed at 300 m, peaked at 738 m, and were much lower at deeper stations. Foraminiferans >300 ?m were the overwhelmingly dominant macrofaunal organisms in the OMZ core. Macrofaunal metazoans reached maximum densities at 140 m depth, with additional peaks at 850, 940 and 1850 m where foraminiferans were less abundant. The polychaete Linopherus sp. was responsible for a macrofaunal biomass peak at 950 m. Apart from large swimming animals (fish and natant decapods), metazoan megafauna were absent between 300 and 900 m (O2 0.2 mL L?1=8.92 ?M). The progressively deeper abundance peaks for foraminiferans (>63 ?m), Linopherus sp. and ophiuroids probably represent lower OMZ boundary edge effects and suggest a link between body size and tolerance of hypoxia. Macro- and megafaunal organisms collected between 800 and 1100 m were dominated by a succession of different taxa, indicating that the lower part of the OMZ is also a region of rapid faunal change. Species diversity was depressed in all groups in the OMZ core, but this was much more pronounced for macrofauna and megafauna than for foraminiferans. Oxygen levels strongly influenced the taxonomic composition of all faunal groups. Calcareous foraminiferans dominated the seasonally and permanently hypoxic sites (136–300 m); agglutinated foraminiferans were relatively more abundant at deeper stations where oxygen concentrations were >0.13 mL L?1(=5.80 ?M). Polychaetes were the main macrofaunal taxon within the OMZ; calcareous macrofauna and megafauna (molluscs and echinoderms) were rare or absent where oxygen levels were lowest. The rarity of larger animals between 300 and 700 m on the Pakistan Margin, compared with the abundant macrofauna in the OMZ core off Oman, is the most notable contrast between the two sides of the Arabian Sea. This difference probably reflects the slightly higher oxygen levels and better food quality on the western side.

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 m 2 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.

Late Quaternary changes in intermediate water oxygenation and oxygen minimum zone, northern Japan: A benthic foraminiferal perspective

A strong oxygen minimum zone (OMZ) currently exists at upper intermediate water depths on the northern Japanese margin, NW Pacific. The OMZ results largely from a combination of high surface water productivity and poor ventilation of upper intermediate waters. We investigated late Quaternary history (last 34 kyr) of ocean floor oxygenation and the OMZ using quantitative changes in benthic foraminiferal assemblages in three sediment cores taken from the continental slope off Shimokita Peninsula and Tokachi, northern Japan, at water depths between 975 and 1363 m. These cores are well located within the present‐day OMZ, a region of high surface water productivity, and in close proximity to the source region of North Pacific Intermediate Water. Late Quaternary benthic foraminiferal assemblages experienced major changes in response to changes in dissolved oxygen concentration in ocean floor sediments. Foraminiferal assemblages are interpreted to represent three main groups representing oxic, suboxic, and dysoxic conditions. Assemblage changes in all three cores and hence in bottom water oxygenation coincided with late Quaternary climatic episodes, similar to that known for the southern California margin. These episodes, in turn, are correlated with orbital and millennial climate episodes in the Greenland ice core including the last glacial episode, Bølling‐Ållerød (B/A), Younger Dryas, Preboreal (earliest Holocene), early Holocene, and late Holocene. The lowest oxygen conditions, marked by dysoxic taxa and laminated sediments in one core, occurred during the B/A and the Preboreal intervals. Suboxic taxa dominated mainly during the last glacial, the Younger Dryas, and most of the Holocene. Dysoxic conditions during the B/A and Preboreal intervals in this region were possibly caused by high surface water productivity at times of reduced intermediate ventilation in the northwestern Pacific. Remarkable similarities are evident in the late Quaternary sequence of benthic foraminiferal assemblage change between the two very distant continental margins of northern Japan and southern California. The oscillations in OMZ strength, reflected by these faunal changes, were widespread and apparently synchronous over wide areas of the North Pacific, reflecting broad changes in intermediate water ventilation and surface ocean productivity closely linked with late Quaternary climate change on millennial and orbital timescales.

South Equatorial Current (SEC) driven changes at DSDP Site 237, Central Indian Ocean, during the Plio-Pleistocene: Evidence from Benthic Foraminifera and Stable Isotopes

This study attempts to analyse paleoceanographic changes in the Central Indian Ocean (Deep Sea Drilling Project Site 237), linked to monsoon variability as well as deep-sea circulation during the Plio-Pleistocene. We used factor and cluster analyses of census data of the 34 most dominant species of benthic foraminifera that enabled us to identify five biofacies: Astrononion umbilicatulum– Uvigerina proboscidea (Au–Up), Pullenia bulloides– Bulimina striata (Pb–Bs), Globocassidulina tumida– Nuttallides umbonifera (Gt–Nu), Gyroidinoides nitidula– Cibicides wuellerstorfi (Gn–Cw) and Cassidulina carinata– Cassidulina laevigata (Cc–Cl) biofacies. Knowledge of the environmental preferences of modern deep-sea benthic foraminifera helped to interpret the results of factor and cluster analyses in combination with oxygen and carbon isotope values. The biofacies indicative of high surface productivity, resulting from a stronger South Equatorial Current (Au–Up and Pb–Bs biofacies), dominate the early Pliocene interval (5.6–4.5 Ma) of global warmth. An intense Indo-Pacific ‘biogenic bloom’ and strong Oxygen Minimum Zone extended to intermediate depths (∼1000–2000 m) over large parts of the Indian Ocean in the early Pliocene. Since 4.5 Ma, the food supply in the Central Indian Ocean dropped and fluctuated while deep waters were corrosive (biofacies Gt–Nu, Gn–Cw). The Pleistocene interval is characterized by an intermediate flux of organic matter (Cc–Cl biofacies).

Glacial‐Holocene δ15N record from the Gulf of Tehuantepec, Mexico: Implications for denitrification in the eastern equatorial Pacific and changes in atmospheric N2O

The intensity of denitrification in the Gulf of Tehuantepec, along the Mexican margin of the eastern tropical North Pacific (ETNP), was reconstructed for the last 23,000 years using the nitrogen isotopic composition of sediment organic matter (δ15Norg) in conjunction with other geochemical tracers of water column productivity. During the last glacial period, δ15Norg values ranged from 6.5 to 8.6‰ with values increasing between 23 and 17 ka. Deglaciation was marked by an abrupt increase in δ15Norg with maximum values (9.9‰) occurring during the Bølling/Allerød period and the earliest Holocene. A decline in δ15Norg values occurred over the last 10,000 years, with late Holocene values being similar to those of the glacial period. Our data suggest that denitrification in this region was minimal during the last glacial and is consistent with the deposition of bioturbated sediments at the study site. The intensity of denitrification increased dramatically during deglaciation and the early Holocene, signaling the presence of a strong oxygen minimum zone. This period is also marked by the deposition of laminated sediments in the region. The δ15Norg records display similar temporal trends to those seen in the Greenland ice core record of atmospheric N2O concentrations. At present, denitrification within the eastern tropical Pacific and Arabian Sea accounts for a major portion of the global marine loss of fixed nitrogen, and a significant fraction of the ocean‐atmosphere N2O flux originates from these areas. Large changes in N2O emissions from these regions in the past would have directly impacted global greenhouse gas concentrations.

Distribution of membrane lipids of planktonic Crenarchaeota in the Arabian Sea.

Intact core tetraether membrane lipids of marine planktonic Crenarchaeota were quantified in water column-suspended particulate matter obtained from four depth intervals ( approximately 70, 500, 1,000 and 1,500 m) at seven stations in the northwestern Arabian Sea to investigate the distribution of the organisms at various depths. Maximum concentrations generally occurred at 500 m, near the top of the oxygen minimum zone, and the concentrations at this depth were, in most cases, slightly higher than those in surface waters. In contrast, lipids derived from eukaryotes (cholesterol) and from eukaryotes and bacteria (fatty acids) were at their highest concentrations in surface waters. This indicates that these crenarchaeotes are not restricted to the photic zone of the ocean, which is consistent with the results of recent molecular biological studies. Since the Arabian Sea has a strong oxygen minimum zone between 100 and 1,000 m, with minimum oxygen levels of <1 microM, the abundance of crenarchaeotal membrane lipids at 500 m suggests that planktonic Crenarchaeota are probably facultative anaerobes. The cell numbers we calculated from the concentrations of membrane lipids are similar to those reported for the Central Pacific Ocean, supporting the recent estimation of M. B. Karner, E. F. DeLong, and D. M. Karl ( Nature 409:507-510, 2001) that the world's oceans contain ca. 10(28) cells of planktonic Crenarchaeota.

Climatically related millennial-scale fluctuations in strength of California margin oxygen-minimum zone during the past 60 k.y.

A strong oxygen-minimum zone (OMZ) currently exists along the California margin because of a combination of high surface-water productivity and poor intermediate-water ventilation. However, the strength of this OMZ may have been sensitive to late Quaternary ocean-circulation and productivity changes along the margin. Although sediment-lamination strength has been used to trace ocean-oxygenation changes in the past, oxygen levels on the open margin are not sufficiently low for laminations to form. In these regions, benthic foraminifera are highly sensitive monitors of OMZ strength, and their fossil assemblages can be used to reconstruct past fluctuations. Benthic foraminiferal assemblages from Ocean Drilling Program Site 1017, off Point Conception, exhibit major and rapid faunal oscillations in response to late Quaternary millennial-scale climate change (Dansgaard-Oeschger cycles) on the open central California margin. These faunal oscillations can be correlated to and are apparently synchronous with those reported from Santa Barbara Basin. Together they represent major fluctuations in the strength of the OMZ which were intimately associated with global climate change--weakening, perhaps disappearing, during cool periods and strengthening during warm periods. These rapid, major OMZ strength fluctuations were apparently widespread on the Northeast Pacific margin and must have influenced the evolution of margin biota and altered biogeochemicalmore » cycles with potential feedbacks to global climate change.« less