Peru Margin Research Articles

Influence of marine denitrification on atmospheric N2O variability during the Holocene

Oceanic denitrification centers are thought to be important marine sources for atmospheric N2O. To consider Holocene variability in this source, we reconstruct the Holocene paleo‐denitrification history of the Peru margin, a major marine denitrification center, using high‐resolution sedimentary δ15N data. This record along with a contemporaneous one from the western Arabian Sea (Altabet et al., 2002) shows similarities with the recently available high‐resolution atmospheric N2O record (Flückiger et al., 2002). While the role of terrestrial processes in the observed N2O changes remains uncertain, these results suggest that variability in marine denitrification in major upwelling centers such as the Peru margin and the Arabian Sea contributed significantly to atmospheric N2O evolution during the Holocene.

Stratified Communities of Active Archaea in Deep Marine Subsurface Sediments

Archaeal 16S rRNA was extracted from samples of deep marine subsurface sediments from Peru Margin site 1227, Ocean Drilling Program leg 201. The amounts of archaeal 16S rRNA in each extract were quantified by serial dilution and reverse transcription (RT)-PCR. The results indicated a 1,000-fold variation in rRNA content with depth in the sediment, with the highest concentrations found near the sediment surface and in the sulfate-methane transition zone (SMTZ). The phylogenetic composition of the active archaeal population revealed by cloning and sequencing of RT-PCR products changed with depth. Several phylotypes affiliated with marine benthic group B (MBGB) dominated clone libraries from the upper part of the SMTZ and were detected only in this layer. Members of the miscellaneous crenarchaeotal group (MCG) dominated clone libraries from the other layers. These results demonstrate that archaeal communities change in activity and community composition over short distances in geochemically distinct zones of deep subseafloor sediments and that these changes are traceable in the rRNA pool. It was shown for the first time that members of both the MCG and MBGB Archaea are more active in the SMTZ than in layers above and below. This indicates that they benefit either directly or indirectly from the anaerobic oxidation of methane. They also appear to be ecophysiologically flexible, as they have been retrieved from a wide range of marine sediments of various geochemical properties.

Analysis of DGGE profiles to explore the relationship between prokaryotic community composition and biogeochemical processes in deep subseafloor sediments from the Peru Margin

The aim of this work was to relate depth profiles of prokaryotic community composition with geochemical processes in the deep subseafloor biosphere at two shallow-water sites on the Peru Margin in the Pacific Ocean (ODP Leg 201, sites 1228 and 1229). Principal component analysis of denaturing gradient gel electrophoresis banding patterns of deep-sediment Bacteria, Archaea, Euryarchaeota and the novel candidate division JS1, followed by multiple regression, showed strong relationships with prokaryotic activity and geochemistry (R(2)=55-100%). Further correlation analysis, at one site, between the principal components from the community composition profiles for Bacteria and 12 other variables quantitatively confirmed their relationship with activity and geochemistry, which had previously only been implied. Comparison with previously published cell counts enumerated by fluorescent in situ hybridization with rRNA-targeted probes confirmed that these denaturing gradient gel electrophoresis profiles described an active prokaryotic community.

Prokaryotic community composition and biogeochemical processes in deep subseafloor sediments from the Peru Margin

The community compositions of Bacteria and Archaea were investigated in deep, sub-seafloor sediments from the highly productive Peru Margin (ODP Leg 201, sites 1228 and 1229, c. 25 km apart) down to nearly 200 m below the seafloor using taxonomic (16S rRNA) and functional (mcrA and dsrA) gene markers. Bacterial and archaeal groups identified from clone libraries of 16S rRNA gene sequences at site 1229 agreed well with sequences amplified from bands excised from denaturing gradient gel electrophoresis (DGGE) depth profiles, with the exception of the Miscellaneous Crenarchaeotic Group (MCG). This suggested that the prokaryotic community at site 1228, obtained from DGGE profiling alone, was reliable. Sites were dominated by Bacteria in the Gammaproteobacteria, Chloroflexi (green non-sulphur bacteria) and Archaea in the MCG and South African Gold Mine Euryarchaeotic Group, although community composition changed with depth. The candidate division JS1 was present throughout both sites but was not dominant. The populations identified in the Peru Margin sediments consisted mainly of prokaryotes found in other deep subsurface sediments, and were more similar to communities from the Sea of Okhotsk (pelagic clays) than to those from the low organic carbon Nankai Trough sediments. Despite broad similarities in the prokaryotic community at the two sites, there were some differences, as well as differences in activity and geochemistry. Methanogens (mcrA) within the Methanosarcinales and Methanobacteriales were only found at site 1229 (4 depths analysed), whereas sulphate-reducing prokaryotes (dsrA) were only found at site 1228 (one depth), and these terminal-oxidizing prokaryotes may represent an active community component present at low abundance. This study clearly demonstrates that the deep subsurface sediments of the Peru Margin have a large diverse and metabolically active prokaryotic population.

Presence of two phylogenetically distinct groups in the deep-sea mussel Acharax (Mollusca: Bivalvia: Solemyidae)

The family Solemyidae represents ancestral protobranch bivalves with the shallow-water genus Solemya and the deep-sea genus Acharax. All known members of this family host symbiotic sulfur-oxidizing bacteria in their gill filaments. Analysis of 18S rRNA gene sequences of Acharax specimens from methane-seeps off Makran (Pakistan), Java (Indonesia), the Aleutian Trench and off the Oregon, Costa Rica, and Peru margins revealed that Solemya spp. and Acharax spp. are well-separated genetically. This supports the current systematic distinction based on morphological criteria. We found 2 clearly distinct clusters within the genus Acharax, with specimens from the Makran, Oregon and Peru (MOP) margins in one (MOP–Acharax) cluster, and those from Java, the Aleutian Trench and Costa Rica (JAC) in the other (JAC–Acharax) cluster. The separation of MOP– and JAC–Acharax clusters from each other and from Solemya (S. reidi and S. velum) is well-supported by phylogenetic calculations employing maximum likelihood and maximum parsimony. Compared to genetic distances among other protobranch groups, distances between the MOP– and JAC–Acharax clusters would justify the affiliation of these clusters to separate species. This implies that species differentiation in Acharax based on shell morphology is likely to underestimate true species diversity within this taxon. Furthermore, our results support the hypothesis that genetic separation of Solemya and Acharax is congruent with the phylogeny of their bacterial endosymbionts.

Microbial stratification in deeply buried marine sediment reflects changes in sulfate/methane profiles

ABSTRACTWe examined sediments collected at Ocean Drilling Program (ODP) Leg 201 Site 1229 on the Peru Margin for microbial populations throughout the sediment column. Heterotrophic cultivation from these sediments yielded numerous colonies from various depths, including 49 bacterial isolates. At ODP Site 1229, there are significant interfaces of sulfate and methane, across which microbial cell numbers increase substantially. At these sulfate/methane transition zones (SMTZs), however, we observed a decrease in the success rate for the cultivation of bacterial colonies. Utilizing both direct plating and enrichment in different media, we cultivated isolates from the upper SMTZ around 30 m below seafloor (mbsf); however, similar attempts yielded no colonies from within the lower zone at 85 mbsf. The phylogenetic relationships of the 16S rRNA gene sequences for the isolates were determined and most were related to other organisms and sequences previously found in the subsurface belonging to the γ‐Proteobacteria, cytophaga–flavobacterium–bacteroides, high G + C Gram‐positives, and Firmicutes groups. The most diverse group of isolates from Site 1229 was found between the SMTZs at 50 mbsf. ODP Leg 201 Site 1228 was examined for comparison and yielded an additional 18 isolates from 16 to 179 mbsf that were similar to those found at Site 1229. Direct plating at Site 1228 also showed decreased colony formation in the area of sulfate/methane transition. Our results suggest that heterotrophic bacterial populations are affected by SMTZs in deeply buried sediment.

Thermosediminibacter oceani gen. nov., sp. nov. and Thermosediminibacter litoriperuensis sp. nov., new anaerobic thermophilic bacteria isolated from Peru Margin

A new group of anaerobic thermophilic bacteria was isolated from enrichment cultures obtained from deep sea sediments of Peru Margin collected during Leg 201 of the Ocean Drilling Program. A total of ten isolates were obtained from cores of 1-2 m below seafloor (mbsf) incubated at 60 degrees C: three isolates came from the sediment 426 m below sea level with a surface temperature of 9 degrees C (Site 1227), one from 252 m below sea level with a temperature of 12 degrees C (Site 1228), and six isolates under sulfate-reducing condition from the lower slope of the Peru Trench (Site 1230). Strain JW/IW-1228P from the Site 1228 and strain JW/YJL-1230-7/2 from the Site 1230 were chosen as representatives of the two identified clades. Based on the 16S rDNA sequence analysis, these isolates represent a novel group with Thermovenabulum and Caldanaerobacter as their closest relatives. The temperature range for growth was 52-76 degrees C with an optimum at around 68 degrees C for JW/IW-1228P and 43-76 degrees C with an optimum at around 64 degrees C for JW/YJL-1230-7/2. The pH(25C) range for growth was from 6.3 to 9.3 with an optimum at 7.5 for JW/IW-1228P and from 5 to 9.5 with an optimum at 7.9-8.4 for JW/YJL-1230-7/2. The salinity range for growth was from 0% to 6% (w/v) for JW/IW-1228P and from 0% to 4.5% (w/v) for JW/YJL-1230-7/2. The G+C [corrected] mol% of the genomic DNA was 46.3 +/- 0.7% (n = 4) for Thermosediminibacter oceani [corrected] JW/IW-1228PT [corrected] and 45.2 +/- 0.7 (n = 6) for Thermosediminibacter litoriperuensis [corrected] JW/YJL-1230-7/2T [corrected] DNA-DNA hybridization yielded 52% similarity between the two strains. According to 16S rRNA gene sequence analysis, the isolates are located within the family, Thermoanaerobacteriaceae. Based on their morphological and physiological properties and phylogenetic analysis, it is proposed that strain JW/IW-1228P(T) is placed into a novel taxa, Thermosediminibacter oceani, gen. nov., sp. nov. (DSM 16646(T)=ATCC BAA-1034(T)), and JW/YJL-1230-7/2(T) into Thermosediminibacter litoriperuensis sp. nov. (DSM 16647(T) =ATCC BAA-1035(T)).

Coexistence of Bacterial Sulfide Oxidizers, Sulfate Reducers, and Spirochetes in a Gutless Worm (Oligochaeta) from the Peru Margin

Olavius crassitunicatus is a small symbiont-bearing worm that occurs at high abundance in oxygen-deficient sediments in the East Pacific Ocean. Using comparative 16S rRNA sequence analysis and fluorescence in situ hybridization, we examined the diversity and phylogeny of bacterial symbionts in two geographically distant O. crassitunicatus populations (separated by 385 km) on the Peru margin (water depth, approximately 300 m). Five distinct bacterial phylotypes co-occurred in all specimens from both sites: two members of the gamma-Proteobacteria (Gamma 1 and 2 symbionts), two members of the delta-Proteobacteria (Delta 1 and 2 symbionts), and one spirochete. A sixth phylotype belonging to the delta-Proteobacteria (Delta 3 symbiont) was found in only one of the two host populations. Three of the O. crassitunicatus bacterial phylotypes are closely related to symbionts of other gutless oligochaete species; the Gamma 1 phylotype is closely related to sulfide-oxidizing symbionts of Olavius algarvensis, Olavius loisae, and Inanidrilus leukodermatus, the Delta 1 phylotype is closely related to sulfate-reducing symbionts of O. algarvensis, and the spirochete is closely related to spirochetal symbionts of O. loisae. In contrast, the Gamma 2 phylotype and the Delta 2 and 3 phylotypes belong to novel lineages that are not related to other bacterial symbionts. Such a phylogenetically diverse yet highly specific and stable association in which multiple bacterial phylotypes coexist within a single host has not been described previously for marine invertebrates.

Direct in situ detection of cells in deep‐sea sediment cores from the Peru Margin (ODP Leg 201, Site 1229)

ABSTRACTMicrobiological investigations of deep‐sea sediments recovered from the Peru Margin during the ODP Leg 201 (Hole 1229A, 1–110 mbsf) demonstrated that microoganisms were a consistent component throughout the profile. Optimization of the dilution factor and DAPI‐staining procedures for direct cell counts allowed the determination of the abundance of the entire microbial community, which was about 108 cells per g dry sediment. Microbial diversity in discrete samples taken from the 110‐m profile was analysed using horseradish‐peroxydase‐rRNA‐probes. In general, the majority of the detected cells belonged to the Eubacteria kingdom with a dominance of sulphate‐reducing bacteria. The composition of the suflate‐reducing community varied with depth. Desulfobacteriaceae were dominant in the uppermost sulphate‐reducing zone and Desulfovibrionaceae at deeper depths in the upward diffusing sulphate‐rich brines. Both sulphate‐reducing groups were also detected in the methanogenic zone. Similarly, Archaea were detected throughout the profile, not only in the methanogenic zone but also in the upper and lower sulphate‐reducing zones.

Initial test of the silicic acid leakage hypothesis using sedimentary biomarkers

Several mechanisms have been proposed for large CO2 changes at glacial Terminations, including shifting the CaCO3:Corg rain ratio by changing surface water nutrient supply, altering the balance between diatom and coccolithophore production. Diatom Si:N is highest in Fe‐stressed high‐latitude waters. Southern Ocean Fe enrichment studies suggest diatom Si demands reduced under Fe‐replete (glacial) conditions, allowing increased silicic acid to leak northward in subducted intermediate water and upwell at lower latitudes. We test this ‘Silicic Acid Leakage’ hypothesis using relative abundances of phytoplankton‐specific biomarkers in Peru margin sediments spanning 0–20 Ka. Results indicate increased coccolithophorid:diatom production from ∼0.5 to 3 between 18.0–15.5 Ka. Temporal correlation with the initial pCO2 rise and early deglacial shift in mode water ventilation implicates a coincidental, possibly causative reorganization of Sub‐Antarctic Mode Water formation and reduced Fe abundance. However, coccolithophorid production subsequently declined, suggesting rain ratio changes were only partly responsible for the CO2 deglacial transition.

Bioturbation by symbiont-bearing annelids in near-anoxic sediments: Implications for biofacies models and paleo-oxygen assessments

Anoxic or nearly anoxic conditions (<4 μM O 2) have long been associated with the absence of bioturbation and animal traces. This premise has guided interpretation of paleoceanographic conditions from rocks and sediments. We recently observed a high-density, living assemblage of highly mobile, symbiont-bearing, burrowing, phallodrilinid oligochaetes within a nearly anoxic basin (<1 μM O 2 [0.02–0.03 ml l −1]) on the Peru margin (305 m). These observations were made during the most intense part of the 1997–98 El Niño when there may have been slight oxygenation of an otherwise anoxic basin, but oligochaete presence prior to this event is likely. The occurrence of symbiont-bearing gutless oligochaetes mainly within the upper 5 cm of the sediment column coincided with a bioturbated zone overlying distinctly laminated sediments. Our observations redefine the lower oxygen limit of macrofaunal bioturbation to ≪2 μM, and indicate a need to modify currently accepted ideas about the relationship between bioturbation and paleo-oxygen concentration. These results also address an ongoing debate about the lifestyles of bioturbating organisms in oxygen-poor settings.

Oceanic conditions in the eastern equatorial Pacific during the onset of ENSO in the Holocene

Records from South America show that modern ENSO (El Nino-Southern Oscillation) did not exist 7000 cal yr B.P. and has developed progressively since then. There has been little information available on oceanic conditions in the eastern equatorial Pacific (EEP) to constrain explanations for ENSO onset. We report quantitative observations on thermocline and mixed-layer conditions in the EEP during ENSO start up. We found important changes in both the thermocline and the mixed layer, indicating increased upwelling of cooler waters since 7000 cal yr B.P. This resulted from change in the source and/or properties of waters supplying the Equatorial Undercurrent, which feeds upwelling along the equator and the Peru margin. Modeling shows that ENSO is sensitive to subsurface conditions in the eastern equatorial Pacific and that the changes in the thermocline we observed were driven by extratropical processes, giving these a role in conditioning the development of ENSO. This is in contrast to models that call for control of equatorial Pacific oceanography by tropical processes only. These infer stronger upwelling and cooler surface waters for the EEP during the mid-Holocene, which is not supported by our results.

Patterns of export production in the eastern equatorial Pacific over the past 130,000 years

Productivity at six core locations in the eastern equatorial Pacific (EEP) was reconstructed with a benthic foraminiferal transfer function. The core records show strong regionality, especially where affected by Peru margin upwelling of deeper Equatorial Undercurrent Water (EUC) (originally coming from the subantarctic). This “Peru margin” record differs from that seen along the equator where divergence leads to shallow upwelling, and it is generally inverse to that seen in cores outside the areas of equatorial upwelling. Principal components analysis shows that the main productivity pattern correlates well to the global oxygen isotope record and has lowest values during isotope stages 2 and 4. In addition to this, equatorial cores show a higher frequency pattern of variation which becomes much more pronounced during MIS 3 and 2. The reconstructions based on benthic foraminifera were tested against those from nonaccumulation rate based inorganic chemical proxies of export production. These were found to correlate well in the region influenced by Peru upwelling, and also to share common features for sites along the equator. All the nonaccumulation rate based paleotracers are consistent with one another and differ from accumulation rate derived proxies. The differences between the two classes of paleotracers may result from uncertainties in calculating actual biogenic fluxes since 230Th‐normalized results conform more to those we obtained. Analysis of planktonic carbon isotope values for the EEP, and their comparison to the record of the Pacific subantarctic, indicates that the subantarctic contribution to the EUC was reduced during MIS 3 and 2.

Structural CO2 in the Apatite of the Late Cretaceous Phosphorite Resources in Egypt: Effect on the Crystal Chemistry and Geologic Implications

Abstract: Phosphorite deposits in Egypt, known as the Duwi Formation, are a part of the Middle East to North Africa phospho‐genic province of late Cretaceous to Paleogene age. Based on the petrographical observation, the phosphatic grains in the phosphorites are classified into phosphatic mudclasts and phosphatic bioclasts. Both of them are composed of francolite. The structural CO2 contents in the francolite range from 3.3 to 7.2 % with an average of 5.3 %. Results indicated that the substitution with CO32‐ of PO43‐ in the francolite decreases the unit cell volume and a‐cell dimension, and increases the c/a ratio. Effect is more obvious in the a‐cell dimension; therefore, it is more significant in distinction between the different apatite species.Lack of covariance between structural CO2 contents in the francolite and the carbonate minerals contents may render the supposition that the phosphorites formed as a result of replacement of preexisting calcareous sediments is doubtful. Similarity in CO2 content in both weathered and fresh samples indicates that the structural CO2 content in the phosphorites is not affected by weathering, and reflects the conditions and CO2 concentration of the depositional environment. Similarity in mineralogy and CO2 contents in the different phosphatic grains and higher CO2 content in the Egyptian phosphorites compared with the authigenic phosphates of Peru margin, which formed by the same mechanism as the Duwi phosphorites, suggest that the phosphatic grains in the Duwi Formation were francolitized during diagenesis by introducing CO2 from the surrounding pore water and diagenesis took place at an elevated temperature. Scattered values of structural CO2 contents suggest the reworking origin of the phosphatic grains in the late Cretaceous phosphorites in Egypt.

Remote vs. local control of changes in eastern equatorial Pacific bioproductivity from the Last Glacial Maximum to the Present

Data from 14 cores, and a transfer function based on benthic foraminifera, were used to map productivity gradients in the eastern equatorial Pacific (EEP) at the Last Glacial Maximum (LGM). These were compared to gradients in the modern ocean. The results support previous work indicating that, during the LGM, productivity was lower across the region in the South Equatorial Current (SEC) under the influence of Peru margin upwelling. Overall, productivity gradients are diminished during the LGM due to three changes: reduced productivity along the Peru margin and the SEC near the equator, increased productivity further south of the equator, and increased productivity in the Panama Basin area. These changes smooth gradients seen in the region today. This reduction of gradients in EEP productivity parallels observations for planktonic foraminiferal carbon isotopes and inferred nutrient concentrations in the thermocline. Reductions in productivity and thermocline nutrients in the SEC downstream of the Peru margin have been previously interpreted as the result of changes in the chemistry and/or upwelling of subantarctic Equatorial Undercurrent Current (EUC) water. Increases in productivity away from the equator may be associated to nitrogen enrichment of the global surface ocean during the LGM. Productivity and planktonic foraminiferal isotope records for the Panama Basin indicate a local process driving highly seasonal productivity for the glacial, perhaps related to changing upper water column stratification. The degree to which overall carbon export to the deep sea changed in the EEP during the LGM depends on the extent of the productivity increase away from the equator. If this increase occurred broadly in the lower latitude subtropical gyres then a marked LGM export increase would be possible despite productivity reductions along the equator.

Tectonic erosion and consequent collapse of the Pacific margin of Costa Rica: Combined implications from ODP Leg 170, seismic offshore data, and regional geology of the Nicoya Peninsula

The convergent margin off the Pacific coast of the Nicoya Peninsula of Costa Rica exhibits evidence for subduction erosion caused by the underthrusting Cocos plate. Critical evidence for efficacy of this process was recovered at the Ocean Drilling Program (ODP) drilling Site 1042 (Leg 170), positioned ∼7 km landward of the Middle America trench axis off the Nicoya Peninsula. The primary drilling objective at this site was to identify the age and origin of a regionally extensive and prominent seismic discontinuity, the so‐called base‐of‐slope sediment (BOSS) horizon or surface. The BOSS horizon, which can be traced landward from near the trench to the Nicoya coastal area and parallel to it for hundreds of kilometers, separates a low‐velocity (∼ 2.0–2.5 km s−1) sequence of slope sediment, from an underlying sequence of much higher‐velocity (&gt;4–4.5 km s−1) rock. Site 1042 reached the acoustically defined BOSS horizon at a below sea level depth of ∼ 3900 m and yielded a carbonate‐cemented calcarenitic breccia of early‐middle Miocene age. Sedimentological, geochemical, paleontological, and cement paragenesis data document that the breccia accumulated in a shallow water depositional environment. On the basis of coastal exposures, the BOSS horizon, as a margin‐wide geologic interface, can be temporally and lithostratigraphically correlated to a regional angular unconformity. This unconformity, known as the Mal Pais unconformity, separates Neogene and younger shelf‐to‐littoral beds from the underlying mafic units of the Mesozoic Nicoya Complex and Cretaceous and early Tertiary sedimentary sequences. At Site 1042 it is inferred that tectonism caused the vertical subsidence of the early Neogene breccia from a shallow to a deep water setting. The Mal Pais unconformity of the BOSS horizon thus connects the rock fabric of the outermost part of margin to that of coastal Nicoya and implies that in the early Neogene the Nicoya shelf extended seaward to near the present trench axis. This circumstance requires that the early Neogene trench axis was at least 50 km seaward of where it is now located. The long‐term effects of subduction erosion, similar to those described for the scientifically drilled Japan, Tonga, and Peru margins, best account for offshore and onshore evidence for a post‐Paleogene history of crustal thinning and landward trench migration of Costa Rica's Pacific margin. During the past 16–17 Myr the calculated mass removal and landward migration rates are 34–36 km³ Myr−1 km−1 of margin, and 3 km Myr−1, respectively. These values are similar to those found for other Pacific margins dominated by nonaccretionary subduction zone processes.

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

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

The link between bottom-simulating reflections and methane flux into the gas hydrate stability zone – new evidence from Lima Basin, Peru Margin

Bottom-simulating reflections (BSRs) are probably the most commonly used indicators for gas hydrates in marine sediments. It is now widely accepted that BSRs are primarily caused by free gas beneath gas-hydrate-bearing sediments. However, our insight into BSR formation to date is mostly limited to theoretical studies. Two endmember processes have been suggested to supply free gas for BSR formation: (i) dissociation of gas hydrates and (ii) migration of methane from below. During a recent campaign of the German Research Vessel Sonne off the shore of Peru, we detected BSRs at locations undergoing both tectonic subsidence and non-sedimentation or seafloor erosion. Tectonic subsidence (and additionally perhaps seafloor erosion) causes the base of gas hydrate stability to migrate downward with respect to gas-hydrate-bearing sediments. This process rules out dissociation of gas hydrates as a source of free gas for BSRs at these locations. Instead, free gas at BSRs is predicted to be absorbed into the gas hydrate stability zone. BSRs appear to be confined to locations where the subsurface structure suggests focusing of fluid flow. We investigated the seafloor at one of these locations with a TV sled and observed fields of rounded boulders and slab-like rocks, which we interpreted as authigenic carbonates. Authigenic carbonates are precipitations typically found at cold vents with methane expulsion. We retrieved a small carbonate-cemented sediment sample from the seafloor above a BSR about 20 km away. This supported our interpretation that the observed slabs and boulders were carbonates. All these observations suggest that BSRs in Lima Basin are maintained predominantly by gas that is supplied from below, demonstrating that this endmember process for BSR formation exists in nature. Results from Ocean Drilling Program Leg 112 showed that methane for gas hydrate formation on the Peru lower slope and the methane in hydrocarbon gases on the upper slope is mostly of biogenic origin. The δ13C composition of the recovered carbonate cement was consistent with biologic methane production below the seafloor (although possibly above the BSR). We speculate that the gas for BSR formation in Lima Basin also is mainly biogenic methane. This would suggest the biologic productivity beneath the gas hydrate zone in Lima Basin to be relatively high in order to supply enough methane to maintain BSRs.

Sulfur diagenesis and partitioning in Holocene Peru shelf and upper slope sediments

Sediments of the Peru margin were sampled in two transects across the shelf and upper slope (75 to 1000 m) in October of 1992. This region is characterized by high surface-water productivity and consequent high organic carbon fluxes, low terrigenous inputs, as well as an intense oxygen-minimum zone (OMZ) impinging on the slope at depths between 75 and 550 m. Sediments analyzed in this study were collected from one ship-deployed box core and five submersible-deployed push cores. Organic carbon (TOC) concentrations in surface sediments range from 1 to 18% on a dry weight basis. Formation of pyrite begins at the sediment/water interface (SWI). Maximum concentrations of pyrite sulfur in the cores from this study range from 0.5 to 1.8 wt.%. Concentrations of pyrite are influenced by availability of reactive iron and an oxidant capable of converting AVS to pyrite. Highly reactive iron is stripped from terrigenous detritus in the core of the OMZ. Some of this iron may contribute to glauconite formation in sediments near the base of the OMZ, where it is concentrated into a glauconitic green sand as a result of reworking and winnowing. In a shallow-water site (109 m) at the upper edge of the OMZ, high concentrations of pyrite, total iron and iron/aluminum ratios indicate that easily reduced iron species are available for sulfidation at the SWI. Pyrite formation at this site is nearly complete in the upper 4 cm of the sediment. In contrast, pyrite concentrations are slightly lower at the surface of two of the deeper-water sites (at 309 m, in the center of the OMZ, and to a lesser extent at 671 m, at the bottom edge of the OMZ), and appear to be influenced, in part, by availability of reactive iron. Concentrations of acid-volatile sulfides (AVS) are low in all sediments (≤ 0.04 wt.%), but are slightly elevated in sediments from the center of the OMZ. The ratio of organic sulfur (S-org)to organic carbon (TOC)at the sediment surface of the cores examined here is remarkably constant (0.012 ± 0.002) and within the range observed in marine biota (0.01 to 0.02), suggesting that organic sulfur at the SWI is dominantly biogenic. Although S-org/TOC ratios increase downcore, there is no clear correlation between concentrations of organic sulfur and organic carbon below the SWI. There is also no clear relation between concentrations of organic sulfur and total iron. Organic sulfur concentrations vary from site to site (0.06 to 0.33 wt.%), but are relatively constant downcore at any given site. Maximum concentrations are in cores from the center of the OMZ.

Foraminifera in the Arabian Sea oxygen minimum zone and other oxygen-deficient settings: taxonomic composition, diversity, and relation to metazoan faunas

Previous work has shown that some foraminiferal species thrive in organically enriched, oxygen-depleted environments. Here, we compare `live’ (stained) faunas in multicorer samples (0–1 cm layer) obtained at two sites on the Oman margin, one located at 412 m within the oxygen minimum zone (OMZ) (O 2=0.13 ml l −1), the other located at 3350 m, well below the main OMZ (O 2∼3.00 ml l −1). While earlier studies have focused on the hard-shelled (predominantly calcareous) foraminifera, we consider complete stained assemblages, including poorly known, soft-shelled, monothalamous forms. Densities at the 412-m site were much higher (16,107 individuals.10 cm −2 in the >63-μm fraction) than at the 3350-m site (625 indiv.10 cm −2). Species richness (E(S 100)), diversity (H′, Fishers Alpha index) and evenness (J′) were much lower, and dominance (R1D) was higher, at 412 m compared with 3350 m. At 412 m, small calcareous foraminifera predominated and soft-shelled allogromiids and sacamminids were a minor faunal element. At 3350 m, calcareous individuals were much less common and allogromiids and saccamminids formed a substantial component of the fauna. There were also strong contrasts between the foraminiferal macrofauna (>300-μm fraction) at these two sites; relatively small species of Bathysiphon, Globobulimina and Lagenammina dominated at 412 m, very large, tubular, agglutinated species of Bathysiphon, Hyperammina, Rhabdammina and Saccorhiza were important at 3350 m. Our observations suggest that, because they contain fewer soft-shelled and agglutinated foraminifera, a smaller proportion of bathyal, low-oxygen faunas is lost during fossilization compared to faunas from well-oxygenated environments. Trends among foraminifera (>63 μm fraction) in the Santa Barbara Basin (590 and 610 m depth; O 2=0.05 and 0.15 ml l −1 respectively), and macrofaunal foraminifera (>300 μm) on the Peru margin (300–1250 m depth; O 2=0.02–1.60 ml l −1), matched those observed on the Oman margin. In particular, soft-shelled monothalamous taxa were rare and large agglutinated taxa were absent in the most oxygen-depleted (<0.20 ml l −1) stations. Foraminifera often outnumber metazoans (both meiofaunal and macrofaunal) in bathyal oxygen-depleted settings. However, although phylogenetically distant, foraminifera and metazoans exhibit similar population responses to oxygen depletion; species diversity decreases, dominance increases, and the relative abundance of the major taxa changes. The foraminiferal macrofauna (>300 μm) were 5 times more abundant than the metazoan macrofauna at 412 m on the Oman margin but 16 times more abundant at the 3350 m site. Among the meiofauna (63–300 μm), the trend was reversed; foraminifera were 17 times more abundant than metazoan taxa at 412 m but only 1.4 times more abundant at 3350 m. An abundance of food combined with oxygen levels which are not depressed sufficiently to eliminate the more tolerant taxa, probably explains why foraminifera and macrofaunal metazoans flourished at the 412-m site, perhaps to the detriment of the metazoan meiofauna.