Global latitudinal species diversity gradient in deep-sea benthic foraminifera

Global scale patterns of species diversity for modern deep-sea benthic foraminifera, an important component of the bathyal and abyssal meiofauna, are examined using comparable data from five studies in the Atlantic, ranging over 138° of latitude from the Norwegian Sea to the Weddell Sea. We show that a pattern of decreasing diversity with increasing latitude characterises both the North and South Atlantic. This pattern is confirmed for the northern hemisphere by independent data from the west-central North Atlantic and the Arctic basin. Species diversity in the North Atlantic northwards from the equator is variable until a sharp fall in the Norwegian Sea (ca. 65°N). In the South Atlantic species diversity drops from a maximum in latitudes less than 30°S and then decreases slightly from 40 to 70°S. For any given latitude, North Atlantic diversity is generally lower than in the South Atlantic. Both ecological and historical factors related to food supply are invoked to explain the formation and maintenance of the latitudinal gradient of deep-sea benthic foraminiferal species diversity. The gradient formed some 36 million years ago when global climatic cooling led to seasonally fluctuating food supply in higher latitudes.     

Regional and seasonal variations of recent benthic deep-sea foraminifera in the Arabian Sea

Assemblages of living deep-sea benthic foraminifera, their densities, vertical distribution pattern, and diversity, were investigated in the intermonsoon period after the northeast monsoon in the Arabian Sea in spring 1997. Foraminiferal numbers show a distinct gradient from north to south, with a maximum of 623 foraminifera in 50 cm³ at the northern site. High percentages of small foraminifera were found in the western and northern part of the Arabian Sea. Most stations show a typical vertical distribution with a maximum in the first centimeter and decreasing numbers with increasing sediment depths. But at the central station, high densities can be found even in deeper sediment layers. Diversity is very high at the northern and western sites, but reduced at the central and southern stations. Data and faunal assemblages were compared with studies carried out in 1995. A principal component analysis of intermonsoon assemblages shows that the living benthic foraminifera can be characterized by five principal component communities. Dominant communities influencing each site differ strongly between the two years. In spring 1997, stations in the north, west and central Arabian Sea were dominated by opportunistic species, indicating the influence of fresh sedimentation pulses or enhanced organic carbon fluxes after the northeast monsoon.     

Influence of seasonal primary productivity on δ13C of North Atlantic deep-sea benthic foraminifera

The stable isotopic composition of benthic foraminifera has been widely used to reconstruct deep-ocean circulation, but questions have been raised about the influence of organic carbon flux on the carbon isotopic composition of deep-sea taxa. We show that annual and seasonality of primary productivity in the North Atlantic do not affect δ¹³C of Planulina wuellerstorfi, but that the intermittency or seasonality of primary production has a significant effect (0.9‰ change over 60° latitude) on δ¹³C of Epistominella exigua, reflecting the influence of pelagic-benthic coupling and microhabitat preferences on test geochemistry. These results support the use of δ¹³C of P. wuellerstorfi in paleocirculation studies and suggest that the δ¹³C of E. exigua can be used to reconstruct seasonality of productivity.     

Temporal variations in the deep-water colonization rates of small benthic foraminifera: the results of an experiment on Cross Seamount

Modern benthic foraminifera are important remineralizers of organic matter and a link between surface-ocean production and life in the deep sea. Assemblages of benthic foraminifera are preserved in the fossil record, providing clues to paleoceanographic conditions. These clues can be fully interpreted only when our knowledge of the biology and ecology of modern species is more complete. To study factors influencing foraminiferal colonization rates, artificial substrates were placed on Cross Seamount (18°40′N, 158°17′W) for 1 to 42 months between 1989 and 1994. The colonization rates of benthic foraminifera onto different substrates (five mineral types were used) at four water depths (800, 975, 1285 and 2000 m) were determined. Both calcareous and agglutinated foraminifera inhabited these artificial substrates at different rates. Many of the agglutinated forms colonized at a uniform rate through time. The colonization rates of other foraminifera, primarily calcareous forms, were not constant through time, nor could the variability be attributed to the controlled variables (water depth and substrate composition). Instead, these temporal variations in colonization rate corresponded with shifts in surface ocean conditions and export fluxes in the central Pacific. Other ecological observations are also presented, including size- and spatial-distributions.     

Deep-sea benthic foraminifera,carbonate dissolution and species diversity in Hardangerfjord,Norway: An initial assessment

This is the first record of live (stained) deep-sea benthic foraminifera in the 850 m deep silled Hardangerfjord, the second deepest fjord in Western Norway. Estimates of organic carbon flux (∼2.5 g Cm⁻² y⁻¹) show that the fjord-values are comparable to similar depths on the continental slope. Accordingly, although these first samples only provide relative abundance data, the low proportion of live to dead individuals in the top cm of the sediment suggests a low foraminifera biomass. Another similarity with the deep sea is that the abiotic environment of the deep basins is stable even though the deepest basins are isolated from the open deep sea by the continental shelf and sills in the outer parts of the fjord suggesting that the deep-sea species are introduced as propagules during deep-water renewals. There is evidence of an increase in dissolution of fragile calcareous tests (e.g., Nonionella iridea) especially in the innermost part of Hardangerfjord since the 1960s and this has led to a relative increase in dead agglutinated assemblages. The presence of larger forms with tests >1 mm provides substrata for the attachment of smaller forms and therefore an increase in species diversity. Indeed, the diversity is comparable both to that of the open deep sea and that of reported macrofauna from the same sites, reflecting similar ecological status. Holtedahl (1965) suggested that there may be some down-slope transport of sediment into the deep basins with the deposition of turbidites. Despite some evidence of transport, no major recent disturbance due to turbidite deposition seems to have occurred and hence Hardangerfjord presents a unique environment with elements of deep-sea faunas in a land-locked setting.     

底栖有孔虫对厦门筼筜湖污染的监测作用

1992年11月至1993年8月对厦门湖及厦门西港部分海域底栖有孔虫及相关的环境化学要素进行了4个季度月的调查，发现湖的底栖有孔虫可分为三个群落，即正常群落、半污染群落和污染群落，分别代表了湖内不同污染程度环境，同时发现了底栖有孔虫的耐得种类和对污染敏感种类，它们具有指示环境的意义。     

Deep penetrating benthic oxygen profiles measured in situ by oxygen optodes

For the first time in situ, deep penetrating O₂ profiles were measured in abyssal sediments in the western South Atlantic. Construction of deep penetrating O₂ optodes and adaptation to a benthic profiling lander are described. The opto-chemical oxygen sensors allow measurements to a depth of 55 cm in marine sediments. A vertical resolution of 0.5 cm was used to determine the O₂ dynamics in those oligotrophic deep sea sediments; the oxygen concentration across the sediment water interface was measured with a resolution of 100 μm. Oxygen penetration depth (OPD), diffusive oxygen uptake (DOU) and oxygen consumption rates were determined at four stations north of the Amazon fan and one at the Mid-Atlantic Ridge. Diffusive oxygen uptake rates ranged from 0.1 to 0.9 mmol m⁻² d⁻¹; the oxygen penetration depth ranged from 8 to 26 cm. Carbon consumption rates calculated from the diffusive oxygen uptake rates were in the range of 0.3–3.0 g C m⁻² a⁻¹. Comparison between in situ and laboratory DOU and OPD measurements confirmed previous findings that core recovery and warming have strong effects on the oxygen dynamics in deep sea sediments. Laboratory measurements yielded a decrease of 50–75% in OPD and consequently an increase in DOU by 1.5 and 18-times. Deep penetrating oxygen optodes provide a new tool to accurately determine oxygen dynamics (and thereby calculate carbon mineralization rates) in oligotrophic sediments. However, oxygen optodes as used in this study do not resolve the diffusive boundary layer (DBL). The data show that deep penetrating O₂ optodes in combination with high-resolution O₂ microelectrodes give a complete picture of the oxygen dynamics, including the DBL, in deep sea sediments.     

Feeding of benthic foraminifera on diatoms and sewage-derived organic matter: an experimental application of lipid biomarker techniques

Foraminiferal ecology at sewage outfalls has been investigated in numerous field studies over the last 30 years. Foraminifera have been frequently used as biomonitors of sewage pollution since they are both abundant and ubiquitous. Sewage outfalls have been demonstrated to have both positive and negative effects on adjacent foraminiferal populations, but it has never been shown conclusively why sewage affects foraminifera in these ways. Such information on the impact mechanisms of sewage pollution is essential if foraminifera are to be used as sewage pollution biomonitors, and also to understand the ecology of these important protists. One possible cause of a positive effect is the direct consumption of sewage-derived particulate organic matter (POM) by the foraminifera themselves. However this hypothesis has never been tested experimentally. Here, lipid (fatty acid and sterol) biomarker techniques were applied to study the ingestion of two potential food items by the foraminiferan Haynesina germanica in the laboratory. An experiment was conducted to confirm that the laboratory conditions were conducive to the survival and feeding of the foraminifera. In this experiment, foraminifera were provided with the pennate diatom Phaeodactylum tricornutum, which was considered to be a suitable food source. After 2 weeks, a four-fold increase in the levels of the diatom fatty acid biomarker, 20:5(n-3), in the foraminifera suggested that they had fed actively on the diatoms and survived under the experimental conditions. These experimental conditions were used in the main experiment, where foraminifera were fed the POM from sewage. Lipid biomarker analysis indicated that H. germanica did not consume secondary treated sewage-derived POM. Neither fatty acid profiles in the sewage nor coprostanol, the diagnostic human faecal sterol, were detected in foraminifera after exposure to the potential sewage food source. However, foraminifera may have consumed bacteria associated with the sewage in the experiment. The findings are discussed in terms of current EU legislation on sewage treatment that has affected the composition of sewage discharges, and therefore possibly reduced the nutritive value of sewage to the marine benthos.     

Ecophenotypic responses of benthic foraminifera to oxygen availability along an oxygen gradient in the California Borderland

Spatial variation in environmental conditions can elicit predictable size and morphological responses in marine organisms through influences on physiology. Thus, spatial and temporal variation in marine organism size and shape are often used to infer paleoenvironmental conditions, such as dissolved oxygen concentrations. Benthic foraminifera commonly serve as a tool for reconstructing past ocean oxygen levels. For example, benthic foraminiferal species assemblages, within‐ and among‐species patterns of test morphology, and geochemical analyses of carbonate tests are often employed to reconstruct past marine conditions. In this study, we measured the sizes and shapes of modern foraminifera representing four species that inhabit a steep oxygen gradient in the Santa Monica Basin on the Southern California Borderland with the aim of quantifying the influence of oxygen availability on foraminiferal morphology, both within and among species. Most foraminifera rely on aerobic respiration, but the four benthic foraminifera from the Santa Monica Basin do not show the predicted size and morphological responses to variations in dissolved oxygen concentrations based on first principles of cell physiology: Bolivina spissa shows no volume or volume‐to‐surface area response, Uvigerina peregrina increases in both test volume and volume‐to‐surface area ratio with decreasing dissolved oxygen concentrations, and both Bolivina argentea and Loxostomum pseudobeyrichi decrease in test volume, but only L. pseudobeyrichi shows a decrease in test volume‐to‐surface area ratio with decreasing oxygen concentrations as expected from physiological predictions. These findings imply that the morphological responses of individual foraminiferal species are not necessarily representative of the responses of other foraminiferal species within the community. Our findings further suggest that these species use physiological strategies such as depressed metabolic rates and alternative energy metabolisms to persist in low oxygen environments and, therefore, cannot be used in any simple way as paleo‐oxygen indicators. Should Proterozoic (1,000–542 million years ago) protists have possessed metabolic strategies similar to foraminifera, the sizes and shapes of protists in the fossil record may not usefully constrain ambient oxygen conditions during the appearance and initial taxonomic radiation of heterotrophic eukaryotes.     

Distribution of benthic foraminifera in Upper Quaternary sediments of the Deryugin Basin (Sea of Okhotsk)

Benthic foraminifera are investigated in sediment core LV28-34-2 (53°51.971′N, 146°47.499′E, sea depth 1431 m, core length 965 cm). The distribution of foraminifera is studied in coarse-grained (>0.125 mm) sediment fractions of 191 samples taken with a step of 5 cm. The core covers the interval from oxygen isotope stage (OIS) 6 up to the Holocene. The foraminiferal assemblages of the penultimate (OIS 6) and last (OIS 5d-2) glaciations are characterized by low abundances and prevalence of Uvigerina auberiana. The specific structure of the OIS 6 assemblages differs from the last glaciation ones by the mass presence of Cassidulina teretis, which characterizes low temperatures and a high influx of organic matter to the sea floor. The major factor responsible for the weak development of benthic foraminifers during the glacial time is the deficiency of food resources. The foraminiferal assemblage of the interglacial optimum (OIS 5e) is composed of both calcareous and agglutinated species (Martinottiella communis).The inflow of Pacific waters was probably more intensive, the bioproductivity was higher, and the critical carbonate compensation depth was shallower at that time than during the glaciations. During the deglaciation, the quantity and structure of the foraminiferal assemblages reflected two pulses of warming (terminations 1B and 1A) with an intervenient cooling event (Younger Dryassic). The assemblages of warm periods were characterized by exclusively high foraminiferal abundances, which sharply decreased during the Younger Dryassic cooling. The specific structure of the foraminiferal assemblage during deglaciation was relatively uniform, being composed of only calcareous taxa typical of highly productive areas of the ocean. The paleoenvironmental conditions were similar to the conditions of the interglacial optimum (OIS 5e), although the preservation of calcareous tests is better. In the Holocene sediments, the concentration of benthic foraminifera is substantially lower than during the deglaciation, which is explainable by their dilution in the sediments by diatoms; the composition of the foraminiferal assemblages is similar to that of their glacial counterparts.     

南海北部海域表层沉积物底栖有孔虫分布特征及影响因素

有孔虫在海洋环境指示方面具有重要意义,对现代环境中有孔虫的分布及影响因素进行研究有助于利用有孔虫对地质历史时期的沉积环境进行还原。我国南海海域辽阔,南海有孔虫的相关研究很早便有报道。对我国南海北部海域表层沉积物中的底栖有孔虫分布特征进行了描述,研究区以玻璃质壳底栖有孔虫占主导,胶结壳其次,瓷质壳最少。根据底栖有孔虫的分布特征划分了4个底栖有孔虫组合,组合1为Heterolepa subpraecincta - Hanzawaia mantaensis - Rotalinoides gaimardii;组合2为Bulimina marginate - Lagena substriata;组合3为Elphidium advenum - Pararotalia nipponica;组合4 为Recurvoides contortus - Ammodisus cretaceous - Bulimina aculeata。对影响该区域底栖有孔虫分布的环境因素进行了探究,结果表明,温度、盐度、底质类型等对底栖有孔虫的分布均存在一定程度的影响,海流对研究区底栖有孔虫时空分布的影响值得进一步探究。     

An in situ iron-enrichment experiment in the western subarctic Pacific (SEEDS): Introduction and summary

To test the iron hypothesis in the subarctic Pacific Ocean, an in situ iron-enrichment experiment (SEEDS) was performed in the western subarctic gyre in July–August 2001. About 350 kg of iron (as acidic iron sulfate) and 0.48 mol of the inert chemical tracer sulfur hexafluoride were introduced into a 10-m deep surface mixed layer over an 80 km² area. This single iron infusion raised dissolved iron levels to 2.9 nM initially. Dissolved iron concentrations rapidly decreased after the infusion, but levels remained close to 0.15 nM even at the end of the 14-day experimental period. During SEEDS there were iron-mediated increases in chlorophyll a concentrations (up to 20 μg l⁻¹), primary production rates, biomass and photosynthetic energy conversion efficiency relative to waters outside the iron-enriched patch. The rapid and very high accumulation of phytoplankton biomass in response to the iron addition appeared to be partly attributable to shallow mixed-layer depth and moderate water temperature in the western subarctic Pacific. However, the main reason was a floristic shift to fast-growing centric diatom Chaetoceros debilis, unlike the previous iron-enrichment experiments in the equatorial Pacific and the Southern Ocean, in both of which iron stimulated the growth of pennate diatoms. The iron-mediated blooming of diatoms resulted in a marked consumption of macronutrients and drawdown of pCO₂. Biological and physiological measurements indicate that phytoplankton growth in the patch became both light- and iron-limited, making phytoplankton biomass relatively constant after day 9. The increase in microzooplankton grazing rate after day 9 also influenced the net growth rate of phytoplankton. There was no significant increase in the export flux of carbon to depth during the 14-day occupation of the experimental site. The export flux between day 4 and day 13 was estimated to be only 13% of the integrated primary production in the iron-enriched patch. The major part of the carbon fixed by the diatom bloom remained in the surface mixed layer as biogenic particulate matter. Our findings support the hypothesis that iron limits phytoplankton growth and biomass in a ‘bottom up’ manner in this area, but the fate of algal carbon remains unknown.     

Distribution patterns of living benthic foraminifera from Cap Breton canyon,Bay of Biscay: Faunal response to sediment instability

Analysis of living (Rose Bengal-stained) benthic foraminifera in 13 multicorer samples taken along the Cap Breton canyon (Bay of Biscay) revealed that the combination of organic-rich material and sediment instability provides very specific benthic ecosystem conditions. The active canyon hosts different foraminiferal assemblages that appear to be determined by different types and frequencies of environmental disturbance at the sites. Most of them are strongly dominated by shallow-infaunal living taxa that combine a tolerance for low-quality organic matter with a high reproductive potential. Foraminiferal assemblages characterized by high densities, very superficially living taxa and strong dominance of bolivinids and buliminids, follow a poor pioneer fauna dominated by Technitella melo. These assemblages are observed in the narrow canyon axis, where frequent sediment resuspension occurs and affects habitat stability. Assemblages studied from sites outside the canyon axis are still dominated by shallow-infaunal species but show lower foraminiferal densities and higher diversities. Deep-infaunal taxa are only present in some inner meanders and more distal stations. These assemblages are typical for ecological niches that are relatively stable and unaffected by re-sedimentation processes. They have attained a more advanced stage of ecosystem stability. They are influenced by neither lateral sediment nor enriched organic matter input.     

Degradation of malachite green dye by Tenacibaculum sp. HMG1isolated from Pacific deep-sea sediments

A deep-sea bacterium from the Pacific Ocean identified as Tenacibaculum sp. HMG1 was found to have strong malachite green(MG) degradation activity. The MG tolerance and decolorizing activities of strain HMG1 were confirmed by bacterial growth and high-performance liquid chromatography(HPLC) analyses. Strain HMG1 was capable of removing 98.8% of the MG in cultures within 12 h and was able to grow vigorously at 20 mg/L MG. A peroxidase gene detected in the genome of strain HMG1 was found to be involved in the MG biodegradation process. The corresponding recombinant peroxidase(r POD) demonstrated high degradative activity at 1 000 mg/L MG. Based on the common candidate intermediates, strain HMG1 was inferred to have one primary MG degradation pathway containing r POD. In addition, five other candidate intermediates of the r POD-MG degradative process were detected. The optimal conditions for MG degradation were determined and showed that strain HMG1 and the r POD enzyme could maintain high bioactivity at a low temperature(20°C), variable p H values(6.0–9.0), higher salinities(100 mmol/L) and other factors, such as multiple metal ions, H_2O_2 and EDTA.MG-tolerant strain Tenacibaculum sp. HMG1 and its peroxidase have prospective applications as environmental amendments for MG degradation during coastal remediation.     

Stable isotopes in deep-sea living (stained) foraminifera from the Mozambique Channel (eastern Africa): multispecies signatures and paleoenvironmental application

Oxygen and carbon isotopes (δ¹⁸O and δ¹³C) have been investigated in carbonate tests of deep-sea foraminifera living in the Mozambique Channel (eastern Africa) to understand how environmental constraints (e.g., organic matter, oxygenation) control the intra- and interspecific variability of isotopic signatures. 197 living individuals, including eight different species, from various microhabitats within the sediment were sorted from sediment samples gathered at two stations on the Malagasy upper slope. Results show that the δ¹⁸O values of foraminiferal taxa were not controlled by microhabitat pattern. They presented tremendous and intriguing intraspecific variability that is not explained by the classical ontogenetic effect. The δ¹³C values of infaunal foraminiferal taxa do not show a 1:1 relationship with the bottom water δ¹³C DIC and do not present a constant offset from it; instead, they appear to be mainly controlled by a microhabitat effect. The lower δ¹³C values of shallow, intermediate, and deep infaunal taxa at the deeper station compared to those seen at the shallower station reflect the enhanced exportation of sedimentary organic matter at the sediment–water interface, and its related mineralization within the upper sediments. The ?δ¹³C between shallow/very shallow infaunal species (i.e., Hoeglundina elegans, Uvigerina hispida) and intermediate/deep infaunal species (i.e., Melonis barleeanus, Globobulimina barbata) permits insight into (1) the exportation of organic matter to the seafloor and (2) the various degradation pathways for organic detritus in the benthic environments off NW Madagascar.     

‘Live’ benthic foraminifera at an abyssal site in the equatorial Pacific nodule province: Abundance,diversity and taxonomic composition

Replicate sediment samples were obtained from 3 closely spaced stations in the Kaplan East (KE) area of the abyssal eastern Equatorial Pacific (∼15°N, 119°W; ∼4100 m water depth), just below the carbonate compensation depth. At each site, 2 (Stns 827, 838) or 3 (Stn 824) complete cores (57 mm i.d.) were subsampled using 2–3 cut-off syringes of 6.6 cm³ cross-sectional area. The 0–1 cm sediment layers (>32 μm fraction) of these 20 subsamples together yielded 12,513 small, rose-Bengal stained benthic foraminifera dominated by agglutinated taxa, most of them morphologically simple monothalamous types or komokiaceans. Almost two-thirds (65%) of specimens were either obvious fragments, mainly of komokiaceans and tubular foraminifera, or single chambers or small groups of chambers believed to be fragments of very fragile komokiaceans. The remaining 4438 specimens (35%) were considered to be complete individuals. Most (78%) of these complete tests were indeterminate agglutinated spheres (termed ‘psammosphaerids’) that constituted 27.6% of all specimens (complete plus fragments). Complete individuals that could be assigned to either described or undescribed species accounted for 983 specimens (22% of complete tests=7.6% of all specimens); only 26 specimens (0.59% of complete individuals) were calcareous and these had invariably lost their tests through dissolution. Some groups exhibited considerable spatial heterogeneity. For example, 45% of the 3455 indeterminate psammosphaerids and 45% of the 3087 Komokiacean-like chambers occurred in single subcores. A total of 252 morphospecies was recognised; 168 were represented by complete individuals and 84 by fragments. There are clear differences between these Pacific assemblages and those from other oceans; in particular, psammosphaerids and isolated komokiacean chambers appear to be much more prevalent in the Pacific compared to the Atlantic Ocean. Some morphospecies present in Kaplan samples are known from the Atlantic but many are not. Such species may either (1) be ubiquitous but undersampled because they are rare or (2) have geographically patterned distributions. Without further sampling, there is no way to distinguish between these 2 possibilities. Fossilisable tests represent a very small component of the KE assemblage. Many of the delicate, monothalamous species that have little fossilisation potential, including the komokiaceans, accumulate stercomata (waste pellets) and may consume organic material and bacteria associated with sediment. Because of their enormous abundance at abyssal depths, these poorly known taxa probably play a substantial role in carbon cycling over vast areas of the Pacific seafloor.     

An optimized method for automated analysis of algal pigments by HPLC

A recent development in algal pigment analysis by high-performance liquid chromatography (HPLC) is the application of automation. An optimization of a complete sampling and analysis protocol applied specifically in automation has not yet been performed. In this paper we show that automation can only be successful if the various methodological aspects of the sampling and analysis protocol are considered in coherence. We introduce an optimized protocol that involves freeze-drying of the sample, subsequent extraction in 90% acetone and the application of water-packing during analysis. The method was evaluated on both natural plankton populations and a broad spectrum of microalgal cultures: Thalassiosira weisflogii (Bacillariophyceae), Emiliania huxleyi (Prymnesiophyceae), Phaeocystis globosa and Phaeocystis antarctica (Prymnesiophyceae) and Pyramimonas sp. (Prasinophyceae). Whereas pigment extracts were unstable in methanol, with recorded chlorophyll a losses from 10% to 60% per day, pigment degradation rates in acetone were generally less than 1% over 18 h storage in the autosampler (4 °C). In addition, it was found that the extraction efficiency of acetone significantly increased upon freeze-drying prior to extraction. Increases as high as 50–60% were measured in P. antarctica. The application of water-packing of the sample during injection resulted in improved peak shape and peak separation, without diluting the pigment concentrations. Automation is especially beneficial for application in the field, when mixed algal assemblages and low biomass put a high demand on the sensitivity as well as reproducibility of the method.