Late Quaternary benthic foraminiferal record of the Bounty Trough, east of New Zealand

The Bounty Trough, east of New Zealand, lies along the southeastern edge of the present-day Subtropical Front (STF), and is a major conduit via the Bounty Channel, for terrigenous sediment supply from the uplifted Southern Alps to the abyssal Bounty Fan. Census data on 65 benthic foraminiferal faunas (>63 µm) from upper bathyal (ODP 1119), lower bathyal (DSDP 594) and abyssal (ODP 1122) sequences, test and refine existing models for the paleoceanographic and sedimentary history of the trough through the last 150 ka (marine isotope stages, MIS 6-1). Cluster analysis allows recognition of six species groups, whose distribution patterns coincide with bathymetry, the climate cycles and displaced turbidite beds. Detrended canonical correspondence analysis and comparisons with modern faunal patterns suggest that the groups are most strongly influenced by food supply (organic carbon flux), and to a lesser extent by bottom water oxygen and factors relating to sediment type. Major faunal changes at upper bathyal depths (1119) probably resulted from cycles of counter-intuitive seaward-landward migrations of the Southland Front (SF) (north-south sector of the STF). Benthic foraminiferal changes suggest that lower nutrient, cool Subantarctic Surface Water (SAW) was overhead in warm intervals, and higher nutrient-bearing, warm neritic Subtropical Surface Water (STW) was overhead in cold intervals. At lower bathyal depths (594), foraminiferal changes indicate increased glacial productivity and lowered bottom oxygen, attributed to increased upwelling and inflow of cold, nutrient-rich, Antarctic Intermediate Water (AAIW) and shallowing of the oxygen-minimum zone (upper Circum Polar Deep Water, CPDW). The observed cyclical benthic foraminiferal changes are not a result of associations migrating up and down the slope, as glacial faunas (dominated by Globocassidulina canalisuturata and Eilohedra levicula at upper and lower bathyal depths, respectively) are markedly different from those currently living in the Bounty Trough. On the abyssal Bounty Fan (1122), faunal changes correlate most strongly with grain size, and are attributed to varying amounts of mixing of displaced and in-situ faunas. Most of the displaced foraminifera in turbiditic sand beds are sourced from mid-outer shelf depths at the head of the Bounty Channel. Turbidity currents were more prevalent during, but not restricted to, glacial intervals.

Benthic foraminifera across the Cretaceous/Paleogene boundary in ODP Hole 198-1210A

Deep-sea benthic foraminifera show important but transient assemblage changes at the Cretaceous/Paleogene (K/Pg) boundary, when many biota suffered severe extinction. We quantitatively analyzed benthic foraminiferal assemblages from lower bathyal-upper abyssal (1500-2000 m) northwest Pacific ODP Site 1210 (Shatsky Rise) and compared the results with published data on assemblages at lower bathyal (~ 1500 m) Pacific DSDP Site 465 (Hess Rise) to gain insight in paleoecological and paleoenvironmental changes at that time. At both sites, diversity and heterogeneity rapidly decreased across the K/Pg boundary, then recovered. Species assemblages at both sites show a similar pattern of turnover from the uppermost Maastrichtian into the lowermost Danian: 1) The relative abundance of buliminids (indicative of a generally high food supply) increases towards the uppermost Cretaceous, and peaks rapidly just above the K/Pg boundary, coeval with a peak in benthic foraminiferal accumulation rate (BFAR), a proxy for food supply. 2) A peak in relative abundance of Stensioeina beccariiformis, a cosmopolitan form generally more common at the middle than at the lower bathyal sites, occurs just above the buliminid peak. 3) The relative abundance of Nuttallides truempyi, a more oligotrophic form, decreases at the boundary, then increases above the peak in Stensioeina beccariiformis. The food supply to the deep sea in the Pacific Ocean thus apparently increased rather than decreased in the earliest Danian. The low benthic diversity during a time of high food supply indicates a stressed environment. This stress might have been caused by reorganization of the planktic ecosystem: primary producer niches vacated by the mass extinction of calcifying nannoplankton may have been rapidly (<10 kyr) filled by other, possibly opportunistic, primary producers, leading to delivery of another type of food, and/or irregular food delivery through a succession of opportunistic blooms. The deep-sea benthic foraminiferal data thus are in strong disagreement with the widely accepted hypothesis that the global deep-sea floor became severely food-depleted following the K/Pg extinction due to the mass extinction of primary producers ("Strangelove Ocean Model") or to the collapse of the biotic pump ("Living Ocean Model").

Relative abundances of twenty-eight dominant benthic foraminifera species of DSDP Hole 23-219 (Table 1s)

Tropical climate is variable on astronomical time scale, driving changes in surface and deep-sea fauna during the Pliocene-Pleistocene. To understand these changes in the tropical Indian Ocean over the past 2.36 Myr, we quantitatively analyzed deep-sea benthic foraminifera and selected planktic foraminifera from >125 µm size fraction from Deep Sea Drilling Project Site 219. The data from Site 219 was combined with published foraminiferal and isotope data from Site 214, eastern Indian Ocean to determine the nature of changes. Factor and cluster analyses of the 28 highest-ranked species distinguished four biofacies, characterizing distinct deep-sea environmental settings. These biofacies have been named after their most dominant species such as Stilostomella lepidula-Pleurostomella alternans (Sl-Pa), Nuttallides umbonifer-Globocassidulina subglobosa (Nu-Gs), Oridorsalis umbonatus-Gavelinopsis lobatulus (Ou-Gl) and Epistominella exigua-Uvigerina hispido-costata (Ee-Uh) biofacies. Biofacies Sl-Pa ranges from ~2.36 to 0.55 Myr, biofacies Nu-Gs ranges from ~1.9 to 0.65 Myr, biofacies Ou-Gl ranges from ~1 to 0.35 Myr and biofacies Ee-Uh ranges from 1.1 to 0.25 Myr. The proxy record indicates fluctuating tropical environmental conditions such as oxygenation, surface productivity and organic food supply. These changes appear to have been driven by changes in monsoonal wind intensity related to glacial-interglacial cycles. A shift at ~1.2-0.9 Myr is observed in both the faunal and isotope records at Site 219, indicating a major increase in monsoon-induced productivity. This coincides with increased amplitude of glacial cycles, which appear to have influenced low latitude monsoonal climate as well as deep-sea conditions in the tropical Indian Ocean.

Lead 210 concentrations in sediments of the Arabian Sea

In order to evaluate bioturbation in abyssal Arabian-Sea sediments of the Indus fan profiles of 210Pb (half-life: 22.3 yr) and 234Th (half-life: 24.1 d) were measured in cores collected during September and October 1995 and April 1997, respectively. The density and composition of epibenthic megafauna and lebensspuren were determined in vertical seafloor photographs during April 1997. Mean eddy-diffusive mixing coefficients according to the distribution of excess 210Pb ( 210Pb-DB) were 0.072±0.028, 0.068±0.055, 0.373±0.119, 0.037±0.009 and 0.079±0.119 cm**2 yr**-1 in the northern, western, central, eastern and southern abyssal Arabian sea, respectively. Mean eddy-diffusive mixing coefficients according to the distribution of excess 234Th (234Th-DB) were 0.53, 1.64 and 0.47 cm**2 yr**-1 in the northern, western and central abyssal Arabian Sea, respectively. Mobile epibenthic megafauna at the western, northern, central and southern study sites were dominated by ophiuroids, holothurians, ophiuroids and natant decapods (the respective densities were 100, 82, 29 and 6 individuals 1000 m**-2). The northern study site was characterized by a high abundance of spoke traces and fecal casts. The central site showed spoke traces and many tracks. The southern site displayed the highest abundance of spoke traces, whereas at the western site hardly any lebensspuren were observed. There is evidence for at least two functional endmember communities in the Arabian Sea. In the northwestern Arabian Sea (WAST) vertical particle displacement seems to be dominated by macrofauna and primarily eddy-diffusive. In the southern Arabian Sea (SAST) non-local and 'incidental' mixing due to spoke-trace producers might become more important and superimpose reduced eddy-diffusive mixing. With respect to biological data CAST is an intermediate location. Given the biological data, average 210Pb-DB is higher and decimeter-scale variability of 210Pb-DB smaller at CAST than expected. These findings indicate that in a mixture of both endmember communities the organisms may interact in way that increases values of biodiffusivity, as reflected by 210Pb-DB, and reduces decimeter-scale 210Pb-DB heterogeneity in comparison to the simple sum of the isolated effects of the endmembers. For time scales <100 years there was no evidence for a relationship between food supply (POC flux) and bioturbation intensity, as reflected by 210Pb-DB and 234Th-DB. Bioturbation intensity should be controlled primarily by the composition of the benthic fauna, its specific adaptation to the environmental setting, and the abundance of each species of the benthic community. Food supply can have only an indirect influence on bioturbation intensity. In certain parts of the ocean the a priori overall positive relationship between POC flux and biodiffusivity might include restricted intervals displaying no or even negative relations.

Live benthic foraminifera in surface sediments of the Indo-Pakistan continental margin

The Indo-Pakistan Continental Margin represents an extreme habitat for benthic foraminifera since (1) high fluxes of organic matter offer a high food supply, (2) an intensified oxygen minimum Zone (OMZ) develops from the base of the euphotic Zone to water depths over 1000 m and (3) the monsoon causes seasonal oscillations within the biogeochemical cycle. At three stations from the uppermost (233 m), the central (658 m) and the deeper part (902 m) of the OMZ, living benthic foraminiferal assemblages were analyzed within the uppermost 10 cm of the sediment column. The ecologic structure of foraminiferal faunas is characterized by high abundances at the sediment surface and a rapid decrease within the uppermost 2 cm of the sediment column. Despite dysoxic to suboxic bottom-water conditions, stained benthic foraminifera occurred in all cores down to the base of the sampled interval. High surface abundances, a high dominance by few endobenthic calcareous taxa and a low diversity, which may result from specific physiological adaptations to almost anoxic conditions and the absence of predators, are recognized in the central part of the OMZ. The upper and lower margins of the OMZ are characterized by higher diversities and lower abundances. The shallowest part of the OMZ is dominated by calcareous foraminifera, whereas agglutinated species are the most common taxa in the deeper part. Comparisons with previous studies show that benthic foraminiferal assemblages, that are influenced by seasonal oscillations controlling food supply and/or the availability of oxygen, show variations in faunal density and species composition. Since there is strong evidence that oxygen is not a limiting factor for some taxa, it seems more likely that the distribution pattern of benthic foraminifera is preferentially controlled by trophic conditions.

Abundance and biomass of in- and epibenthic invertebrate macrofauna in the German Bight from 1969 to 2000

In order to examine the long-term development of offshore macrozoobenthic soft-bottom communities of the German Bight, four representative permanent stations (MZB-SSd, -FSd, -Slt, -WB) have been sampled continuously since 1969. Inter-annual variability and possible long-term trends were analysed based on spring-time samples from 1969 until 2000. This is part of the ecological long-term series of the AWI and is supplemented by periodic large-scale mapping of the benthos. The main factors influencing the development of the benthic communities are biological interactions, climate, food supply (eutrophication) and the disturbance regime. The most frequent disturbances are sediment relocations during strong storms or by bottom trawling, while occasional oxygen deficiencies and extremely cold winters are important disturbance events working on a much larger scale. Benthic communities at the sampling stations show a large inter-annual variability combined with a variation on a roughly decadal scale. In accordance with large-scale system shifts reported for the North Sea, benthic community transitions occurred between roughly the 1970ies, 80ies and 90ies. The transitions between periods are not distinctly marked by strong changes but rather reflected in gradual changes of the species composition and dominance structure.

Age determination of sediment cores from the Northeast Atlantic

Radiocarbon dating series, bulk sediment, and organic carbon flux from various Atlantic deep-sea regions reveal that the thickness of the bioturbated zone increases by 2 cm if food supply increases by 1 gC/m**2/yr (r = 0.8). Bulk sediment accumulation rates do not influence the depth of bioturbational mixing under normal pelagic sedimentary conditions. We believe that this relationship between nutrient supply and benthic mixing can be used for a quantitative and time-variable unmixing procedure to improve high-resolution stratigraphic correlations and paleoclimatic interpretations of deep-sea records.

(Appendix) Analysis of benthic foraminifera from ODP Leg 181 and DSDP Leg 90

Canonical correspondence analysis indicates that the distribution of Neogene benthic foraminiferal faunas (>63 µm) in seven DSDP and ODP sites (500-4500 m water depth) east of New Zealand (38-51°S, 170°E-170°W) is most strongly influenced by depth (water mass stratification), and secondly by age (palaeoceanographic changes influencing faunal composition and biotic evolution). Stratigraphic faunal changes are interpretted in terms of the pulsed sequential development of southern, and later northern, polar glaciation and consequent cooling of bottom waters, increased vertical and lateral stratification of ocean water masses, and increased overall and seasonal surface water productivity. Oligocene initiation of the Antarctic Circumpolar Current and Deep Western Boundary Current (DWBC), flowing northwards past New Zealand, resulted in extensive hiatuses throughout the Southwest Pacific, some extending through into the Miocene. Planktic foraminiferal fragmentation index values indicate that carbonate dissolution was significant at abyssal depths throughout most of the Neogene, peaking at upper abyssal depths in the late Miocene (11-7 Ma), with the lysocline progressively deepened thereafter. Miocene abyssal faunas are dominated by Globocassidulina subglobosa and Oridorsalis umbonatus, with increasing Epistominella exigua after 16 Ma at upper abyssal depths. Peak abundances of Epistominella umbonifera indicate increased input of cold Southern Component Water to the DWBC at 7-6 Ma. Faunal association changes imply establishment of the modern Oxygen Minimum Zone (upper Circumpolar Deep Water) in the latest Miocene. Significant latitudinal differences between the benthic foraminiferal faunas at lower bathyal depths indicate the existence of an oceanic front along the Chatham Rise (location of present Subtropical Front), since the early late Miocene at least, with more pulsed productivity (higher E. exigua) along the south side. Modern Antarctic Intermediate Water faunal associations were established north of the Chatham Rise at 10-9 Ma, and south of it at 3-1.5 Ma. Middle-upper bathyal faunas on the Campbell Plateau are dominated by reticulate bolivinids during the early and middle Miocene, indicative of sustained productivity above relatively sluggish, suboxic bottom waters. Faunal changes and hiatuses indicate increased current vigour over the Campbell Plateau from the latest Miocene on. Surface water productivity (food supply) appears to have increased in three steps (at times of enhanced global cooling) marked by substantially increased relative abundance of: (1) Abditodentrix pseudothalmanni, Alabaminella weddellensis, Cassidulina norvangi (16-15 Ma, increased pulsed productivity); (2) Bulimina marginata f. aculeata, Nonionella auris, Trifarina angulosa, Uvigerina peregrina (3-1.5 Ma, increased overall productivity); and (3) Cassidulina carinata (1-0.5 Ma, increased overall productivity). Three intervals of deep-sea benthic foraminiferal taxonomic turnover are recognised (16-15, 11.5-10, 2-0.5 Ma) corresponding to intervals of enhanced global cooling and possible productivity changes. The late Pliocene-middle Pleistocene extinction, associated with increasing Northern Hemisphere glaciation, culminating in the middle Pleistocene climatic transition, was more significant in the study area than the earlier Neogene turnovers.

One size fits all: stability of metabolic scaling under warming and ocean acidification in echinoderms

Responses by marine species to ocean acidification (OA) have recently been shown to be modulated by external factors including temperature, food supply and salinity. However the role of a fundamental biological parameter relevant to all organisms, that of body size, in governing responses to multiple stressors has been almost entirely overlooked. Recent consensus suggests allometric scaling of metabolism with body size differs between species, the commonly cited 'universal' mass scaling exponent (b) of ¾ representing an average of exponents that naturally vary. One model, the Metabolic-Level Boundaries hypothesis, provides a testable prediction: that b will decrease within species under increasing temperature. However, no previous studies have examined how metabolic scaling may be directly affected by OA. We acclimated a wide body-mass range of three common NE Atlantic echinoderms (the sea star Asterias rubens, the brittlestars Ophiothrix fragilis and Amphiura filiformis) to two levels of pCO2 and three temperatures, and metabolic rates were determined using closed-chamber respirometry. The results show that contrary to some models these echinoderm species possess a notable degree of stability in metabolic scaling under different abiotic conditions; the mass scaling exponent (b) varied in value between species, but not within species under different conditions. Additionally, we found no effect of OA on metabolic rates in any species. These data suggest responses to abiotic stressors are not modulated by body size in these species, as reflected in the stability of the metabolic scaling relationship. Such equivalence in response across ontogenetic size ranges has important implications for the stability of ecological food webs.

Seawater carbonate chemistry and biological processes of Mytilus edulis during experiments, 2011

Progressive ocean acidification due to anthropogenic CO2 emissions will alter marine ecosytem processes. Calcifying organisms might be particularly vulnerable to these alterations in the speciation of the marine carbonate system. While previous research efforts have mainly focused on external dissolution of shells in seawater under saturated with respect to calcium carbonate, the internal shell interface might be more vulnerable to acidification. In the case of the blue mussel Mytilus edulis, high body fluid pCO2 causes low pH and low carbonate concentrations in the extrapallial fluid, which is in direct contact with the inner shell surface. In order to test whether elevated seawater pCO2 impacts calcification and inner shell surface integrity we exposed Baltic M. edulis to four different seawater pCO2 (39, 142, 240, 405 Pa) and two food algae (310-350 cells mL-1 vs. 1600-2000 cells mL-1) concentrations for a period of seven weeks during winter (5°C). We found that low food algae concentrations and high pCO2 values each significantly decreased shell length growth. Internal shell surface corrosion of nacreous ( = aragonite) layers was documented via stereomicroscopy and SEM at the two highest pCO2 treatments in the high food group, while it was found in all treatments in the low food group. Both factors, food and pCO2, significantly influenced the magnitude of inner shell surface dissolution. Our findings illustrate for the first time that integrity of inner shell surfaces is tightly coupled to the animals' energy budget under conditions of CO2 stress. It is likely that under food limited conditions, energy is allocated to more vital processes (e.g. somatic mass maintenance) instead of shell conservation. It is evident from our results that mussels exert significant biological control over the structural integrity of their inner shell surfaces.

Station characteristics and copepod fatty alcohol, fatty acid and lipid composition during MSM02/4 in Fram Strait

The biodiversity of pelagic deep-sea ecosystems has received growing scientific interest in the last decade, especially in the framework of international marine biodiversity initiatives, such as Census of Marine Life (CoML). While a growing number of deep-sea zooplankton species has been identified and genetically characterized, little information is available on the mechanisms minimizing inter-specific competition and thus allowing closely related species to co-occur in the deep-sea pelagic realm. Focussing on the two dominant calanoid copepod families Euchaetidae and Aetideidae in Fram Strait, Arctic Ocean, the present study strives to characterize ecological niches of co-occurring species, with regard to vertical distribution, dietary composition as derived from lipid biomarkers, and trophic level on the basis of stable isotope signatures. Closely related species were usually restricted to different depth layers, resulting in a multi-layered vertical distribution pattern. Thus, vertical partitioning was an important mechanism to avoid inter-specific competition. Species occurring in the same depth strata usually belonged to different genera. They differed in fatty acid composition and trophic level, indicating different food preferences. Herbivorous Calanus represent major prey items for many omnivorous and carnivorous species throughout the water column. The seasonal and ontogenetic vertical migration of Calanus acts as a short-cut in food supply for pelagic deep-sea ecosystems in the Arctic.

(Table 1) Time series of ostracod population density off Bokniseck shore ('Hausgarten'), western Baltic Sea

Seasonal distributions of ostracode species from the Bay of Kiel, western Baltic Sea (Bokniseck, 'Hausgarten') were studied at seven observation stations located between 6 - 23.5 m water depth. During the period from December 1973 to March 1975, 175 samples were taken every two weeks from the sediment surface at each station. Environmental factors were measured simultaneously with sampling. Most of the ostracode species were present in all of the samples throughout the year; four species were found to be seasonal. On the other hand, the population densities of each of the examined ostracode species, as revealed by countings, indicate marked seasonal variations. These variations are attributed to changes in length rate and timing of the reproductive cycles of the different ostracode species, effected mainly by food supply. Maxima in the abundance of the ostracode species with short (1/2 - 1 month) life cycles, occurred simultaneously with peaks of food supply in spring and autumn, whereas the maxima abundance of ostracodes with longer (10 - 12 months) life cycles was delayed.

Investigation of the seasonality of N. pachyderma in the North Atlantic Ocean during Heinrich Stadial 1 using an ecosystem modeling approach

Fossil shells of planktonic foraminifera serve as the prime source of information on past changes in surface ocean conditions. Because the population size of planktonic foraminifera species changes throughout the year, the signal preserved in fossil shells is biased towards the conditions when species production was at its maximum. The amplitude of the potential seasonal bias is a function of the magnitude of the seasonal cycle in production. Here we use a planktonic foraminifera model coupled to an ecosystem model to investigate to what degree seasonal variations in production of the species Neogloboquadrina pachyderma may affect paleoceanographic reconstructions during Heinrich Stadial 1 (~18-15 cal. ka B.P.) in the North Atlantic Ocean. The model implies that during Heinrich Stadial 1 the maximum seasonal production occurred later in the year compared to the Last Glacial Maximum (~21-19 cal. ka B.P.) and the pre-industrial era north of 30 ºN. A diagnosis of the model output indicates that this change reflects the sensitivity of the species to the seasonal cycle of sea-ice cover and food supply, which collectively lead to shifts in the modeled maximum production from the Last Glacial Maximum to Heinrich Stadial 1 by up to six months. Assuming equilibrium oxygen isotopic incorporation in the shells of N. pachyderma, the modeled changes in seasonality would result in an underestimation of the actual magnitude of the meltwater isotopic signal recorded by fossil assemblages of N. pachyderma wherever calcification is likely to take place.