Lake sediments from Store Koldewey, Northeast Greenland

A 2.9 m long sedimentary record was studied from a small lake, here referred to as Duck Lake, located at 76°25'N, 18°45'W on Store Koldewey, an elongated island off the coast of Northeast Greenland. The sediments were investigated for their geophysical and biogeochemical characteristics, and for their fossil chironomid assemblages. Organic matter began to accumulate in the lake at 9.1 cal. kyr BP, which provides a minimum age for the deglaciation of the basin. Although the early to mid-Holocene is known as a thermal maximum in East Greenland, organic matter accumulation in the lake remained low during the early Holocene, likely due to late plant immigration and lack of nutrient availability. Organic matter accumulation increased during the middle and late Holocene, when temperatures in East Greenland gradually decreased. Enhanced soil formation probably led to higher nutrient availability and increased production in the lake. Chironomids are abundant throughout the record after 9.1 cal. kyr BP and seem to react sensitively to changes in temperature and nutrient availability. It is concluded that relative temperature reconstructions based on biogeochemical data have to be regarded critically, particularly in the period shortly after deglaciation when nutrient availability was low. Chironomids may be a suitable tool for climatic reconstructions even in those high arctic environments. However, a better understanding of the ecology of chironomids under these extreme conditions is needed.

Sedimentology and paleontology on core Lz1104 from Lake Hjort, Greenland

Two sediment cores of 70 and 252 cm length were recovered from Hjort Sø, a small lake on Store Koldewey, Northeast Greenland, and studied with a multidisciplinary approach in order to reconstruct the local environmental history and to test the relevance of proxies for paleoenvironmental information. The basal sediments from the longer core are dominated by clastic matter, which was likely deposited during deglaciation of the lake basin. These clastic sediments are overlain by gyttja, which is also present throughout the shorter core. AMS radiocarbon dating was conducted on plant macrofossils of 11 samples from the gyttja in both cores. A reliable chronology was established for both cores, which dated the onset of organic accumulation at 9,500 cal. year BP. The Holocene temperature development, with an early to mid Holocene thermal maximum, is best reflected in the grain-size composition. Nutrient availability was apparently low during the early Holocene and led to low productivity in the lake and its vicinity. From ca. 7,000 cal. year BP, productivity in the lake increased significantly, probably induced by external nutrient input from goose excrements. From this time, micro- and macrofossil remains reflect relatively well the climate history of East Greenland, with a cooling during the middle Holocene, the medieval warming, and the Little Ice Age. The amount of organic matter in the sequence seems to be more affected by lake ice cover or by nutrient supply from the catchment than by temperature changes. The record from Hjort Sø thus reveals the difficulties in interpreting sedimentary records from high arctic regions.

Layer description, carbon and nitrogen content and stable isotope record of bulk sedimentary organic matter from Laguna Potrok Aike

An investigation of stable isotope (d13C TOC and d15N TN) and elemental parameters (TOC, TN contents and TOC/TN ratios) of bulk organic matter (<200 µm) from sediment cores recovered from the Patagonian lake Laguna Potrok Aike (Argentina) in the framework of the ICDP deep drilling project PASADO provided insights into past changes in lake primary productivity and environmental conditions in South Patagonia throughout the last Glacial-Interglacial transition. Stratigraphically constrained cluster analyses of all proxy parameters suggest four main phases. From ca 26,100 to 17,300 cal. years BP, lacustrine phytoplankton was presumably the predominant organic matter source in an aquatic environment with low primary productivity rates. At around 17,300 cal. years BP, abrupt and distinct shifts of isotopic and elemental values indicate that the lacustrine system underwent a rapid reorganization. Lake primary productivity (phytoplankton and aquatic macrophytes) shows higher levels albeit with large variations during most of the deglaciation until 13,000 cal. years BP. The main causes for this development can be seen in improved growing conditions for primary producers because of deglacial warming in combination with expedient availability of nutrients and likely calm wind conditions. After 13,000 cal. years BP, decreased d13C TOC values, TOC, TN contents and TOC/TN ratios indicate that the lake approached a new state with reduced primary productivity probably induced by unfavourable growing conditions for primary producers like strengthened winds and reduced nutrient availability. The steady increase in d15N TN values presumably suggests limitation of nitrate supply for growth of primary producers resulting from a nutrient shortage after the preceding phase with high productivity. Nitrate limitation and consequent decreased lacustrine primary productivity continued into the early Holocene (10,970-8400 cal. years BP) as reflected by isotopic and elemental values.

Abundance of living coccolithophores in the northern Arabian Sea (Table 2)

The present study was conducted to provide information about living coccolithophores from the northern Arabian Sea as potential proxies in palaeoceanographic studies. In all, 71 plankton samples from 16 stations collected in September 1993 were analysed for their contents of living coccolithophores. Absolute abundances range from less than 400 coccospheres per litre in surface waters to 35 000 spheres per litre at intermediate water depths. From 49 identified taxa, nine species contribute significant cell numbers of more than 2000 coccospheres per litre and comprise more than 10% of the communities in at least one sample. Important species are (in approximate order of cell abundances): Gephyrocapsa oceanica, Florisphaera profunda, Oolithotus antillarum, Calciosolenia murrayi, Umbellosphaera irregularis, Emiliania huxleyi, Umbellosphaera tenuis, Calciopappus rigidus, and Algirosphaera robusta. At most profiles, a vertical succession of coccolithophore species was found. Calciosolenia murrayi and C. rigidus were restricted to surface waters, whereas high numbers of F. profunda and A. robusta occurred at depths below 40 m. The coccolithophore communities reflected the local oceanographic situation and seemed to be more dependent on mixed layer depth and nutrient availability than on temperature and salinity changes. Additionally, synecologic competition with diatoms in part controlled the species composition and generally reduced the abundance of coccolithophores. Synecological and ecological tolerances of species were discussed with the help of cluster analysis.

A 34-month sediment trap experiment at the Cape Verde Ocean Observatory (CVOO, December 2009 - October 2012)

Anticyclonic mesoscale eddies (ACME) have been proposed as a mechanism by which new nutrients are episodically delivered into the euphotic zone, thereby enhancing new production as well as shifting phytoplankton community structure. In this paper, we report on a 34-month sediment trap experiment at the Cape Verde Ocean Observatory (CVOO; ca. 18°N, 24°E; December 2009-October 2012), occasionally influenced by ACME passages. The typically oligotrophic, weakly seasonal particle flux pattern at the CVOO is strongly modified by the appearance of a highly productive and low oxygen ACME. Out of four recorded diatom flux maxima at CVOO, three were associated with the passage of ACMEs. The recorded diatom maxima events support the view that local ACME dynamics promotes upward nutrient supply into the euphotic zone leading to a rapid response of diatoms. This response is clearly reflected by the flux seasonality: between 40% and 60% of the total annual diatom flux at the CVOO site was intercepted in a relatively short time interval (<60 days). A highly diverse diatom community characterized the diatom fluxes throughout. Along with the ACME passages, small species of the genus Nitzschia, and Thalassionema nitzschioides var. parva dominated and delivered a major portion of the opal and organic carbon into deeper waters at site CVOO. Several pelagic, warm-water background species became dominant during intervals with low nutrient availability in the euphotic zone. Results of our interannual time-series suggest that ACMEs impact on total diatom production and the species-specific composition of the assemblage north of the Cave Verde Islands, and can strengthen the biological pump in open-ocean, oligotrophic subtropical regions of the world ocean. Our observations are useful for testing biogeochemical ocean models and will also help in improving the knowledge of processes and mechanisms behind interannual time-series of bulk components and microorganisms in pelagic and hemipelagic ocean areas.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Biogenic opal, carbonat concentrations and stable oxygen isotope ratios of foraminifera from sediment cores of the Southern Ocean

We present records of biogenic opal percentage and burial rate in 12 piston cores from the Atlantic and Indian sectors of the Southern Ocean. These records provide a detailed, quantitative description of changing patterns of opal deposition over the last 450 kyr. The striking regional coherence of these records suggests that dissolution in the deep sea and sediment pore waters does not obscure the surface productivity signal, and therefore these opal time series can be used in combination with other surface water tracers to make inferences about the chemistry and circulation of the Southern Ocean under different global climate conditions. Three broad depositional patterns can be distinguished. Northernmost records (39°-42°S latitude) are characterized by enhanced opal burial during glacial periods and strong 41 kyr periodicity. Records from cores just north of the present Antarctic Polar Front (46°-49°S) show even larger increases in opal burial rate during glacial intervals, but have variance concentrated in the 100 and 23 kyr bands. Southernmost records (51°-55°S) are completely out of phase with those to the north, with greatly reduced opal burial rates during glacial periods. Taken as a whole, the opal records show no evidence for the increased total Antarctic productivity predicted by recent geochemical models of atmospheric CO2 variability. The areal expansion of Southern Ocean sea ice over the present zone of high siliceous productivity provides one plausible explanation for the glacial-interglacial opal patterns. The excess silica not taken up in this zone during glacial periods would contribute to greater nutrient availability and thus higher productivity in the subantarctic region. However, local circulation changes may act to modify this basic signal, possibly accounting for the observed differences in the opal variance spectra.

N2 fixation in eddies of the eastern tropical South Pacific Ocean

Mesoscale eddies play a major role in controlling ocean biogeochemistry. By impacting nutrient availability and water column ventilation, they are of critical importance for oceanic primary production. In the eastern tropical South Pacific Ocean off Peru, where a large and persistent oxygen-deficient zone is present, mesoscale processes have been reported to occur frequently. However, investigations into their biological activity are mostly based on model simulations, and direct measurements of carbon and dinitrogen (N2) fixation are scarce. We examined an open-ocean cyclonic eddy and two anticyclonic mode water eddies: a coastal one and an open-ocean one in the waters off Peru along a section at 16°S in austral summer 2012. Molecular data and bioassay incubations point towards a difference between the active diazotrophic communities present in the cyclonic eddy and the anticyclonic mode water eddies. In the cyclonic eddy, highest rates of N2 fixation were measured in surface waters but no N2 fixation signal was detected at intermediate water depths. In contrast, both anticyclonic mode water eddies showed pronounced maxima in N2 fixation below the euphotic zone as evidenced by rate measurements and geochemical data. N2 fixation and carbon (C) fixation were higher in the young coastal mode water eddy compared to the older offshore mode water eddy. A co-occurrence between N2 fixation and biogenic N2, an indicator for N loss, indicated a link between N loss and N2 fixation in the mode water eddies, which was not observed for the cyclonic eddy. The comparison of two consecutive surveys of the coastal mode water eddy in November 2012 and December 2012 also revealed a reduction in N2 and C fixation at intermediate depths along with a reduction in chlorophyll by half, mirroring an aging effect in this eddy. Our data indicate an important role for anticyclonic mode water eddies in stimulating N2 fixation and thus supplying N offshore.

(Appendix B) Dinoflagellate cysts in Southeast Pacific Ocean surface sediments

orty-eight surface sediment samples from the southeast (SE) Pacific (25-53°S) are investigated for the determination of the spatial distribution of organic-walled dinoflagellate cysts along the western South American continental margin. Fifty-five different taxa are recorded and reflect oceanic or coastal assemblages. The oceanic assemblages are characterised by low cyst concentrations and the dominance of autotrophs, while the coastal assemblages generally contain a higher number of cysts, which are mainly produced by heterotrophic species. Highest cyst concentrations are observed in the active upwelling system offshore Concepción (35-37°S). Brigantedinium spp., Echinidinium aculeatum, Echinidinium granulatum/delicatum and cysts of Protoperidinium americanum dominate assemblages related to upwelling. Echinidinium aculeatum appears to be the best indicator for the presence of all year round active upwelling cells. Other protoperidinioid cysts may also occur in high relative abundances in coastal regions outside active upwelling systems, if the availability of nutrients, co-responsible for the presence/absence of their main food sources such as diatoms and other protists, is sufficient. The importance of nutrient availability as a determining environmental variable influencing cyst signals on a regional scale (SE Pacific) is demonstrated through statistical analyses of the data. Because of the importance of nutrients, uncertainties about the outcomes of quantitative sea-surface temperature (SST) reconstructions (Modern Analogue Technique) based on dinoflagellate cysts may arise, since no interaction between different hydrographical variables is considered in this approach. The combination of the SE Pacific surface sample dataset with other published cyst data from the Southern Hemisphere resulted in a database which includes 350 samples: the 'SH350 database'. This database is used to test the accuracy of the quantitative reconstructions by calculating and comparing the estimated versus observed values for each site. An attempt to perform quantitative SST reconstructions on the last 25 cal ka of site ODP1233 (41°S; 74°27'W) is made and again stresses the importance of other environmental variables such as nutrient availability in determining the dinoflagellate cyst assemblages.

(Table 1) Types of giant magnetofossils in ODP holes

Magnetotactic bacteria biomineralize magnetic minerals with precisely controlled size, morphology, and stoichiometry. These cosmopolitan bacteria are widely observed in aquatic environments. If preserved after burial, the inorganic remains of magnetotactic bacteria act as magnetofossils that record ancient geomagnetic field variations. They also have potential to provide paleoenvironmental information. In contrast to conventional magnetofossils, giant magnetofossils (most likely produced by eukaryotic organisms) have only been reported once before from Paleocene-Eocene Thermal Maximum (PETM; 55.8 Ma) sediments on the New Jersey coastal plain. Here, using transmission electron microscopic observations, we present evidence for abundant giant magnetofossils, including previously reported elongated prisms and spindles, and new giant bullet-shaped magnetite crystals, in the Southern Ocean near Antarctica, not only during the PETM, but also shortly before and after the PETM. Moreover, we have discovered giant bullet-shaped magnetite crystals from the equatorial Indian Ocean during the Mid-Eocene Climatic Optimum (~40 Ma). Our results indicate a more widespread geographic, environmental, and temporal distribution of giant magnetofossils in the geological record with a link to "hyperthermal" events. Enhanced global weathering during hyperthermals, and expanded suboxic diagenetic environments, probably provided more bioavailable iron that enabled biomineralization of giant magnetofossils. Our micromagnetic modelling indicates the presence of magnetic multi-domain (i.e., not ideal for navigation) and single domain (i.e., ideal for navigation) structures in the giant magnetite particles depending on their size, morphology and spatial arrangement. Different giant magnetite crystal morphologies appear to have had different biological functions, including magnetotaxis and other non-navigational purposes. Our observations suggest that hyperthermals provided ideal conditions for giant magnetofossils, and that these organisms were globally distributed. Much more work is needed to understand the interplay between magnetofossil morphology, climate, nutrient availability, and environmental variability.

Interactive effects of ocean acidification and nitrogen limitation on the diatom Phaeodactylum tricornutum

Climate change is expected to bring about alterations in the marine physical and chemical environment that will induce changes in the concentration of dissolved CO2 and in nutrient availability. These in turn are expected to affect the physiological performance of phytoplankton. In order to learn how phytoplankton respond to the predicted scenario of increased CO2 and decreased nitrogen in the surface mixed layer, we investigated the diatom Phaeodactylum tricornutum as a model organism. The cells were cultured in both low CO2 (390 µatm) and high CO2 (1000 µatm) conditions at limiting (10 µmol/L) or enriched (110 µmol/L) nitrate concentrations. Our study shows that nitrogen limitation resulted in significant decreases in cell size, pigmentation, growth rate and effective quantum yield of Phaeodactylum tricornutum, but these parameters were not affected by enhanced dissolved CO2 and lowered pH. However, increased CO2 concentration induced higher rETRmax and higher dark respiration rates and decreased the CO2 or dissolved inorganic carbon (DIC) affinity for electron transfer (shown by higher values for K1/2 DIC or K1/2 CO2). Furthermore, the elemental stoichiometry (carbon to nitrogen ratio) was raised under high CO2 conditions in both nitrogen limited and nitrogen replete conditions, with the ratio in the high CO2 and low nitrate grown cells being higher by 45% compared to that in the low CO2 and nitrate replete grown ones. Our results suggest that while nitrogen limitation had a greater effect than ocean acidification, the combined effects of both factors could act synergistically to affect marine diatoms and related biogeochemical cycles in future oceans.

Diatom isotope data (d18Odiatom and d30Sidiatom) from IODP Site 303-U1304 (North Atlantic)

The Last Interglacial (LIG), corresponding to Marine Isotope Stage (MIS) 5e, provides a reference of interglacial climate variability in the absence of anthropogenic forcing. Using an expanded section of the LIG gained at Integrated Ocean Drilling Program Site U1304 in the Subarctic Atlantic, we demonstrate that the early MIS 5e was marked by oceanographic conditions conducive for high diatom production and accumulation. The appearance of diatom-dominated laminated oozes ~3 k.y. after the beginning of MIS 5e at ca. 125 ka coincides with a shift to higher d30Sidiat values together with the dominance of Thalassiothrix longissima, indicative of increased nutrient availability and silicic acid utilization in surface waters. Though the Subarctic Front provided the physical conditions for high diatom production and deposition, these processes alone are insufficient to explain the high rates of siliceous productivity and the formation of diatomaceous sediments. Instead, the additional presence of an increased nutrient pool provided by Subantarctic Mode Water played the decisive role in initiating and sustaining diatom production. The high diatom productivity and the occurrence of diatomaceous sediments in the late Quaternary challenge the current hypothesis of a silica-depleted North Atlantic during the LIG.

Dinoflagellate cysts in western equatorial Atlantic surface sediments

In contrast to the wide range of studies carried out in temperate and high-latitude oceanic regions, only a few studies have focused on recent and Holocene organic-walled dinoflagellate cyst assemblages from the tropics. This information is, however, essential for fully understanding the ability of species to adapt to different oceanographic regimes, and ultimately their potential application to local and regional palaeoenvironmental and palaeoceanographic reconstructions. Surface sediment samples of the western equatorial Atlantic Ocean north of Brazil, an area greatly influenced by Amazon River discharge waters, were therefore analysed in detail for their organic-walled dinoflagellate cyst content. A diverse association of 43 taxa was identified, and large differences in cyst distribution were observed. The cyst thanatocoenosis in bottom sediments reflects the seasonal advection of Amazon River discharge water through the Guyana Current and the North Equatorial Countercurrent well into the North Atlantic. To establish potential links between cyst distribution and the environmental conditions of the upper water column, distribution patterns were compared with mean temperature, salinity, density and stratification gradients within the upper water column (0-100 m) over different times of the year, using correspondence analysis and canonical correspondence analysis. The analyses show that differences in these parameters only play a subordinate role in determining species distribution. Instead, nutrient availability, or related factors, dominates the distribution pattern. The only possible indicators of slightly reduced salinities are Trinovantedinium applanatum and Lingulodinium machaerophorum. Four assemblage groups of cyst taxa with similar environmental affinities related to specific water masses/currents can be distinguished and have potential for palaeoenvironmental reconstruction.

Diatom oxygen and silicon isotopes data from IODP Hole 323-U1341B in the Bering Sea through the late Pliocene and over the onset of major Northern Hemisphere Glaciation

The Bering Sea is one of the most biologically productive regions in the marine system and plays a key role in regulating the flow of waters to the Arctic Ocean and into the subarctic North Pacific Ocean. Cores from Integrated Ocean Drilling Program (IODP) Expedition 323 to the Bering Sea provide the first opportunity to obtain reconstructions from the region that extend back to the Pliocene. Previous research at Bowers Ridge, south Bering Sea, has revealed stable levels of siliceous productivity over the onset of major Northern Hemisphere Glaciation (NHG) (circa 2.85-2.73 Ma). However, diatom silica isotope records of oxygen (d18Odiatom) and silicon (d30Sidiatom) presented here demonstrate that this interval was associated with a progressive increase in the supply of silicic acid to the region, superimposed on shift to a more dynamic environment characterized by colder temperatures and increased sea ice. This concluded at 2.58 Ma with a sharp increase in diatom productivity, further increases in photic zone nutrient availability and a permanent shift to colder sea surface conditions. These transitions are suggested to reflect a gradually more intense nutrient leakage from the subarctic northwest Pacific Ocean, with increases in productivity further aided by increased sea ice- and wind-driven mixing in the Bering Sea. In suggesting a linkage in biogeochemical cycling between the south Bering Sea and subarctic Northwest Pacific Ocean, mainly via the Kamchatka Strait, this work highlights the need to consider the interconnectivity of these two systems when future reconstructions are carried out in the region.

Coccolithophores in surface sediments of the Arabian Sea (TAble 1)

One hundred surface sediment samples of the Arabian Sea (Indian Ocean) were investigated and relative abundances of coccoliths were compared to mean annual gradients of temperature, salinity, chlorophyll, PO4 and mixed layer depth. Total coccolith concentrations ranged from 42*10**6/g sediment in coastal areas to more than 19000*10**6/g sediment in oceanic regions. The general distribution does not seem to be dependent on coccolithophore productivity in surface waters alone, but also on the diluting input of terrigenous material. A total of 27 taxa were identified. The main species dominating the assemblages were Gephyrocapsa oceanica, Emiliania huxleyi and Florisphaera profunda with a combined average abundance of more than 70%. Several species and species groups reflect with their distribution the environmental parameters of the overlying water masses and may be successfully used to improve palaeoclimatic reconstructions, e.g. (a) F. profunda exhibits a high similarity or even positive correlation to the mean annual mixed layer depth, (b) calciosolenids can be described as coastal or shelf species. While temperature and salinity gradients do not seem to be crucial for coccolithophores in this region, the mean mixed layer depth as well as the PO4 concentration (representative for total nutrient availability) may control in part the coccolithophore assemblages. According to the results of a cluster analysis and the distribution pattern of all species, it was possible to differentiate three main coccolithophore assemblages. A G. oceanica dominated assemblage mainly occurs in the northern part of the study area and can be described as 'high nutrient assemblage'. The second assemblage, dominated by F. profunda, may be typical for oligotrophic and stable conditions in open ocean waters. A third assemblage, with high amounts of 'coastal species', characterises coastal conditions on the shelves.