Skeletal trade-offs in coralline algae in response to ocean acidification

Ocean acidification is changing the marine environment, with potentially serious consequences for many organisms. Much of our understanding of ocean acidification effects comes from laboratory experiments, which demonstrate physiological responses over relatively short timescales. Observational studies and, more recently, experimental studies in natural systems suggest that ocean acidification will alter the structure of seaweed communities. Here, we provide a mechanistic understanding of altered competitive dynamics among a group of seaweeds, the crustose coralline algae (CCA). We compare CCA from historical experiments (1981-1997) with specimens from recent, identical experiments (2012) to describe morphological changes over this time period, which coincides with acidification of seawater in the Northeastern Pacific. Traditionally thick species decreased in thickness by a factor of 2.0-2.3, but did not experience a change in internal skeletal metrics. In contrast, traditionally thin species remained approximately the same thickness but reduced their total carbonate tissue by making thinner inter-filament cell walls. These changes represent alternative mechanisms for the reduction of calcium carbonate production in CCA and suggest energetic trade-offs related to the cost of building and maintaining a calcium carbonate skeleton as pH declines. Our classification of stress response by morphological type may be generalizable to CCA at other sites, as well as to other calcifying organisms with species-specific differences in morphological types.

(Table 3) Li concentrations and isotopic compositions in foraminiferal shells, carbonate sediments, and associated pore waters from ODP Sites 131-806 and 138-851

An improved procedure for lithium isotope analysis using Li3PO4 as the ion source has been investigated for application to geological samples. The 7Li/6Li ratio is measured using double rhenium filament thermal ionization mass spectrometry in which isotopic fractionation is minimized at high temperatures. The method produces a stable, high intensity Li+ ion beam that allows measurement of nanogram quantities of lithium. This results in a reduction in sample size of up to 1000 times relative to that required for the established Li2BO2+ method while maintaining a comparable precision of better than 1? (1 sigma). Replicate analyses of the NBS L-SVEC Li2CO3 standard yielded a mean value of 12.1047+/-0.0043 (n=21), which is close to the reported absolute value of 12.02+/-0.03. Intercalibration with a wide range of geological samples shows excellent agreement between the Li3PO4 and Li2BO2+ techniques. Replicate analyses of seawater and a fresh submarine basalt display high precision results that agree with previous measurements. Taking advantage of the high ionization efficiency of the phosphate ion source, we have made the first measurements of the lithium concentration (by isotope dilution) and isotopic composition of calcareous foraminiferal tests and other marine carbonates. Preliminary results indicate that substantial lithium exchange occurs between carbonate sediments and their interstitial waters. In addition, a possible link between lithium paleoceanography and paleoclimate during the last 1000 ky may be derived from planktonic foraminiferal tests. This highly sensitive technique can be applied in the examination of low lithium reservoirs and thereby provide insight into some fundamental aspects of lithium geochemistry.

Geochemistry of sediments from the upwelling off Morocco

The high-productive upwelling area off Morocco is part of one of the four major trade-wind driven continental margin upwelling zones in the world oceans. While coastal upwelling occurs mostly on the shelf, biogenic particles derived from upwelling are deposited mostly at the upper continental slope. Nutrient-rich coastal water is transported within the Cape Ghir filament region at 30°N up to several hundreds of kilometers offshore. Both upwelling intensity and filament activity are dependent on the strength of the summer Trades. This study is aimed to reconstruct changes in trade wind intensity over the last 250,000 years by the analysis of the productivity signal contained in the sedimentary biogenic particles of the continental slope and beneath the Cape Ghir filament. Detailed geochemical and geophysical analyses (TOC, carbonate, C/N, delta13Corg, delta15N, delta13C of benthic foraminifera, delta18O of benthic and planktic foraminifera, magnetic susceptibility) have been carried out at two sites on the upper continental slope and one site located further offshore influenced by the Cape Ghir filament. A second offshore site south of the filament was analyzed (TOC, magnetic susceptibility) to distinguish the productivity signal related to the filament signal from the general offshore variability. Higher productivity during glacial times was observed at all four sites. However, the variability of productivity during glacial times was remarkably different at the filament-influenced site compared to the upwelling-influenced continental slope sites. In addition to climate-related changes in upwelling intensity, zonal shifts of the upwelling area due to sea-level changes have impacted the sedimentary productivity record, especially at the continental slope sites. By comparison with other proxies related to the strength and direction of the prevailing winds (Si/Al ratio as grain-size indicator, pollen) the productivity record at the filament-influenced site reflects mainly changes in trade-wind intensity. Our reconstruction reveals that especially during glacial times trade-wind intensity was increased and showed a strong variability with frequencies related to precession.

Compilation of 220 sediment core d13C data from the glacial Atlantic Ocean

We compare a compilation of 220 sediment core d13C data from the glacial Atlantic Ocean with three-dimensional ocean circulation simulations including a marine carbon cycle model. The carbon cycle model employs circulation fields which were derived from previous climate simulations. All sediment data have been thoroughly quality controlled, focusing on epibenthic foraminiferal species (such as Cibicidoides wuellerstorfi or Planulina ariminensis) to improve the comparability of model and sediment core carbon isotopes. The model captures the general d13C pattern indicated by present-day water column data and Late Holocene sediment cores but underestimates intermediate and deep water values in the South Atlantic. The best agreement with glacial reconstructions is obtained for a model scenario with an altered freshwater balance in the Southern Ocean that mimics enhanced northward sea ice export and melting away from the zone of sea ice production. This results in a shoaled and weakened North Atlantic Deep Water flow and intensified Antarctic Bottom Water export, hence confirming previous reconstructions from paleoproxy records. Moreover, the modeled abyssal ocean is very cold and very saline, which is in line with other proxy data evidence.

Siliceous phytoplankton fluxes off northwest Africa

Four years of observations (1988-1991) of downward fluxes of diatoms and silicoflagellates at a trap site off Cape Blanc (ca. 20°N, 20°W), northwest Africa, are presented. Significant variations in flux and species composition were observed as well as a marked drop in the export of biogenic opal (and diatoms) from 1988 to 1989; fluxes remained low thereafter. We hypothesize that this diminution might be related to decreased coastal upwelling intensity and offshore spreading of the typical chlorophyll filament, and/or a lesser silicate content of upwelling waters off Cape Blanc. In addition, the more seaward positioning of the mooring may have influenced the fluxes. At all times, diatoms were the most prominent contributors to the biogenic opal flux, and diatom fluxes closely paralleled total mass flux fluctuations. Although species composition varied seasonally, no significant qualitative variations were observed from year to year. In general, the dominance of neritic diatoms, such as Thalassionema nitzschioides var. nitzschioides, resting spores of Chaetoceros and Cyclotella litoralis, reflected the continuous offshore influence of coastal upwelling at the Cape Blanc trap site, with stronger intensity in spring/summer. In contrast, the occurrence of pelagic diatoms (e.g. Nitzschia bicapitata, N. interruptestriata, T. nitzschioides var. parva and Fragilariopsis doliolus), and high silicoflagellate fluxes (mainly Dictyocha messanensis) were linked to inshore transport of oceanic waters, generally in winter. With the exception of some fragile, pelagic diatoms, dominant species found in the settled material also occurred in the underlying sediments, suggesting that diatom thanatocoenosis downcore (Organisms preserved from the top to the bottom in sediment core) can be used as a reliable indicator of the intensity and persistence of the offshore spreading of coastal upwelling.

Seawater carbonate chemistry and structural integrity of the coralline algae Lithothamnion glaciale in a laboratory experiment

The uptake of anthropogenic emission of carbon dioxide is resulting in a lowering of the carbonate saturation state and a drop in ocean pH. Understanding how marine calcifying organisms such as coralline algae may acclimatize to ocean acidification is important to understand their survival over the coming century. We present the first long-term perturbation experiment on the cold-water coralline algae, which are important marine calcifiers in the benthic ecosystems particularly at the higher latitudes. Lithothamnion glaciale, after three months incubation, continued to calcify even in undersaturated conditions with a significant trend towards lower growth rates with increasing pCO2. However, the major changes in the ultra-structure occur by 589 µatm (i.e. in saturated waters). Finite element models of the algae grown at these heightened levels show an increase in the total strain energy of nearly an order of magnitude and an uneven distribution of the stress inside the skeleton when subjected to similar loads as algae grown at ambient levels. This weakening of the structure is likely to reduce the ability of the alga to resist boring by predators and wave energy with severe consequences to the benthic community structure in the immediate future (50 years).

Planktic foraminifera analyses of surface sediments and satellite data in the Canary Islands region

The Canary Islands region occupies a key position with respect to biogeochemical cycles, with the zonal transition from oligotrophic to nutrient-rich waters and the contribution of Saharan dust to the particle flux. We present the distribution of geochemical proxies (TOC, carbonate, d15N, d13Corg, C/N-ratio) and micropaleontological parameters (diatoms, dinoflagellates, foraminifera, pteropods), in 80 surface-sediment samples in order to characterise the influence of coastally upwelled water on the domain of the subtropical gyre. Results of the surface-sediment analyses confirmed the high biomass gradient from the coast to the open ocean inferred from satellite data of surface chlorophyll or SST. The distribution of total dinoflagellate cysts, the planktic foraminifera species Globigerina bulloides, the diatom resting spore Chaetoceros spp., and TOC concentration coincided well with the areas of strong filament production off Cape Ghir and Cape Yubi. The warm-water planktic foraminifera Globigerinoides ruber (white), the diatom Nitzschia spp., and the d15N-values showed the opposite trend with high values in the open ocean. Factor analyses on the planktic foraminifera species distribution indicated three major assemblages in the Canary Islands region that represent the present surface-water conditions from the upwelling influenced region via a mixing area towards the subtropical gyre.

Distribution of coccoliths in surface sediments of the south-eastern South Atlantic Ocean

Recent coccoliths from 52 surface sediment samples recovered from the south-eastern South Atlantic were examined qualitatively and quantitatively in order to assess the controlling mechanisms for their distribution patterns, such as ecological and preservational factors, and their role as carbonate producers. Total coccolith abundances range from 0.2 to 39.9 coccoliths*10**9/ g sediment. Four assemblages can be delineated by their coccolith content characterising the northern Benguela, the middle to southern Benguela, the Walvis Ridge and the deeper water. Distinctions are based on multivariate ordination techniques applied on the relative abundances of the most abundant taxa, Emiliania huxleyi, Calcidiscus leptoporus, Gephyrocapsa spp., Coccolithus pelagicus and subtropical to tropical species. The coccolith distribution seems to be temperature and nutrient controlled co-varying with the seaward extension of the upwelling filament zone in the Benguela. A preservation index (CEX') based on the differential dissolution behaviour of the delicate E. huxleyi and Gephyrocapsa ericsonii versus the robust C. leptoporus is applied in order to detect the position of the coccolith lysocline. Although some samples were recognised as dissolution-affected, the distribution of the coccoliths in the surface-sediments reflects the different oceanographic surface-water conditions. Mass estimations of the coccolith carbonate reveal coccoliths to be only minor contributors to the carbonate preserved in the surface sediments. The mean computed coccolith carbonate content is 17 wt.%, equivalent to a mean contribution of 23% to the bulk carbonate.

(Table 3) Interannual variability of total mass flux at the deep sediment trap site CBmeso between 1988 and 2006

This article will review major features of the 'giant' Cape Blanc filament off Mauritania with regard to the transport of chlorophyll and organic carbon from the shelf to the open ocean. Within the filament, chlorophyll is transported about 400 km offshore. Modelled particle distributions along a zonal transect at 21°N showed that particles with a sinking velocity of 5 m d**-1 are advected offshore by up to 600 km in subsurface particle clouds generally located between 400 m and 800 m water depth, forming an Intermediate Nepheloid Layer (INL). It corresponds to the depth of the oxygen minimum zone. Heavier particles with a sinking velocity of 30 m d**-1 are transported from the shelf within the Bottom Layer (BL) of more than 1000 m thickness, largely following the topography of the bottom slope. The particles advected within the BL contribute to the enhanced winter-spring mass fluxes collected at the open-ocean mesotrophic sediment trap site CB-13 (200 nm offshore), due to a long distance advection in deeper waters. The lateral contribution to the deep sediment trap in winter-spring is estimated to be 63% and 72% for organic carbon and total mass, respectively, whereas the lateral input for both components on an annual basis is estimated to be in the order of 15%. Biogenic opal increases almost fivefold from the upper to the lower mesotrophic CB-13 trap, also pointing to an additional source for biogenic silica from eutrophic coastal waters. Blooms obviously sink in smaller, probably mesoscale-sized patches with variable settling rates, depending on the type of aggregated particles and their ballast content. Generally, particle sinking rates are exceptionally high off NW Africa. Very high chlorophyll values and a large size of the Cape Blanc filament in 1998-1999 are also documented in enhanced total mass and organic carbon fluxes. An increasing trend in satellite chlorophyll concentrations and the size of the Cape Blanc filament between 1997 and 2008 as observed for other coastal upwelling areas is not documented.