Coccolith Sr/Ca ratios in the eastern Mediterranean: Production versus export processes

Samples collected by two sediment traps located southwest of Crete in the eastern Mediterranean (EMED) [48A (1953 m) and 48B (950 m)] from June 2005 to May 2006 were used to study fluxes of organic carbon, carbonate and coccolithophores in combination with the variations of Sr/Ca ratios in different individually picked coccolith species. Considering the complexity of the EMED, we validate the use of Sr/Ca ratios as productivity proxy and unravel the varied processes which may influence it. We examined the relationship between the seasonal peaks in export fluxes and the Sr/Ca ratio in coccoliths of three upper photic zone coccolithophores species collected in the traps, Calcidiscus leptoporus, Helicosphaera carteri and Emiliania huxleyi. We aimed at testing whether high export fluxes are correlated with high Sr/Ca ratios, suggestive of higher nutrient-stimulated production, or Sr/Ca ratios are unchanged during high export periods, suggestive of increased export efficiency or scavenging. Periods of enhanced trap fluxes in March and June result from surface water blooms recognized in satellite imagery. An additional peak flux was found in January, but this peak represents re-suspended or recycled material in the water column.The amplitude of seasonal variations in the Sr/Ca ratios of the three investigated species is small in both traps. In the shallow trap, a decrease in the Sr/Ca ratio of C. leptoporus occurred synchronously with minimal fluxes. The other two species were not measured for this period. In the deep trap, no such decrease in Sr/Ca was observed during minimal fluxes, in either C. leptoporus or H. carteri, probably due to a long residence of coccoliths in the water column, recycling and low export efficiency. Absolute Sr/Ca ratios for all species are lower than in other more productive environments like the Bay of Bengal, Arabian Sea, or Sargasso Sea. We conclude that Sr/Ca ratios in coccoliths of surface sediments in the EMED reflect mainly spring–summer bloom conditions averaged over hundreds to thousands of years.In addition, the origin of varying calcite thickness in H. carteri was investigated. The similarity of average Sr/Ca ratios in differently-calcified specimens confirms that coccolith thickness variations in this species result from primary biomineralization processes and not from variable overgrowth by (low Sr) abiogenic calcite in the water column or the sediments.     

Strontium/Calcium ratio, carbon and oxygen stable isotopes in coccolith carbonate from different grain-size fractions in South Atlantic surface sediments

In this study, the Strontium/Calcium (Sr/Ca) ratio, and the carbon and oxygen isotopic compositions of coccoliths are investigated in three different grain-size fractions (<20 μm, 15-5 μm, <5 μm) of 17 surface sediment samples from the Equatorial and South Atlantic. The results are compared to environmental parameters in order to assess the factors controlling the observed coccolith geochemical patterns. Isotopic and geochemical composition of coccolith species in surface sediment samples from the South Atlantic greatly varies according to the different grain-size fractions. However, even if the absolute values show a great offset, the general trends are comparable. The δ¹⁸O values show a decreasing trend with increasing temperature. The δ¹³C and Sr/Ca ratio are mainly influenced by productivity of coccolithophores, which is in turn controlled by different factors, such as temperature, nutrient supply and productivity of other phytoplankton groups. Dilution and dissolution are negligible factors in these open marine samples. Therefore, coccolith abundance in bulk sediment is the best approximation for productivity of coccolithophores. The various coccolith species fractionate Sr differently, as is best shown by the 5-15 μm fraction where three species (Calcidiscus leptoporus, Helicosphaera carteri and Coccolithus pelagicus) predominantly occur.     

Paleoceanographic interpretations of coccoliths and oxygen-isotopes from the sediment surface of the southwest Indian Ocean

The distribution of forty-four coccolithophore species in one hundred deep-sea core-tops from the southwest Indian Ocean is described. Three coccolith assemblages have been recognised (Maputo, Agulhas Current and deep water) by the relative abundances of four ecologically significant coccolithophore species (Gephyrocapsa oceanica, Emiliania huxleyi, Calcidiscus leptoporus and Umbilicosphaera sibogae). Their biogeographical distribution appears to be related to water temperature, nutrient concentration and dissolution.The degree of preservation of coccoliths and foraminifera indicates that the carbonate lysocline lies somewhere between 3500 and 4000 m, resulting in the concentration of dissolution-resistant microfossils below this depth.Stable oxygen isotope ratios of the planktonic foraminiferal species Globigerinoides sacculifer range between −1.5 to −1.0‰ PDB (equal to 22.8–25.1°C) and occur in a narrow band on the sea floor beneath the “A” route of the Agulhas Current.These values are about 0.5 per mil PDB lighter than samples analyzed on either side of this band and can be explained by the Agulhas Current's elevated temperature at the ocean surface of 2–3°C. Thus an oxygen isotope imprint of the Agulhas Current exists beneath it on the sea floor.The Agulhas Current is probably the major factor influencing sedimentation, sediment-distribution patterns and geological features in the study area. At present it is voluminous and fast flowing, possibly eroding sediments up to 2500 m below the surface.The oxygen-isotope ratios and nannoplankton counts obtained in this study indicate, however, that the majority of samples are most probably recent or at least not older than 85,000 years. This implies that sediments are accumulating on the ocean floor and that the Agulhas Current does not have a pronounced erosional influence, at least in areas from which cores were retrieved for this study.     

Vertical and lateral variations in coccolithophore community structure across the subtropical frontal zone in the South Atlantic Ocean

In this study the coccolithophore compositions of 11 plankton depth stations along a N–S transect from the South Atlantic Subtropical Gyre to the Subantarctic Zone were examined qualitatively and quantitatively. The lateral and vertical distribution patterns of not only the most abundant taxa but also of the morphotypes of distinct species complexes, such as Calcidiscus leptoporus, Emiliania huxleyi, and Umbellosphaera tenuis were the focus. Geographic variation among morphotypes mirrors different ecological affinities of the members of a species complex. Multivariate statistics were used to infer the relationship between a set of known environmental data and species concentrations. The results of the detrended Canonical Correspondence Analysis (CCA) revealed the presence of distinct species assemblages. The Subtropical Gyre assemblage within the upper 50 m of the photic zone is mainly composed of Umbellosphaera irregularis, U. tenuis types III and IV, Discosphaera tubifera, Rhabdosphaera clavigera, S. pulchra and E. huxleyi var. corona, adapted to warm and oligotrophic conditions. In the deeper photic zone abundant Florisphaera profunda, Gephyrocapsa ericsonii and Oolithotus spp. are encountered, benefiting from higher nutrient concentrations in the vicinity of the nutricline. A well-defined Subtropical Frontal Zone (STFZ) association is clearly dominated by E. huxleyi types A and C throughout the upper 100 m of the water column. Secondary contributors in the upper photic zone are Syracosphaera spp. (mainly S. histrica, S. molischii), Michaelsarsia elegans, Ophiaster spp. and U. tenuis type II. This assemblage is associated with cooler, nutrient-rich waters. E. huxleyi type B is found deeper in the water column. Here it is accompanied by Algirosphaera robusta, G. muellerae, and S. anthos indicating a tolerance of lower light availability in environments with elevated productivity. C. leptoporus spp. leptoporus shows relatively high cell numbers in all sampled water levels throughout the STFZ. Interestingly, its coccoliths are often smaller 5 μm in lith diameter. The mean coccolithophore assemblages of a station were compared to the underlying surface sediment assemblages. For the most part, the distribution of the morphotypes is reflected in the sedimentary archive, thus proving their potential as paleoecological proxies.     

Geographical and seasonal differences in morphology and dynamics of the coccolithophore Calcidiscus leptoporus

Calcidiscus leptoporus is a cosmopolitan coccolithophore species, composed of three morphotypes characterised by differences in morphology and size. The seasonal dynamics of this species have been studied in four sediment trap and plankton time series covering different biogeographic settings. Investigated parameters were: variations in absolute and relative abundances of the three morphotypes, average size variations of the C. leptoporus assemblages, and intra-morphotype size variations. For each time series, and for the complete data set, the relationship between C. leptoporus dynamics and some environmental parameters was investigated. Seasonal variations can be recognised in the four time series, but the biogeographic pattern governing this seasonality is complex. The two best documented morphotypes show comparable seasonal fluctuations in absolute abundances, mainly co-varying with temperature and related factors such as water mixing. However, their relative abundances change with varying nutrient contents in the surface water masses, suggesting slight differences in their respective ecological preferences. Average size of the C. leptoporus assemblage provides information about the morphotype composition and allows a comparison of the seasonal variations observed in the present study and Holocene geographic patterns reported in the literature. Additionally, intra-morphotype size variations might be related to environmental changes.     

Movement of oceanic fronts south of Australia during the last 10 ka: interpretation of calcareous nannoplankton in surface sediments from the Southern Ocean

Preservation of calcareous nannoplankton in surface sediment samples from the Southern Ocean south of Australia and adjacent to New Zealand record a single assemblage. The dominant species are Emiliania huxleyi, Gephyrocapsa muellerae, Calcidiscus leptoporus, Helicosphaera carteri and Coccolithus pelagicus. The assemblage varies little in abundance and diversity with minor correlation to present-day overlying surface water masses and oceanic fronts. Increase in abundance of H. carteri and C. pelagicus in the region of the Subtropical Front may reflect higher nutrients associated with this front. The assemblage, although altered by dissolution, represents a warmer climatic interval than present-day with the presence of preferentially dissolved, warm-water species preserved as far south as the Polar Front. The presence of warm-water species under sub-Antarctic waters at the Polar Front is interpreted as a relic population from the Holocene climatic optimum of 10–8 ka. The absence of coccoliths in sediments poleward of the Polar Front suggests an equatorward shift of this front following the climatic optimum, resulting in increased productivity of siliceous phytoplankton associated with the colder waters and increased dissolution of coccoliths. Movement of the Subtropical Front for the same interval is not recorded in the preserved coccoliths. The more heavily calcified form of E. huxleyi which dominates the living assemblage north of the Subtropical Front is subject to dissolution in this region and is poorly preserved in the sediment assemblage.     

Miocene variability of Calcidiscus gr. leptoporus and possible evolutionary relationship with another Coccolithaceae: Umbilicosphaera gr. sibogae.

Bibliographic data concerning the variability of coccosphere and coccoliths during the life-cycle of two extant Coccolithophorid species (Calcidiscus gr. leptoporus and Umbilicosphaera gr. sibogae) are summarized and completed by new observations on the structure of the proximal shield by fossil coccoliths. Thorough investigation of the proximal shield of C. gr. leptoporus in the latest Miocene (Messinian) assemblages from three oceanic localities (central Pacific, eastern equatorial Atlantic and southwestern Indian oceans) indicates the co-occurrence of two types of coccoliths: (1) typical C. gr. leptoporus and (2) specimens constituted by a distal shield similar to that of C. leptoporus, but with a proximal shield comparable to that of U. sibogae, although no typical U. sibogae occurs on the basis of the distal views. Such intermediate coccoliths have been previously described from the Badenian (middle Miocene) of central Europe. Whereas the authors considered them as a new species (Cycloperfolithus carlae), they are interpreted here as transitional morphotypes revealing an evolutionary link between C. gr. leptoporus and U. gr. sibogae, during the late Miocene. Consequently, the palaeontological 'species' carlae is hereby transferred to the genus Calcidiscus. These observations also point out the possibility of a diachronous evolution of the two shields of the same coccolith type, underlining the necessity for a better knowledge of the proximal shield structure, usually somewhat neglected.     

A GENETIC MARKER TO SEPARATE EMILIANIA HUXLEYI (PRYMNESIOPHYCEAE) MORPHOTYPES1

Emiliania huxleyi (Lohm.) Hay and Mohler is a ubiquitous unicellular marine alga surrounded by an elaborate covering of calcite platelets called coccoliths. It is an important primary producer involved in oceanic biogeochemistry and climate regulation. Currently, E. huxleyi is separated into five morphotypes based on morphometric, physiological, biochemical, and immunological differences. However, a genetic marker has yet to be found to characterize these morphotypes. With the use of sequence analysis and denaturing gradient gel electrophoresis, we discovered a genetic marker that correlates significantly with the separation of the most widely recognized A and B morphotypes. Furthermore, we reveal that the A morphotype is composed of a number of distinct genotypes. This marker lies within the 3′ untranslated region of a coccolith associated protein mRNA, which is implicated in regulating coccolith calcification. Consequently, we tentatively termed this marker the coccolith morphology motif.     

Response of calcareous nannofossil assemblages to paleoenvironmental changes through the mid-Pleistocene revolution at Site 1090 (Southern Ocean)

Quantitative study on calcareous nannofossil assemblages has been performed in high time resolution (2–3 kyr) at the Ocean Drilling Program Site 1090. The location of this site in the Southern Ocean is crucial for the comprehension of thermohaline circulation and frontal boundary dynamics, and for testing the employ of nannoflora as paleoceanographical tool. The chronologically well constrained investigated record spans between Marine Isotope Stage (MIS) 35 and 15, through an interval of global paleoclimate and paleoceanographical modification also known as mid-Pleistocene revolution (MPR). Measures of ecological (Shannon–Weaver diversity and paleoproductivity) and dissolution indices together with spectral and wavelet analyses carried out on the acquired time series provide valuable information for interpretation of data in terms of paleoecology and paleoceanography. Assemblages are mainly represented by dominant small Gephyrocapsa, common Calcidiscus leptoporus s.l., Coccolithus pelagicus s.l., Gephyrocapsa (4-5.5 μm), the extinct Pseudoemiliania lacunosa and Reticulofenestra spp. (R. asanoi and Reticulofenestra sp.). Morphotypes discriminated within Calcidiscus leptoporus s.l. and Coccolithus pelagicus s.l., reveal that they may have had different ecological preferences during Pleistocene with respect to the present. The composition and fluctuation in nannofossil assemblage and their comparison with the available Sea Surface Temperature (SST) and C-org curves suggest a primary ecological response to paleoenvironmental changes; relationships to different surface water features and boundary dynamics, as well as to different efficiencies and motions of the intermediate and deep water masses have been inferred. A more northward position of Subantarctic Front (SAF) during most of the Early Pleistocene record has been highlighted based on assemblage composition characterised by common Calcidiscus leptoporus s.l., Coccolithus pelagicus s.l., medium Gephyrocapsa (4–5.5 μm), and by the rarity or absence of Umbilicosphaera spp., Rhabdosphaera spp., Pontosphaera spp., Oolithotus fragilis. Exceptions are the more intense interglacials MIS 31, 17, and probably MIS 15, when a southward displacement of frontal system occurred, coincident with peaks in abundance of Helicosphaera spp. and Syracosphaera spp. Higher nutrient content and more dynamic conditions occurred between MIS 32 and MIS 25, in relation to shallower location of nutrient-rich Antarctic Intermediate Water (AAIW) core and to reduction of glacial–interglacial variability. A nannofossil barren interval is coincident with the known stagnation of South Atlantic deep water circulation during MIS 24, when North Atlantic Deep Water (NADW) was reduced or suppressed and an enhanced northward deep penetration of the more corrosive Circumpolar Deep Water (CPDW) took place. An event of strong instability in nutricline dynamics characterised the transition MIS 23–22 as suggested by sharp fluctuations in paleoproductivity proxies, linked to major changes in oceanographic circulation and to the first distinct increase of larger ice volumes at this time. From MIS 21 upward the nannofossil variations seem to be primarily controlled by glacial–interglacial cyclicity and temperature fluctuations. The cyclic fluctuation recognised in nannofossil abundance seems to be linked to orbitally-forced climatic variation, primarily to the obliquity periodicity recorded in the patterns of C. leptoporus intermediate (5–8 μm) and C. pelagicus pelagicus (6–10 μm); however no obvious and linear relations may be always observed between nannoflora fluctuation and Milankovitch parameters, suggesting more complex and unclear relationships between nannofossils and environmental change.     

The Effect of cytoskeleton inhibitors on coccolith morphology in Coccolithus braarudii and Scyphosphaera apsteinii

The calcite platelets of coccolithophores (Haptophyta), the coccoliths, are among the most elaborate biomineral structures. How these unicellular algae accomplish the complex morphogenesis of coccoliths is still largely unknown. It has long been proposed that the cytoskeleton plays a central role in shaping the growing coccoliths. Previous studies have indicated that disruption of the microtubule network led to defects in coccolith morphogenesis in Emiliania huxleyi and Coccolithus braarudii. Disruption of the actin network also led to defects in coccolith morphology in E. huxleyi, but its impact on coccolith morphology in C. braarudii was unclear, as coccolith secretion was largely inhibited under the conditions used. A more detailed examination of the role of actin and microtubule networks is therefore required to address the wider role of the cytoskeleton in coccolith morphogenesis. In this study, we have examined coccolith morphology in C. braarudii and Scyphosphaera apsteinii following treatment with the microtubule inhibitors vinblastine and colchicine (S. apsteinii only) and the actin inhibitor cytochalasin B. We found that all cytoskeleton inhibitors induced coccolith malformations, strongly suggesting that both microtubules and actin filaments are instrumental in morphogenesis. By demonstrating the requirement for the microtubule and actin networks in coccolith morphogenesis in diverse species, our results suggest that both of these cytoskeletal elements are likely to play conserved roles in defining coccolith morphology.     

Further evidence for the heterococcolith–holococcolith combination Calcidiscus leptoporus–Crystallolithus rigidus

New findings of combination coccospheres bearing heterococcoliths of Calcidiscus leptoporus and holococcoliths of Crystallolithus rigidus are documented. These findings confirm previous suggestions that these two “species” are separate phases of the life-cycle of a single species.     

Extant calcareous nannoplankton in the Australian Sector of the Southern Ocean (austral summers 1994 and 1995)

Living calcareous nannoplankton in the region between Australia and Antarctica are distributed in five assemblages associated with distinct physico-chemical properties of surface and subsurface water masses. Temperature and salinity ranges for living assemblages were 2–15.7°C and 33.7–35.56‰, respectively, with maximum cell densities for austral summer 1994 found at 9.63°C and 34.44‰, and for austral summer 1995 at 12.8°C and 35.17‰. Nutrients (phosphate, silicate and nitrate) increase poleward and vertically from surface to depth. Abundance and diversity of calcareous nannoplankton decrease in a poleward direction with major shifts located across both the Subtropical and the Subantarctic Fronts. Higher cell densities were found below 50 m equatorward of the Subtropical Front and above 50 m poleward of this front. Poleward of the Antarctic Divergence coccolithophores are absent from all samples. Three different morphotypes of Emiliania huxleyi were identified, one of which has a distribution associated with the Subtropical Front. Of the subordinate species Syracosphaera spp, Calciosolenia murrayi and Umbellosphaera tenuis dominate equatorward of the Subtropical Front with Syracosphaera spp and Calcidiscus leptoporus dominant poleward of this front. A peculiar community of weakly calcified species is recorded for the first time outside the Weddell Sea.     

Late Neogene nannofossil biostratigraphy and paleoceanography of the northeastern Gulf of Mexico and adjacent areas

Two 305 m cored sections from the northwest Florida continental shelf contain a nearly complete record of late Neogene hemipelagic sedimentation. One of the sites, south and east of De Soto Canyon, is isolated from terrigenous sediment except for sediment transported in suspension. This site contains a continuous record from the late Miocene to the Recent. The second site, on the western rim of De Soto Canyon, is more expanded and continuous from the late Pliocene to the Recent. A hiatus separates the late Pliocene from the middle Miocene. Six prominent nannofossil biohorizons were recognized within the Pleistocene, seven within the Pliocene, and three within the Miocene; in addition one biohorizon marks the base of the Pleistocene and another the base of the Pliocene.Nearly all carbonate in the sediment is pelagic. Terrigenous detrital sedimentation was controlled by glacioeustatic sea level fluctuations during the Pleistocene, and sea level changes are probably responsible for fluctuations in the ratio of pelagic carbonate to clayey detritus in pre-Pleistocene sediments also. Carbonate content, coarse fraction percent, and relative abundances of environmentally sensitive nannoplankton species suggest important paleoceanographic changes in the northeastern Gulf of Mexico and adjacent areas. Fluctuations in the relative abundance of the solution-resistant coccoliths of the genusCyclococcolithus indicate that waters at a depth of 600–1000 m were more corrosive during the late Miocene than they are today. The decrease in carbonate dissolution during the late Miocene probably was a response to gradual constriction of the Central American passage and the consequent restriction of flow of corrosive water from the Pacific. Short term fluctuations in dissolution during the Pliocene and Pleistocene are related to climatic cycles.Productivity variations in the surface waters, recorded mainly by the relative abundance of small and large morphotypes of closely related coccolith species, indicate that productivity increased during the Pliocene, but the most dramatic change — a major oceanwide increase in productivity — occurred during the Pleistocene, during and just prior to the Jaramillo magnetic event about 0.9 m.y. ago. Surprisingly the late Miocene Messinian event did not leave a significant imprint in the northeastern Gulf of Mexico.     

Calcareous nannoplankton in surface sediments of the East China Sea

Twenty-eight species of coccoliths were identified in surface sediments of the East China Sea. The species composition of the coccolith assemblages is similar to that of the North Pacific Water Mass, but differs from the latter by a greater dominance ofEmiliania huxleyi andGephyrocapsa oceanica (together making up over 90% specimens). The coccolith species composition shows considerable differences between the continental shelf and the Okinawa Trough areas, reflecting different water masses. On the other hand, the coccolith content in sediments is apparently controlled largely by the deposition rate of terrigenous material.     

Coccolithophores in the equatorial Atlantic Ocean: response to seasonal and Late Quaternary surface water variability

The present study was initiated to ascertain the significance of coccolithophores as a proxy for paleoceanographic and paleoproductivity studies in the equatorial Atlantic. Data from a range of different samples, from the plankton, surface sediments as well as sediment cores are shown and compared with each other.In general, the living coccolithophores in the surface and subsurface waters show considerable variation in cell numbers and distribution patterns. Cell densities reached a maximum of up to 300×10³ coccospheres/l in the upwelling area of the equatorial Atlantic. Here, Emiliania huxleyi is the dominant species with relatively high cell numbers, whereas Umbellosphaera irregularis and Umbellosphaera tenuis are characteristic for oligotrophic surface waters. Although they are observed in high relative abundances, these species only occur in low absolute numbers. The lower photic zone is dominated by high abundances and considerable cell numbers of Florisphaera profunda.The geographical distribution pattern of coccoliths in surface sediments reflects the conditions of the overlying surface water masses. However, abundances of the oligotrophic species Umbellosphaera irregularis and Umbellosphaera tenuis are strongly diminished, causing an increase in relative abundance of the lower photic zone taxa Florisphaera profunda and Gladiolithus flabellatus.During the past 140,000 years the surface water circulation of the equatorial Atlantic has changed drastically, as can be seen from changes in the coccolithophore species composition, absolute coccolith numbers, as well as coccolith accumulation rates. Significant increases in coccolith numbers and accumulation rates is observed in the southern equatorial Atlantic during the last glacial interval (oxygen isotope stages 2–4), which we attribute to enhanced upwelling intensities and advection of cool nutrient rich waters at this site. In the western equatorial Atlantic we observe an opposite trend with decreasing numbers of coccoliths during glacial periods, which probably is caused by a deepening of the thermocline.     

Light-dependent coccolith formation in the two forms of Coccolithus pelagicus

Summary Two methods were employed for measuring coccolith formation and photosynthesis in coccolithophorids. The first method was based on measurements of ¹⁴C radioactivity of cells on membrane filters before and after acid treatment. The second method involved a conversion of ¹⁴C in coccoliths or whole cells to BaCO₃ prior to counting. It was observed that in determinations of photosynthetic (or total) ¹⁴C by the first method, the count rate produced by a given amount of the isotope was 30–40% lower in the non-motile and motile forms of Coccolithus pelagicus than in C. huxleyi. There was no similarly great discrepancy in determinations of coccolith ¹⁴C.Light-dependent coccolith formation was demonstrated in both forms of C. pelagicus. The non-motile form may deposit several times more carbon in its coccoliths than it assimilates photosynthetically. In the motile form, coccolith carbon amounts to less than 2% of photosynthetic carbon.     

Evolution of the coccolith genus Lotharingius during the Late Pliensbachian-Early Toarcian interval in Asturias (N Spain). Consequences of the Early Toarcian environmental perturbations

The evolution of the coccolith genus Lotharingius was investigated in 18 samples from the West Rodiles section (Asturias, Northern Spain), Upper Pliensbachian-Lower Toarcian in age. In each sample, the length and width of the coccolith and of its central area were measured on 150 specimens of Lotharingius. A total of 2700 specimens were measured, and 10,800 measurements were obtained. Based on the results of the morphometric analysis, the stratigraphic ranges, the abundance patterns and the light microscope observations, the aims of this work are to determine whether the specimens analysed correspond to different species, to distinct ecophenotypes or to an anagenetic lineage of morphotypes of a single species with overlapping stratigraphic ranges, and to test the influence of the paleoenvironmental perturbations during the Early Toarcian on the studied genus. Three species of Lotharingius were distinguished: L. hauffii, L. sigillatus and L. crucicentralis. Based on central area size and structure, two morphotypes (A and B) of L. hauffii were differentiated. All the parameters measured show a great overlap between the morphotypes and species. An increase in both coccolith and central area size and a development of more complex central area structures were observed through the studied section. L. hauffii is the smallest form, followed by L. sigillatus, and finally L. crucicentralis is the largest species and has the most complex central area structure. A clear decrease in size or “dwarfing” of the species was observed in the samples corresponding to the Early Toarcian environmental perturbations. L. crucicentralis was not identified in these samples. We hypothesize that the changes in size and abundance of the Lotharingius species could be related to the unfavourable palaeoenvironmental conditions for the biomineralization of their coccoliths.     

ROLE OF THE LIGHT-DARK CYCLE AND MEDIUM COMPOSITION ON THE PRODUCTION OF COCCOLITHS BY EMILIANIA HUXLEYI (HAPTOPHYCEAE)1

Production of coccoliths by cells of Emiliania huxleyi (Lohmann) Hay and Mohler was measured during exposure of the cells to two diel light-dark cycles (16:8 h). During the light period about eight coccoliths per cell were formed at a constant rate of one coccolith per 2 h. Cells divided during the first half of the dark period. No coccolith production took place during the dark period. With electron microscopy we found early-stage, coccolith-production compartments in cells after mitosis while still in the dark. No calcification was observed in these compartments. Cells grown on enriched seawater (Eppley's medium) tended to produce enough coccoliths to cover the cell in a single layer. When these cells reached the stationary phase coccolith production stopped. Coccolith production was induced by removal of extracellular coccoliths. Cells grown on medium containing 2% of the nitrate and phosphate of Eppley's medium tended to produce coccoliths in the stationary phase. This resulted in the formation of multiple layers of coccoliths. The multiple covering was restored after decalcification of stationary cells. Formation of multiple layers of coccoliths may help the cells reach deeper, nutrient-rich water by increasing the sinking rate of the cells.     

Increased coccolith production by Emiliania huxleyi cultures enriched with dissolved inorganic carbon

Cells of Emiliania huxleyi grown on Eppley's medium enriched with dissolved inorganic carbon (DIC) developed multiple layers of coccoliths. The maximum diameter of cells grown in the presence of 13.2 mM DIC was 12.3 m, whereas that of cells grown in the presence of 1.5 mM DIC was 8.0 m. Although enrichment of Eppley's medium with DIC increased both coccolith production and cell growth, coccolith production was enhanced to a greater extent than cell growth. The enrichment of Eppley's medium with DIC was used to enhance production of coccolith particles by E. huxleyi. Repeated-batch culture, in which DIC, Ca²⁺, nitrate and phosphate concentrations in the medium were maintained by replacing the culture medium, was carried out in a closed photobioreactor. During repeated-batch culture, a maximum coccolith yield of 560 mg/l for 2 days and a maximum biomass yield of 810 mg/l for 2 days were achieved. Enrichment and maintenance of DIC is therefore an efficient method for the production of large quantities of coccoliths.     

The last 100,000 years in the western Mediterranean: sea surface water and frontal dynamics as revealed by coccolithophores

A quantitative analysis was carried out on coccolith assemblages from two Pleistocene cores (K1 and K10) from the western Mediterranean. The distribution of selected coccolithophore species provides new paleoclimatic and paleoceano-graphic data. A continuous sequence from the top of Isotope Stage 5 to the Holocene was recorded. The reversal in dominance between Gephyrocapsa muellerae and Emiliania huxleyi was dated in both cores at ca. 73 ka. At about 47 ka, E. huxleyi shows a regular increase, whereas G. muellerae progressively decreases in abundance. During interglacial periods, high concentrations of coccoliths are observed, whereas in glacial times, coccoliths are more diluted and the percentage of reworked forms increases as a consequence of the higher terrigenous input. After taking careful account of the dilution factor, we conclude that the production of coccolithophores was higher during warm periods. Maxima in coccolith concentrations coincide with highstand episodes, probably as a result of the intensification of the Atlantic flux into the Mediterranean across the Gibraltar Strait. This intensification could have produced an increase in nutrient content in the surface Mediterranean waters. During cold periods, the western Mediterranean front underwent a reduction in activity, probably due to an increase in the saline and/or thermal gradients between the superficial waters, and intermediate waters in the Liguro Provençal basin.