(Table T1) Radiolaria abundance of ODP Hole 175-1082A

A primary objective of Leg 175 was to investigate the upwelling history of the Benguela Current. Upwelling along the coast is found over the shelf in several well-established cells, as well as along the shelf-slope break, and extends over the 1000-m isobath. Streaming filaments along the coast also carry upwelled water off shore (Shannon, 1985). The upwelled nutrient-rich waters are sourced from the South Atlantic central water mass, which is a mixture of subtropical and subantarctic water masses. Below the central water mass lies Antarctic intermediate water (Shannon and Hunter, 1988, doi:10.2989/025776188784480735; Stramma and Peterson, 1989, doi:10.1175/1520-0485(1989)019<1440:GTITBC>2.0.CO;2). The upwelling system supports a robust marine community (Shannon and Pillar, 1986) where radiolarians are abundant (Bishop et al., 1978, doi:10.1016/0146-6291(78)90010-3). The endemic nature of radiolarians makes them useful in reconstructing the paleocirculation patterns. The biogeographic distribution of many species is limited by water-mass distribution. In a given geographic region, species may also have discrete depth habitats. However, their depth of occurrence can change worldwide because the depths of water masses vary with latitude (Boltovskoy, 1999). Consequently, species found at shallow depths at high latitudes (cold-water fauna) are observed deeper in the water column at lower latitudes. The low-latitude submergence of cold-water species broadens their distribution, resulting in species distributions that can cover multiple geographic regions (Kling, 1976, doi:10.1016/0011-7471(76)90880-9; Casey, doi:10.1016/0031-0182(89)90017-5; 1971; Boltovskoy, 1987, doi:10.1016/0377-8398(87)90014-4). Since radiolarian distribution is closely related to water-mass distribution and controlled by climatic conditions rather than geographic regions, similar assemblages characterize the equatorial, subtropical, transition, subpolar, and polar regions of ocean basins (Petrushevskaya, 1971a; Casey, 1989, doi:10.1016/0031-0182(89)90017-5; Boltovskoy, 1999). Numerous radiolarian species found in water masses in the Angola and Benguela Current systems have also been observed in plankton net samples, sediment traps, and surface-sediment studies in the Atlantic sector of the Southern Ocean, where they exhibited particular water-mass affinities (Abelmann, 1992a, doi:10.1007/BF00243107; Abelmann 1992b, doi:10.1007/BF00243108; Abelmann and Gowing, 1997, doi:10.1016/S0377-8398(96)00021-7). This report presents data on the radiolarian fauna recovered from Site 1082 sediments in the form of a survey of species reflecting the latitudinal migration of the Angola-Benguela Front and upwelling. The data constitute a time series of relative radiolarian abundances at very high resolution (every 20 cm) of the upper 12 m of Hole 1082A.

(Table 2) Mineralogy, stable isotopes, and mineral percentages of the carbonate fraction of samples from the South China Sea

Chemoherm carbonates, as well as numerous other types of methane seep carbonates, were discovered in 2004 along the passive margin of the northern South China Sea. Lithologically, the carbonates are micritic containing peloids, clasts and clam fragments. Some are highly brecciated with aragonite layers of varying thicknesses lining fractures and voids. Dissolution and replacement is common. Mineralogically, the carbonates are dominated by high magnesium calcites (HMC) and aragonite. Some HMCs with MgCO3 contents of between 30-38 mol%-extreme-HMC, occur in association with minor amounts of dolomite. All of the carbonates are strongly depleted in d13C, with a range from -35.7 to -57.5 per mil PDB and enriched in d18O (+ 4.0 to + 5.3 per mil PDB). Abundant microbial rods and filaments were recognized within the carbonate matrix as well as aragonite cements, likely fossils of chemosynthetic microbes involved in carbonate formation. The microbial structures are intimately associated with mineral grains. Some carbonate mineral grains resemble microbes. The isotope characteristics, the fabrics, the microbial structure, and the mineralogies are diagnostic of carbonates derived from anaerobic oxidation of methane mediated by microbes. From the succession of HMCs, extreme-HMC, and dolomite in layered tubular carbonates, combined with the presence of microbial structure and diagenetic fabric, we suggest that extreme-HMC may eventually transform into dolomites. Our results add to the worldwide record of seep carbonates and establish for the first time the exact locations and seafloor morphology where such carbonates formed in the South China Sea. Characteristics of the complex fabric demonstrate how seep carbonates may be used as archives recording multiple fluid regimes, dissolution, and early transformation events.

(Table 1) Stable isotope data of chemoherm carbonates at Hydrate Ridge on the Cascadia continental margin

Two active chemoherm build-ups growing freely up into the oceanic water column, the Pinnacle and the South East-Knoll Chemoherms, have been discovered at Hydrate Ridge on the Cascadia continental margin. These microbially-mediated carbonate formations rise above the seafloor by several tens of meters and display a pinnacle-shaped morphology with steep flanks. The recovered rocks are pure carbonates dominated by aragonite. Based on fabric and mineralogic composition different varieties of authigenic aragonite can be distinguished. Detailed visual and petrographic investigations unambiguously reveal the involvement of microbes during the formation of the carbonates. The fabric of the cryptocrystalline and fibrous aragonite can be described as thrombolitic. Fossilized microbial filaments in the microcrystalline aragonite indicate the intimate relationship between microbes and carbonates. The strongly 13C-depleted carbon isotope values of the samples (as low as -48.1 per mill PDB) are characteristic of methane as the major carbon source for the carbonate formation. The methane-rich fluids from which the carbonates are precipitated originate most probably from a gas reservoir below the bottom-simulating reflector (BSR) and rise through fault systems. The d18O values of the aragonitic chemoherm carbonates are substantially higher (as high as 5.0 per mill PDB) than the expected equilibrium value for an aragonite forming from ambient seawater (3.5 per mill PDB). As a first approximation this indicates formation from glacial ocean water but other factors are considered as well. A conceptual model is presented for the precipitation of these chemoherm carbonates based on in situ observations and the detailed petrographic investigation of the carbonates. This model explains the function of the consortium of archaea and sulfate-reducing bacteria that grows on the carbonates performing anaerobic oxidation of methane (AOM) and enabling the precipitation of the chemoherms above the seafloor surrounded by oxic seawater. Beggiatoa mats growing on the surface of the chemoherms oxidize the sulfide provided by sulfate-dependent anaerobic oxidation of methane within an oxic environment. The contact between Beggiatoa and the underlying microbial consortium represents the interface between the overlying oxic water column and an anoxic micro-environment where carbonate formation takes place.

SeaWiFS chlorophyll imagery of the iron-fertilized area of SOIREE

The growth of populations is known to be influenced by dispersal, which has often been described as purely diffusive (Kierstead and Slobodkin, 1953; Okubo, 1980). In the open ocean, however, the tendrils and filaments of phytoplankton populations provide evidence for dispersal by stirring (Gower et al., 1980, doi:10.1038/288157a0; Holligan et al., 1993, doi:10.1029/93GB01731). Despite the apparent importance of horizontal stirring for plankton ecology, this process remains poorly characterized. Here we investigate the development of a discrete phytoplankton bloom, which was initiated by the iron fertilization of a patch of water (7 km in diameter) in the Southern Ocean (Boyd et al., 2000, doi:10.1038/35037500). Satellite images show a striking, 150-km-long bloom near the experimental site, six weeks after the initial fertilization. We argue that the ribbon-like bloom was produced from the fertilized patch through stirring, growth and diffusion, and we derive an estimate of the stirring rate. In this case, stirring acts as an important control on bloom development, mixing phytoplankton and iron out of the patch, but also entraining silicate. This may have prevented the onset of silicate limitation, and so allowed the bloom to continue for as long as there was sufficient iron. Stirring in the ocean is likely to be variable, so blooms that are initially similar may develop very differently.

Diagenesis of siliceous particles in ODP Hole 114-699A

A number of neogenic opaline structures, not previously reported in the literature, as well as other neogenic phases are described from four Oligocene to Pliocene biosiliceous sediment samples from Hole 699A. The possible influence of microbes on the formation or the morphology of some of them is discussed. The samples, which are early Pliocene, early to middle Miocene, and late Oligocene (two) in age, were histologically fixed aboard ship upon retrieval. Investigations of the samples used SEM (with Edax/Tracor) and XRD methods. Diagenesis has affected all four samples, but the most extensive development of neoformed structures occurs in the Miocene and uppermost Oligocene samples, where microbial filaments (0.05 to 10 ?m long), microbial colonies, and siliceous microhemispheroids (0.2 to 0.7 µm diameter) were observed. The latter encrust filaments, diatoms, and detrital grains to varying degrees. Other neoformed structures include (1) flakes formed by coalesced microhemispheroids, some of which are guided by short, stubby filaments, which occur only in the Miocene and uppermost Oligocene samples, and (2) flakes characterized by smooth or microfissured surfaces, which grow on diatom frustules and in pore spaces and have a more widespread distribution. The XRD data indicate possible cristobalite formation in the Miocene and uppermost Oligocene samples; we believe that the neoformed opaline structures (encrusted filaments and microhemispheroids) may represent an early phase of opal-CT. The timing of neoformation of most of these features appears to have been fairly recent, continuing even at the time of sampling. There appears to be no direct correlation of this incipient, lower Miocene-uppermost Oligocene diagenetic layer and the pore-water chemistry profiles; a massive increase in shear strength in these sediments, however, may indicate some cementation. Smectite was identified by XRD as the most prominent clay mineral in these generally clay-poor sediments. Honeycombed minerals with filamentous edges, which could correspond to smectite, were observed with SEM in the pore spaces.

Animation of micro-computed tomography of bioerosion trace Entobia megastoma made by East Australian bioeroding sponge Cliona celata

Endolithic bioerosion is difficult to analyse and to describe, and it usually requires damaging of the sample material. Sponge erosion (Entobia) may be one of the most difficult to evaluate as it is simultaneously macroscopically inhomogeneous and microstructurally intricate. We studied the bioerosion traces of the two Australian sponges Cliona celata Grant, 1826 (sensu Schönberg 2000) and Cliona orientalis Thiele, 1900 with a newly available radiographic technology: high resolution X-ray micro-computed tomography (MCT). MCT allows non-destructive visualisation of live and dead structures in three dimensions and was compared to traditional microscopic methods. MCT and microscopy showed that C. celata bioerosion was more intense in the centre and branched out in the periphery. In contrast, C. orientalis produced a dense, even trace meshwork and caused an overall more intense erosion pattern than C. celata. Extended pioneering filaments were not usually found at the margins of the studied sponge erosion, but branches ended abruptly or tapered to points. Results obtained with MCT were similar in quality to observations from transparent optical spar under the dissecting microscope. Microstructures could not be resolved as well as with e.g. scanning electron microscopy (SEM). Even though sponge scars and sponge chips were easily recognisable on maximum magnification MCT images, they lacked the detail that is available from SEM. Other drawbacks of MCT involve high costs and presently limited access. Even though MCT cannot presently replace traditional techniques such as corrosion casts viewed by SEM, we obtained valuable information. Especially for the possibility to measure endolithic pore volumes, we regard MCT as a very promising tool that will continue to be optimised. A combination of different methods will produce the best results in the study of Entobia.

Spatial compartmentalization of free radical formation and mitochondrial heterogeneity in bivalve gills revealed by live-imaging techniques, supplementary material

Live-imaging techniques (LIT) utilize target-specific fluorescent dyes to visualize biochemical processes using confocal and multiphoton scanning microscopy, which are increasingly employed as non-invasive approach to physiological in-vivo and ex-vivo studies. Here we report application of LIT to bivalve gills for ex-vivo analysis of gill physiology and mapping of reactive oxygen (ROS) and nitrogen (RNS) species formation in the living tissue. Our results indicate that H2O2, HOO. and ONOO- radicals (assessed through C-H2DFFDA staining) are mainly formed within the blood sinus of the filaments and are likely to be produced by hemocytes as defense against invading pathogens. The oxidative damage in these areas is controlled by enhanced CAT (catalase) activities recorded within the filaments. The outermost areas of the ciliated epithelial cells composing the filaments, concentrated the highest mitochondrial densities (MTK Deep Red 633 staining) and the most acidic pH values (as observed with ageladine-a). These mitochondria have low (depolarized) membrane potentials (D psi m) (JC-1 staining), suggesting that the high amounts of ATP required for ciliary beating may be in part produced by non-mitochondrial mechanisms, such as the enzymatic activity of an ATP-regenerating kinase. Nitric oxide (NO, DAF-2DA staining) produced in the region of the peripheral mitochondria may have an effect on mitochondrial electron transport and possibly cause the low membrane potential. High DAF-2DA staining was moreover observed in the muscle cells composing the wall of the blood vessels where NO may be involved in regulating blood vessel diameter. On the ventral bend of the gills, subepithelial mucus glands (SMG) contain large mucous vacuoles showing higher fluorescence intensities for O2.- (DHE staining) than the rest of the tissue. Given the antimicrobial properties of superoxide, release of O2.- into the mucus may help to avoid the development of microbial biofilms on the gill surface. However, cells of the ventral bends are paying a price for this antimicrobial protection, since they show significantly higher oxidative damage, according to the antioxidant enzyme activities and the carbonyl levels, than the rest of the gill tissue. This study provides the first evidence that one single epithelial cell may contain mitochondria with significantly different membrane potentials. Furthermore, we provide new insight into ROS and RNS formation in ex-vivo gill tissues which opens new perspectives for unraveling the different ecophysiological roles of ROS and RNS in multifunctional organs such as gills.

Photosynthetic production in the Central Arctic during POLARSTERN cruise ARK-XXVII/3 (IceArc) in 2012

The ice-covered Central Arctic Ocean is characterized by low primary productivity due to light and nutrient limitations. It has been speculated that the recent reduction in ice cover could lead to a substantial increase in primary production, but still little is known as to the fate of the ice-associated primary production, and of nutrient supply with increasing warming. This study presents results from the Central Arctic Ocean collected during summer 2012, when sea-ice reached a minimum extent since the onset of satellite observations. Net primary productivity (NPP) was measured in water column, sea ice and melt ponds by 14CO2 uptake at different irradiances. Photosynthesis vs. irradiance (PI) curves were established in laboratory experiments and used to upscale measured NPP to the deep Eurasian Basin (north of 78°N) using the irradiance-based Central Arctic Ocean Primary Productivity model (CAOPP). In addition, new annual production was calculated from the seasonal nutrient drawdown in the mixed layer since last winter. Results show that ice algae can contribute up to 60% to primary production in the Central Arctic at the end of the season. The ice-covered water column had lower NPP rates than open water probably due to light limitation. According to the nutrient ratios in the euphotic zone, nitrate limitation was detected in the Siberian Seas (Laptev Sea area), while silicate was the main limiting nutrient at the ice margin influenced by Atlantic waters. Although sea-ice cover was substantially reduced in 2012, total annual new production in the Eurasian Basin was 17 ± 7 Tg C/yr, which is similar to previous estimates. However, when including the contribution by sub-ice algal filaments, the annual production for the deep Eurasian Basin (north of 78°N) is 16 Tg C/yr higher than estimated before. Our data suggest that sub-ice algae might be responsible for potential local increases in NPP due to higher light availability under the ice, and their ability to benefit from a wider area of nutrients as they drift with the ice.

The distributions, C- and O-isotopic compositions, and frequency of giant-ooids

Early Triassic oceans were characterized by deposition of a number of "anachronistic facies", including microbialites, seafloor carbonate cement fans, and giant ooids. Giant ooids were particularly prevalent in Lower Triassic sections across South China and exhibit unusual features that may provide insights into marine environmental conditions following the end-Permian mass extinction. The section at Moyang (Guizhou Province) contains abundant giant ooids ranging in size between 2 and 6 mm (maximum 12 mm) and exhibiting various cortical structures, including regular, deformed, compound, regenerated and "domed". Preservation of ooid cortical structure is generally good as indicated by petrographic observations, and trace element and carbon isotope analyses suggest that diagenesis occurred in a closed diagenetic system. All ooids exhibit fine concentric laminae, frequently alternating between light-colored coarsely crystalline and dark-colored finely crystalline layers probably reflecting variation in organic content or original mineralogy. Under scanning electron microscope, biomineralized filaments or biofilms and tiny carbonate fluorapatite (CFA) crystals are commonly found in the finely crystalline layers. We infer that the precipitation of CFA was related to adsorption of P via microbial activity on the surfaces of ooids following episodic incursions of deep waters rich in carbon dioxide, hydrogen sulfide and phosphate into shallow-marine environments. Giant ooid precipitation may have been promoted in shallow ramp settings during these events by increased watermass agitation and supersaturation with respect to calcium carbonate, as well as reduced carbonate removal rates through biotic skeletal formation. Spatio-temporal distribution data reveal that giant ooids were widespread in the Tethyan region during the Early Triassic, and that they were most abundant immediately after the end-Permian crisis and disappeared gradually as metazoans repopulated marine environments.

Biogenic flux and primary production of sediment traps in the Atlantic and the Southern Ocean

Fluxes of organic carbon normalised to a depth of 1000 m from 18 sites in the Atlantic and the Southern Ocean are presented, comprising nine biogeochemical provinces as defined by Longhurst et al. (1995. Journal of Plankton Research 17, 1245-1271). For comparison with primary production, we used a recent compilation of primary production values derived from CZCS data (Antoine et al., 1996. Global Biogeochemical Cycles 10, 57-69). In most cases, the seasonal patterns stood reasonably well in accordance with the carbon fluxes. Particularly, organic carbon flux records from two coastal sites off northwest and southwest Africa displayed a more distinct correlation to the primary production in sectors (1 x 1°) which are situated closer to the coastal environments. This was primarily caused by large upwelling filaments streaming far offshore, resulting in a cross-shelf carbon transport. With respect to primary production, organic carbon export to a water depth of 1000 m, and the fraction of primary production exported to a depth of 1000 m (export fraction=EF1000), we were able to distinguish between: (1) the coastal environments with highest values (EF1000=1.75-2.0%), (2) the eastern equatorial upwelling area with moderately high values (EF1000=0.8-1.1%), (3) and the subtropical oligotrophic gyres that yielded lowest values (EF1000=0.6%). Carbon export in the Southern Ocean was low to moderate, and the EF1000 value seems to be quite low in general. Annual organic carbon fluxes were proportional to primary production, and the export fraction EF1000 increased with primary production up to 350 gCm**-2 yr**-1. Latitudinal variations in primary production were reflected in the carbon flux pattern. A high temporal variability of primary production rates and a pronounced seasonality of carbon export were observed in the polar environments, in particular in coastal domains, although primary production (according to Antoine et al., 1996. Global Biogeochemical Cycles 10, 57-69), carbon fluxes, and the export fraction remained at low.

Seawater carbonate chemistry and dissolution rates by boring microflora during ex situ experiments with dead corals (Porites lobata), 2009

Eight-month-old blocks of the coral Porites lobata colonized by natural Hawaiian euendolithic and epilithic communities were experimentally exposed to two different aqueous pCO2 treatments, 400 ppmv and 750 ppmv, for 3 months. The chlorophyte Ostreobium quekettii dominated communities at the start and at the end of the experiment (65-90%). There were no significant differences in the relative abundance of euendolithic species, nor were there any differences in bioeroded area at the surface of blocks (27%) between pCO2 treatments. The depth of penetration of filaments of O. quekettii was, however, significantly higher under 750 ppmv (1.4 mm) than under 400 ppmv (1 mm). Consequently, rates of carbonate dissolution measured under elevated pCO2 were 48% higher than under ambient pCO2 (0.46 kg CaCO3 dissolved m2/a versus 0.31 kg /m2/a). Thus, biogenic dissolution of carbonates by euendoliths in coral reefs may be a dominant mechanism of carbonate dissolution in a more acidic ocean.

d15N of trap WR2

Temporal changes in d15N values of sinking particles collected with sediment traps in the Benguela upwelling regime off southwest Africa mirrored variations in the input of inorganic nitrogen to the surface water. Reductions in d15N (to as low as 2.5 per mil) corresponded to low sea surface temperatures during austral spring and late austral autumn/early winter, indicating increased nitrate availability due to the presence of recently upwelled water. High particulate fluxes accompanied the low d15N values and sea surface temperatures, reflecting increased productivity, fueled by the upwelled nutrients. High d15N values (up to 13.1 per mil) coincided with high sea surface temperatures and low particle fluxes. In this area, the seaward extension of upwelling filaments, which usually occurs twice yearly, brings nutrient-rich water to the euphotic zone and leads to elevated productivity and relatively lower d15N values of the particulate nitrogen. Satellite images of ocean chlorophyll show that productivity variations coincide with d15N changes. The observed isotopic pattern does not appear to have been caused by variations in the species composition of the phytoplankton assemblage. Calculations based on d15N of the sinking particulate nitrogen show that the surface nitrate pool was more depleted during late austral summer/early fall and mid-winter and that supply exceeded demand during the intense spring bloom and in late austral fall. The main uncertainty associated with these estimates is the effect of diagenesis on d15N and possible variability in preservation of the isotope signal between periods of high and low particle flux.

Planktonic foraminifera in the Arabian Sea

Variations in primary productivity (PP) have been reconstructed in eutrophic, mesotrophic and oligotrophic parts of the Arabian Sea over the past 135 000 years applying principal component analysis and transfer function to planktic foraminiferal assemblages. Temporal variation in paleoproductivity is most pronounced in the mesotrophic northern (NAST site) and oligotrophic eastern (EAST site) Arabian Sea, and comparatively weak in the western eutrophic GeoB 3011-1 site in the upwelling area off Oman. Higher PP during interglacials (250-320 g C/m**2 year) than during cold stages (210-270 g C/m**2 year) at GeoB 3011-1 could have been caused by a strengthened upwelling during intensified summer monsoons and increased wind velocities. At NAST, during interglacials, PP is estimated to exceed g C/m**2 year 1, and during glacials to be as low as 140-180 g C/m**2 year. These fluctuations may result from a (1) varying impact of filaments that are associated to the Oman coastal upwelling, and (2) from open-ocean upwelling associated to the Findlater Jet. At EAST, highest productivity of about 380 g C/m**2 year is documented for the transition from isotope stage 5 to 4. We suggest that during isotope stages 2, 4, 5.2, the transition 5/4, and the end of stage 6, deep mixing of surface waters was caused by moderate to strong winter monsoons, and induced an injection of nutrients into the euphotic layer leading to enhanced primary production. The deepening of the mixed layer during these intervals is confirmed by an increased concentration of deep-dwelling planktic foraminiferal species. A high-productivity event in stage 3, displayed by estimated PP values, and by planktic foraminifera and radiolaria flux and accumulation rate, likely resulted from a combination of intensified SW monsoons with moderate to strong NE monsoons. Differential response of Globigerina bulloides, Globigerinita glutinata and mixed layer species to the availability of food is suited to subdivide productivity regimes on a temporal and spatial scale.

Investigation of time series stations along a productivity gradient north of the Canary Islands

Three stations along a productivity gradient north of the Canary Islands were investigated for surface-water properties, particle flux, and composition (biogenic and lithogenic components, and stable nitrogen isotope composition, delta15N) and export production. Investigation sites along the east-west transect off the NW African upwelling margin included the European Station for Time-Series in the Ocean, Canary Islands (ESTOC), one location contiguous to the NW African upwelling zone in the Eastern Boundary Current (EBC) and one station north of the island La Palma (LP). The seasonality of surface-water properties along the transect was mainly influenced by the winter cooling and simultaneous phytoplankton maximum and, in addition at EBC, by nearby upwelling. Accordingly, particle flux and composition along the transect were closely linked to the winter bloom sedimentation and upwelling related enhanced plankton biomass stemming from the primary upwelling and the Cape Yubi filament at EBC. During all seasons, particle flux was highest at EBC and had the highest contribution of biogenic opal and lithogenic components, and the lowest delta15N compared to the offshore stations. But contrary to what would be expected from the productivity gradient, particle flux did not decrease from ESTOC to LP. Below the upper several hundred meters, particle flux was enhanced by additional particle input along the entire transect, manifested by an increase of flux with depth and lower delta15N values. We offer a scenario in which intermediate nepheloid layers originating from the primary upwelling as well as particle dispersion from upwelling filaments, mainly the Cape Ghir filament, impact on the trap stations as far as 700 km into the open ocean. This study contributes to our understanding of the poorly resolved biogeochemical transition between the productive shelf and subtropical gyre provinces.

Seawater samples along several inshore-offshore transects off NW Africa and SE Africa

For the optimal use in palaeoceanographic studies of the stable oxygen isotopic signal and elemental composition of the calcareous photosynthetic dinoflagellate Thoracosphaera heimii, it is essential to gain detailed information about its calcification depth and spatial distribution. We therefore studied the vertical and horizontal distribution patterns of T. heimii in the upper water column (0-200 m) along three transects: an inshore-offshore gradient off Cape Blanc (CB), a south-north transect from CB to the Portuguese coast and a north-south transect off Tanzania. We compared concentrations of living cysts (cells with cell content) with chlorophyll-a, salinity and temperature measurements at the sampling depth. In order to explore the seasonal variability in cyst production, three transect off CB were sampled at three different times of the year. Living T. heimii cysts were found in the upper 160 m of the water column with highest concentrations in the photic zone indicating that the calcification of T. heimii occurs in the upper part of the water column. Maximal abundances of living cysts were found relatively often in or just above the deep chlorophyll maximum (DCM), the depth of which varies regionally from about 20-40 m off CB to about 80 m off Tanzania and along the transect from CB to the Portuguese Coast. However, there was no significant correlation at the 95% confidence level between the cyst concentrations and temperature, salinity and chlorophyll-a concentrations at the sampling depths observed. In both the Atlantic and Indian Oceans, the highest abundances of T. heimii were observed in regions where the upper water masses contained relatively low nutrient concentrations that are influenced only sporadically, or not at all, by enhanced photic zone mixing related to the presence of upwelling cells or river outflow plumes at or close to the sampling sites. The seasonal production of cysts by T. heimii appears to be negatively related to the presence of upwelling filaments across the sampling sites. Our study suggests that turbulence of the upper water masses is a major environmental factor influencing T. heimii production.