Dinoflagellate cysts from surface sediments of Saldanha Bay, South Africa: an indication of the potential risk of harmful algal blooms

The distribution and abundance of dinoflagellate cysts from recent coastal sediments in Saldanha Bay, was investigated, and compared to the cyst assemblages of the adjacent coastal upwelling system as reflected in the sediments off Lambert's Bay on the southern Namaqua shelf. Twenty-two cyst types were identified from three sample sites off Lambert's Bay with recorded abundances between 1726 and 1863 cysts ml⁻¹ wet sediment. At least 21 distinctive cyst types were identified from 32 sample sites within Saldanha Bay. Cyst abundance in Saldanha Bay was relatively low, averaging 116 cysts ml⁻¹ wet sediment. The region off Lambert's Bay is especially susceptible to the formation of harmful algal blooms attributed to high biomass dinoflagellate blooms. Owing to these blooms and the retentive circulation characteristics of this area, cyst formation and deposition is high. Blooms can be advected into Saldanha Bay, but their development and duration in the Bay is restricted by the system of exchange that operates between the Bay and the coastal upwelling system, in that there is a net export of surface waters from the Bay. Consequently, fewer cysts are formed and deposited within the Bay thereby reducing the likelihood of in situ bloom development initiated from the excystment of cysts.     

Die Anwendung organischwandiger Dinoflagellatenzysten zur Rekonstruktion von Paläoumwelt, Paläoklima und Paläozeanographie: Möglichkeiten und Grenzen

Over the past two decades, organic-walled dinoflagellate cysts (dinocysts) have been increasingly used as paleoenvironmental and paleoclimatic indicators in marine sediments from the Mesozoic and Cenozoic. Because dinocysts are acid-resistant and abundant in nearly all aquatic settings, the dinocyst-based environmental and paleoclimatic information is complementary to the data derived from calcareous and siliceous microfossil groups such as foraminifers, calcareous nannoplankton, diatoms, and radiolarians. In some cases, the environmental information of dinocysts even exceeds that of calcareous and siliceous microfossils. The present contribution provides an outline of the present-day knowledge on dinocyst paleoecology. It reviews the potential of dinocysts for the reconstruction of sea-surface productivity, temperature, and salinity as well as their application in sequence stratigraphy. Finally, it evaluates the applicability of dinocysts in reconstructing the paleo-oxygenation in the bottom and pore waters of marine environments and in the lower water column of epeiric seas.     

The potential of organic-walled dinoflagellate cysts for the reconstruction of past sea-surface conditions in the Southern Ocean

In this study we investigate the potential of organic-walled dinoflagellate cysts (dinocysts) as tools for quantifying past sea-surface temperatures (SST) in the Southern Ocean. For this purpose, a dinocyst reference dataset has been formed, based on 138 surface sediment samples from different circum-Antarctic environments. The dinocyst assemblages of these samples are composed of phototrophic (gonyaulacoid) and heterotrophic (protoperidinioid) species that provide a broad spectrum of palaeoenvironmental information. The relationship between the environmental parameters in the upper water column and the dinocyst distribution patterns of individual species has been established using the statistical method of Canonical Correspondence Analysis (CCA). Among the variables tested, summer SST appeared to correspond to the maximum variance represented in the dataset.To establish quantitative summer SST reconstructions, a Modern Analogue Technique (MAT) has been performed on data from three Late Quaternary dinocyst records recovered from locations adjacent to prominent oceanic fronts in the Atlantic sector of the Southern Ocean. These dinocyst time series exhibit periodic changes in the dinocyst assemblage during the last two glacial/interglacial-cycles. During glacial conditions the relative abundance of protoperidinioid cysts was highest, whereas interglacial conditions are characterised by generally lower cyst concentrations and increased relative abundance of gonyaulacoid cysts. The MAT palaeotemperature estimates show trends in summer SST changes following the global oxygen isotope signal and a strong correlation with past temperatures of the last 140 000 years based on other proxies. However, by comparing the dinocyst results to quantitative estimates of summer SSTs based on diatoms, radiolarians and foraminifer-derived stable isotope records it can be shown that in several core intervals the dinocyst-based summer SSTs appeared to be extremely high. In these intervals the dinocyst record seems to be highly influenced by selective degradation, leading to unusual temperature ranges and to unrealistic palaeotemperatures. We used the selective degradation index (kt-index) to determine those intervals that have been biased by selective degradation in order to correct the palaeotemperature estimates. We show that after correction the dinocyst based SSTs correspond reasonably well with other palaeotemperature estimates for this region, supporting the great potential of dinoflagellate cysts as a basis for quantitative palaeoenvironmental studies.     

The potential of organic-walled dinoflagellate cysts for the reconstruction of past sea-surface conditions in the Southern Ocean

In this study we investigate the potential of organic-walled dinoflagellate cysts (dinocysts) as tools for quantifying past sea-surface temperatures (SST) in the Southern Ocean. For this purpose, a dinocyst reference dataset has been formed, based on 138 surface sediment samples from different circum-Antarctic environments. The dinocyst assemblages of these samples are composed of phototrophic (gonyaulacoid) and heterotrophic (protoperidinioid) species that provide a broad spectrum of palaeoenvironmental information. The relationship between the environmental parameters in the upper water column and the dinocyst distribution patterns of individual species has been established using the statistical method of Canonical Correspondence Analysis (CCA). Among the variables tested, summer SST appeared to correspond to the maximum variance represented in the dataset.To establish quantitative summer SST reconstructions, a Modern Analogue Technique (MAT) has been performed on data from three Late Quaternary dinocyst records recovered from locations adjacent to prominent oceanic fronts in the Atlantic sector of the Southern Ocean. These dinocyst time series exhibit periodic changes in the dinocyst assemblage during the last two glacial/interglacial-cycles. During glacial conditions the relative abundance of protoperidinioid cysts was highest, whereas interglacial conditions are characterised by generally lower cyst concentrations and increased relative abundance of gonyaulacoid cysts. The MAT palaeotemperature estimates show trends in summer SST changes following the global oxygen isotope signal and a strong correlation with past temperatures of the last 140 000 years based on other proxies. However, by comparing the dinocyst results to quantitative estimates of summer SSTs based on diatoms, radiolarians and foraminifer-derived stable isotope records it can be shown that in several core intervals the dinocyst-based summer SSTs appeared to be extremely high. In these intervals the dinocyst record seems to be highly influenced by selective degradation, leading to unusual temperature ranges and to unrealistic palaeotemperatures. We used the selective degradation index (kt-index) to determine those intervals that have been biased by selective degradation in order to correct the palaeotemperature estimates. We show that after correction the dinocyst based SSTs correspond reasonably well with other palaeotemperature estimates for this region, supporting the great potential of dinoflagellate cysts as a basis for quantitative palaeoenvironmental studies.     

The Cenomanian–Turonian boundary event, OAE2 and palaeoenvironmental change in epicontinental seas: New insights from the dinocyst and geochemical records

Organic-walled dinoflagellate cyst (dinocyst) and geochemical records across the Cenomanian–Turonian boundary (CTB) are presented for the NW European reference section at Eastbourne, England. Dinocyst and nannofossil fertility indexes indicate that an upwelling-driven productivity pulse accompanied a eustatic sea-level fall that preceded, by at least 40 kyr, the global rise in δ¹³C values that marks the onset of Oceanic Anoxic Event 2 (OAE2) and the deposition of black shales in the deep ocean. A marine productivity collapse in the latest Cenomanian is evidenced by the falling absolute and relative abundance of peridinioid dinocysts, believed to be the product of heterotrophic dinoflagellates. This biotic change accompanied transgression and sharply rising sea-surface temperatures, following an Atlantic-wide episode of short-lived cooling. Geochemical tracers provide evidence of Caribbean–Colombian plateau volcanism. The increase in water depth caused by the latest Cenomanian transgression resulted in less eutrophic waters in epicontinental seas, where CTB biotic turnover was driven largely by water-mass changes rather than oxygen depletion. The species richness/absolute abundance ratio of dinocysts is proposed as a water-mass stability proxy.     

Calcareous dinoflagellate cysts in South and equatorial Atlantic surface sediments: diversity,distribution, ecology and potential for palaeoenvironmental reconstruction

Several marine, peridiniphycidean dinoflagellate species produce calcareous cysts during their life cycle, which are relatively resistant to chemical and physical degradation and are therefore often found in large quantities in oceanic bottom sediments. Although the use of these calcareous cysts as proxies for palaeoenvironmental and palaeoclimatic reconstructions has seen many advances over the last decade, until now only relatively patchy and regional information was available on their recent distribution patterns and ecology, especially at the species level. In this paper, comprehensive calcareous cyst diversity and distribution data have been compiled from published and unpublished work for 167 South and equatorial Atlantic Ocean surface sediments, ranging from 20°N to 50°S, and 30°E to 65°W. The main aim has been to focus on the complex, often non-linear, relationships between individual species’ distributions and the physicochemical and trophic conditions of the overlying (sub)surface waters through the use of x–y graphs of cyst abundance vs. (sub)surface water environmental parameters, and detrended correspondence analyses. Ten cyst species and the calcareous vegetative coccoid species Thoracosphaera heimii were observed in the bottom sediments, each species showing its own characteristic distribution pattern in relation to the environmental conditions of the upper water masses above them (e.g. sea surface temperature, productivity, stratification). The sensitive reactions of various species to unique combinations of environmental parameters shows that each species has its own specific ecological traits, thus rejecting the bundled use of ‘calcareous cyst accumulation’ as a general proxy for oligotrophy or stratification in future palaeoenvironmental analyses. The acquired ‘reference’ data set of this study is large and diverse enough to allow its future application in quantitative palaeoenvironmental reconstruction models, and shows that there is still an enormous reconstruction potential concealed in many fossil calcareous dinoflagellate cyst assemblages.     

Does dinocyst wall composition really reflect trophic affinity? New evidence from ATR micro-FTIR spectroscopy measurements

Attenuated total reflection (ATR) microscope Fourier transform infrared (micro-FTIR) spectroscopy was used to investigate the dinosporin composition in the walls of modern, organic-walled dinoflagellate resting cysts (dinocysts). Variable cyst wall compositions were observed, which led to the erection of four spectrochemical groups, some with striking similarities to other resistant biomacromolecules such as sporopollenin and algaenan. Furthermore, possible proxies derivable from the spectrochemical composition of modern and fossil dinocysts were discussed. The color of the dinocyst walls was reflected in the spectral data. When comparing that color with a standard and the results of a series of bleaching experiments with oxidative agents, eumelanin was assigned as a likely pigment contributing to the observed color. Following this assignment, the role of eumelanin as an ultraviolet sunscreen in colored dinocysts was hypothesized, and its implications on the autofluorescence and morphological preservation of dinocysts were further discussed. Unlike what had previously been assumed, it was shown that micro-FTIR data from dinocysts cannot be used to unambiguously infer trophic affinities of their associated cells. Finally, using methods with high spatial resolutions (synchrotron transmission micro-FTIR and optical photothermal infrared spectroscopy), it was shown that dinocyst wall layers are chemically homogenous at the probed scales. This study fills a large knowledge gap in our understanding of the chemical nature of dinocyst walls and has nuanced certain assumptions and interpretations made in the past.     

Changes in style and intensity of production in the Southeastern Atlantic over the last 70,000 yr

Accomplishing reliable paleo-reconstructions of productivity and upwelling conditions in eastern boundary current systems requires the use of cores collected in a basin-wide spatial pattern. Based on diatom assemblage analysis and the concentration and the bulk biogenic components of three gravity cores recovered from the Benguela Upwelling System (BUS) between 19° and 25°S, I describe rapid paleoceanographic changes that occurred during the last 70 ka B.P. in the southeastern Atlantic. The pattern of biogenic production and accumulation differs to varying degrees among the three core sites along the SW African coast. The highest sedimentation and accumulation rates at 25°S off Lüderitz conform with the present-day, well-known pattern of highest productivity and most intense coastal upwelling. Highest diatom values at 25°S during MIS3 points to more intense upwelling due to the combination of strong seaward-extending upwelling filaments, shoaling of the upwelled water, and the influence of silicate-rich waters of Antarctic origin. Productivity decreased along the central BUS throughout MIS2, when the siliceous–calcareous productivity regime shifted toward a system dominated by calcareous producers. Although intensity and strength of winds created adequate conditions for upwelling during MIS2, diatom production decreased. The complete replacement of the upwelling-associated diatom flora by a non-upwelling-related diatom community during MIS1 reflects weakened upwelling, weakened seaward extension of the upwelling filaments, and dominance of warmer surface waters. Combining changes in the composition of the diatom assemblage and variations of the bulk biogenic components allows for reliable reconstruction of paleoproductivity and upwelling changes for the SE Atlantic during the last 70 ka B.P.     

Distributions of calcareous dinoflagellate cysts in surface sediments of the western equatorial Atlantic Ocean,and their potential use in palaeoceanography

Only very few studies focus on recent calcareous dinoflagellate cyst diversity, geographic distribution and ecology, so that information on the distribution patterns and environmental affinities of individual cyst species is extremely limited. This information is, however, essential if we want to use calcareous dinoflagellate cysts for palaeoenvironmental reconstruction. Surface sediment samples from the generally oligotrophic western equatorial Atlantic Ocean, offshore northeast Brazil, were therefore quantitatively analysed for their calcareous dinoflagellate cyst content, including the calcareous vegetative coccoid Thoracosphaera heimii. Seven calcareous dinoflagellate cyst species/morphotypes and T. heimii were encountered in high concentrations throughout the area. Substantial differences in the distribution patterns were observed. The highest concentrations of cysts are found in sediments of the more oligotrophic, oceanic regions, beyond the influence of Amazon River discharge waters. Dinoflagellates producing calcareous cysts thus appear to be capable of surviving low nutrient concentrations and produce large numbers of cysts in relatively stable and predictable environments affected by minimal seasonality. To test for the environmental affinities of individual species, distribution patterns in surface sediments were compared with temperature, salinity, density and stratification gradients within the upper water column (0–100 m) over different times of the year, using principal components analysis and redundancy analysis. T. heimii and four of the seven encountered cyst species (Sphaerodinella? albatrosiana, two morphotypes of Sphaerodinella? tuberosa and Scrippsiella regalis) relate to these parameters significantly and the variations in the cyst associations appear to be associated with the different surface water currents characterising the area. The results imply that calcareous dinoflagellate cyst distributions can potentially be used to distinguish between different open oceanic environments and they could, therefore, be useful in tracing water mass movements throughout the late Quaternary.     

Production and retention of biogenic matter in the southeast Beaufort Sea during 2003�C2004: insights from annual vertical particle fluxes of organic carbon and biogenic silica

Regional variability in the annual fluxes of particulate organic carbon (POC) and biogenic silica (Si) at the periphery of the Mackenzie Shelf (Beaufort Sea) was investigated using eight long-term sediment traps moored at ~100-m depth. Relatively high autochthonous POC and Si fluxes were recorded in the Mackenzie Trough (4.1 and 8.9 g m⁻² year⁻¹ respectively) and off Cape Bathurst (6.6 and 79 g m⁻² year⁻¹), two areas where upwelling events are frequently observed. Diatomaceous new production was minimum on the mid-slope of the Mackenzie Shelf (2.8 g C m⁻² year⁻¹), moderate in the Mackenzie Trough (14.5 g C m⁻² year⁻¹), and highest off Cape Bathurst (128.7 g C m⁻² year⁻¹). High annual autochthonous POC flux corresponded to high diatom production. Among sites, the vertical attenuation of the POC flux increased with diatomaceous new production. Hence, the retention of autochthonous POC in the surface layer (<100 m) was highest (95%) at the highly productive site off Cape Bathurst, intermediate (72%) in the moderately productive Mackenzie Trough, and low (4%) at the unproductive mid-slope of the shelf. Our results indicate that, on Arctic shelves, upwelling and the production of diatoms increase the fraction of the POC which is retained in the surface layer and diverted to the pelagic food web. In the relatively unproductive waters of the Arctic Ocean, biological hot spots such as the one identified off Cape Bathurst where the food web promotes retention rather than vertical export could be disproportionately important as feeding grounds for higher trophic levels.     

Late Albian dinoflagellate cyst paleobiogeography as indicator of asymmetric sea surface temperature gradient on both hemispheres with southern high latitudes warmer than northern ones

Late Albian temperature sensitive dinoflagellate cysts are characterized in order to better understand mid-Cretaceous sea surface temperature gradients. Distribution maps of thirty-seven species recovered from one hundred fifty outcrops and deep sea drilling holes (ODP, DSDP) Sites located from low and high paleolatitudes over the two hemispheres (75°N–70°S) are encountered. Fifty years of published data available in eighty-seven articles have been considered and synthesized using a database coupled with a Geographical Information System (GIS). The continuous and disjoint biogeographic patterns of dinocyst species along latitudes define seven climatic belts, four in the northern hemisphere (high, mid-high, mid-low, low latitude belts) and three in the southern hemisphere (low, mid-low, mid-high latitude belts). Dinocysts restricted along latitudes are temperature sensitive species. Limit ranges of temperature sensitive dinocysts of mid-low and mid-high latitude belts reveal mixing belts, located at 40–45°N and 50°–70°S. They represent major palaeofrontal systems as paleosubtropical fronts with strongly mixed water column. The northern frontal system (40–45°N) was located as in the modern ocean. The large southern frontal system (50°S–70°S) was 10°–20° poleward to Antarctica. Semi-quantitative temperature range limits for mid-Cretaceous dinoflagellates and SST gradients in the two hemispheres are suggested by setting dinocyst climatic belts against estimated δ¹⁸O temperature curve from fish teeth. A paleoecological classification is suggested. Latitudinal distribution of extant temperature sensitive dinoflagellate cysts follows the asymmetric modern temperature gradient. Asymmetric latitudinal ranges of Late Albian dinocyst species of mid-low latitude belts restricted between 45°N and 70°S demonstrate asymmetric temperature gradients with southern high latitudes being warmer than northern high latitudes.     

Organic-walled dinoflagellate cyst production in relation to upwelling intensity and lithogenic influx in the Cape Blanc region (off north-west Africa)

Fossil dinoflagellate cyst assemblages are increasingly used in paleoclimatic research to establish paleoenvi‐ronmental reconstructions. To obtain reliable reconstructions, it is essential to know which physical factors influence the cyst production. Most information about the relationship between variations in physical parameters and cyst production is known from middle and higher latitudes. Information from the (sub)tropics is rare. To increase this information, the temporal variation in cyst assemblages from the upwelling area off north‐west Africa (off Mauritania) has been compared to environmental conditions of the upper water column. Samples were collected by the sediment trap CB9, off north‐west Africa (Cape Blanc, 21°15′2″N, 20°42′2″W) between 11 June 1998 and 7 November 1999 at 27.5‐day intervals. Off Cape Blanc, upwelling occurs throughout the year with variable intensity. This region is also characterized by frequently occurring Saharan dust storms. Seasonal variations in dust input, upwelling intensity and sea surface temperature are reflected by the production of organic‐walled dinoflagellate cyst assemblages. Several cyst taxa are produced throughout the sampling interval, with the highest fluxes at times of strongest upwelling relaxation and/or dust input (Echinidinium aculeatum Zonneveld, Echini‐din ium delicatum Zonneveld, Echinidinium granulaturn Zonneveld, Echinidinium spp., Impagidinium aculeatum (Wall) Lentin et Williams, Impagidinium sphaeri‐cum (Wall) Lentin et Williams, Protoperidinium americanum (Gran et Braarud) Balech, Protoperidinium stellatum (Wall in Wall et Dale) Rochon etal., Protoperidinium spp., Selenopemphix nephroides (Benedek) Benedek et Sarjeant and Selenopemphix quanta (Bradford) Matsuoka). Species such as, for example, Bitectatodinium spongium (Zonneveld) Zonneveld et Jurkschat and Impagidinium patulum (Wall) Stover et Evitt do not show any production pattern related to a particular season of the year or to specific environmental conditions in the upper water column. The production of cysts of Protoperidinium monospinum (Paulsen) Zonneveld et Dale is restricted to intervals with increased nutrient concentrations in upper waters when sea surface temperatures at the sampling site is below approximately 24°C.     

SEABIRDS OF THE SENEGAL UPWELLING AND ADJACENT WATERS

Seabird distributions were observed off Senegal during the upwelling season there, in February-March 1976, and are interpreted against an oceanographic background. Gannets Sula bassana , Pomarine Skuas Stercorarius pomarinus , Great Skuas Catharacta skua , Lesser Black-backed Gulls Larus fuscus , Black-headed Gulls L. ridibundus , Sandwich Terns Sterna sandvicensis and Royal Terns S. maxima were the dominant species on the shelf and slope adjacent to the inshore upwelling zone. Grey Phalaropes Phalaropus fulicarius were the only seabirds associated with an oceanic 'front' farther offshore; it is suggested that this and similar boundary zones are important feeding areas during the pelagic phase of this species' annual cycle.
Leach's Storm-Petrels Oceanodroma leucorhoa were seen east to 20W, but the predominant storm-petrel of the waters between 23W and the continental slope was the Madeiran O. castro. It is suggested that the hatching period of Madeiran Storm-Petrels on the Cape Verde Islands is timed to take advantage of the arrival there of young fish in the surface waters, spawned off the African coast and drifted offshore. However Madeiran Storm-Petrels make no direct use of the coastal upwelling; nor do Little Shearwaters Puffinus assimilis or White-faced Storm-Petrels Pelagodroma marina , the other winter-breeding Cape Verdes seabirds, which probably feed close to the Islands.
The Senegal upwelling is underexploited by seabirds, in contrast to comparable upwelling systems elsewhere. It is suggested that this is due to the seasonal timing of the upwelling, the scarcity of secure nest-sites for breeding seabirds, and the competing attractions of the Benguela upwelling in the South Atlantic.     

Dinoflagellate cyst distribution in marine surface sediments off West Africa (17–6°N) in relation to sea-surface conditions, freshwater input and seasonal coastal upwelling

An organic-walled dinoflagellate cyst analysis was carried out on 53 surface sediment samples from West Africa (17–6°N) to obtain insight in the relationship between their spatial distribution and hydrological conditions in the upper water column as well as marine productivity in the study area.Multivariate analysis of the dinoflagellate cyst relative abundances and environmental parameters of the water column shows that sea-surface temperature, salinity, marine productivity and bottom water oxygen are the factors that relate significantly to the distribution patterns of individual species in the region.The composition of cyst assemblages and dinoflagellate cyst concentrations allows the identification of four hydrographic regimes; 1) the northern regime between 17 and 14°N characterized by high productivity associated with seasonal coastal upwelling, 2) the southern regime between 12 and 6°N associated with high-nutrient waters influenced by river discharge 3) the intermediate regime between 14 and 12°N influenced mainly by seasonal coastal upwelling additionally associated with fluvial input of terrestrial nutrients and 4) the offshore regime characterized by low chlorophyll-a concentrations in upper waters and high bottom water oxygen concentrations.Our data show that cysts of Polykrikos kofoidii, Selenopemphix quanta, Dubridinium spp., Echinidinium species, cysts of Protoperidinium monospinum and Spiniferites pachydermus are the best proxies to reconstruct the boundary between the NE trade winds and the monsoon winds in the subtropical eastern Atlantic Ocean. The association of Bitectatodinium spongium, Lejeunecysta oliva, Quinquecuspis concreta, Selenopemphix nephroides, Trinovantedinium applanatum can be used to reconstruct past river outflow variations within this region.     

The Apectodinium acme and terrestrial discharge during the Paleocene-Eocene thermal maximum: new palynological, geochemical and calcareous nannoplankton observations at Tawanui, New Zealand

Manifestations of profound perturbations in biogeochemical systems during the Paleocene-Eocene thermal maximum (PETM) include a prominent global negative δ¹³C and a pronounced increase in the relative abundance of dinoflagellate cysts (dinocysts) assigned to the genus Apectodinium. While motile representatives of Apectodinium were most likely thermophilic and heterotrophic, the underlying causes of this dinoflagellate response are not well understood. Here we provide new insight by examining the palynology, chemistry and calcareous nannoplankton across the PETM in a continental slope section at Tawanui, New Zealand. Across the PETM, marked changes in the relative abundance of Apectodinium vary antithetically with significant changes in the δ¹³C of carbonate and organic matter. In general, the high relative abundance of Apectodinium relates to enhanced concentrations of dinocysts, signifying a ‘bloom’ of Apectodinium in surface waters during the PETM. Changes in Apectodinium and δ¹³C records correspond to variations in many other parameters, including a smaller negative shift in bulk carbonate δ¹³C than expected, increased terrestrial palynomorphs, elevated TOC and C/N ratios, lower carbonate contents, higher SiO₂ and Al₂O₃ contents, and lower Si/Al ratios. All of these variations can be explained by an increase in delivery of terrigenous material to the continental margin. A peak in the relative abundance of Glaphyrocysta dinocysts at the onset of the PETM may indicate greater down slope transport of neritic material. Changes in calcareous nannoplankton abundances suggest increased nutrient availability in surface waters during the PETM. The combined results show that Apectodinium-dominated assemblages, global perturbations in carbon isotopes and enhanced terrigenous delivery closely correspond in time at Tawanui. A sudden and massive carbon injection to the ocean-atmosphere system may have enhanced weathering and increased terrigenous inputs to continental margins during the PETM. We further suggest that these inputs caused the Apectodinium acme by elevating primary productivity in marginal seas.     

Scrippsiella trochoidea and scrippsiella regalis, nov. comb. (peridiniales, dinophyceae): a comparison

Culture experiments on dinoflagellates from the Atlantic Ocean revealed Scrippsiella regalis (Gaarder) Janofske, nov. comb., a calciodinelloid species with a spherical spiny calcareous cyst. This calcareous cyst was collected previously from plankton and sediment samples, where it was described as the coccolithophorid Discosphaera regalis Gaarder or was often mistaken for the cyst of Scrippsiella trochoidea (von Stein) Loeblich III. The morphological features of both the cellulosic theca and the calcareous cyst of S. trochoidea and S. regalis were compared with respect to their systematic position and the emendation of taxa. Both species were found to have different distribution patterns. Scrippsiella trochoidea is known only from the neritic environment, whereas S. regalis has been found mostly in oceanic samples. The preservation of these spiny calcareous dinocysts in the (fossil) sediment was dependent on the ultrastructure of the calcareous layer of the cyst wall.     

THE CALCAREOUS RESTING CYST OF PENTAPHARSODINIUM TYRRHENICUM COMB. NOV. (DINOPHYCEAE)1

While investigating dinoflagellate cyst assemblages in surface sediments of the Gulfs of Naples and Salerno (Mediterranean Sea), we found a new calcareous resting cyst. This cyst has a round to oval body surrounded by a thick mineral layer, which gives it the shape of a Napoleon hat, with a flat, oval face bearing the archeopyle and a convex keel on the opposite side. The cyst shape is variable in both natural samples and clonal cultures. The organic membrane underlying the calcareous covering is resistant to acetolysis, thus demonstrating the presence of sporopolleninlike material. The cyst germinated into a motile stage having the same morphological features and thecal plate pattern as Peridinium tyrrhenicum Balech. We believe the validity of the genus Pentapharsodinium Indelicato & Loeblich should be accepted. Based on the comparative examination of the species we collected and of a similar species, Pentapharsodinium trachodium Indelicato & Loeblich, we propose Pentapharsodinium tyrrhenicum as a new combination for Peridinium tyrrhenicum. The genus Pentapharsodinium also includes P. dalei Indelicato & Loeblich (= Peridinium faeroense Dale), which produces spiny, organic-walled cysts. The presence of species forming calcareous cysts and species producing noncalcareous cysts in the same genus raises questions about maintaining the family Calciodinellaceae. This family should only include calcareous cyst-forming peridinioids, in order to maintain a unified system of classification of fossil and recent dinoflagellates.     

Diatoms in surface sediments of the Indonesian Archipelago and their relation to hydrography

Diatoms from 53 surface sediments (water depth 350–7200 m) of the Indonesian Archipelago were studied to determine the recent distribution of important assemblages. Three significant assemblages were distinguished, each related to hydrographic parameter(s) of the overlying water mass. The first assemblage is related to the warm saline surface waters of Pacific and Indian Ocean origin; the second assemblage reflects the low-salinity lobe in Makassar Strait; and the third corresponds to major seasonal upwelling areas in the Arafura Sea and south of Java.The normalized ratio of typically Pacific Ocean and Indian Ocean species reflects the fluctuations of the Pacific versus Indian Ocean inflow during the Late Quaternary.Absolute diatom abundances (ADA; in valves g^–1 carbonate-free dry wt) and diatom accumulation rates (DAR; in valves CM^–2 ka^–1) of autochthonous species did not correspond to daily rates of primary production in the photic zone and consequently cannot be used for paleoproductivity estimates.Besides vertical transport some lateral transport of diatoms occurs, as was demonstrated by the presence of three groups of allochthonous species. They are indicators of productivity in the littoral environment, bottom currents and river outflow.     

Calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea: distribution patterns and influence of main environmental gradients

The distribution of calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea was quantitatively analysed. The samples contain 11 cyst species and the vegetative coccoid Thoracosphaera heimii. Cyst abundance increases towards the deeper parts of the basins and is generally higher in the eastern Mediterranean Sea. Three major distribution characteristics exist: (1) different assemblages in oceanic and neritic regions, (2) little agreement with the associations of areas studied so far like the Atlantic Ocean, and (3) a unique oceanic assemblage in the eastern Mediterranean Sea. A gradual change in cyst assemblages from the western to the eastern Mediterranean Sea was observed and statistically compared with the main environmental gradients in the upper water column. Temperature, nitrate concentration and possibly salinity appear to be the most important factors controlling cyst production. Three groups containing cysts with similar environmental preferences can be distinguished: (1) an eastern Mediterranean group related to relatively high temperature and salinity but low nitrate concentration, (2) a group of more or less consistently abundant cosmopolitan species tolerating or even preferring relatively low temperature and salinity but high nitrate concentration, and (3) a group containing species that are possibly adapted to neritic environments and have probably been transported from coastal areas into the studied regions. In contrast to other calcareous plankton, calcareous dinoflagellate cysts correlate strongly with the main environmental gradients in the Mediterranean Sea, bearing a high potential for palaeoenvironmental reconstructions.     

Gradients in microbial methanol uptake: productive coastal upwelling waters to oligotrophic gyres in the Atlantic Ocean

Methanol biogeochemistry and its importance as a carbon source in seawater is relatively unexplored. We report the first microbial methanol carbon assimilation rates (k) in productive coastal upwelling waters of up to 0.117±0.002 d⁻¹ (∼10 nmol l⁻¹d⁻¹). On average, coastal upwelling waters were 11 times greater than open ocean northern temperate (NT) waters, eight times greater than gyre waters and four times greater than equatorial upwelling (EU) waters; suggesting that all upwelling waters upon reaching the surface (⩽20 m), contain a microbial population that uses a relatively high amount of carbon (0.3–10 nmol l⁻¹d⁻¹), derived from methanol, to support their growth. In open ocean Atlantic regions, microbial uptake of methanol into biomass was significantly lower, ranging between 0.04–0.68 nmol l⁻¹d⁻¹. Microbes in the Mauritanian coastal upwelling used up to 57% of the total methanol for assimilation of the carbon into cells, compared with an average of 12% in the EU, and 1% in NT and gyre waters. Several methylotrophic bacterial species were identified from open ocean Atlantic waters using PCR amplification of mxaF encoding methanol dehydrogenase, the key enzyme in bacterial methanol oxidation. These included Methylophaga sp., Burkholderiales sp., Methylococcaceae sp., Ancylobacter aquaticus, Paracoccus denitrificans, Methylophilus methylotrophus, Methylobacterium oryzae, Hyphomicrobium sp. and Methylosulfonomonas methylovora. Statistically significant correlations for upwelling waters between methanol uptake into cells and both chlorophyll a concentrations and methanol oxidation rates suggest that remotely sensed chlorophyll a images, in these productive areas, could be used to derive total methanol biological loss rates, a useful tool for atmospheric and marine climatically active gas modellers, and air–sea exchange scientists.