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.     

Preservation of organic-walled dinoflagellate cysts in different oxygen regimes: a 10,000 year natural experiment

The occurrence of organic-walled dinoflagellate cysts in (fossil) sediments depends on several factors, including as the ecological preferences of the cyst-forming dinoflagellates, cyst production, transport and preservation. Although laboratory experiments have shown that several cyst species are sensitive to chemical treatment, no information about the selective preservation of dinoflagellate cyst species in natural environments has previously been presented. Here, we present data on the effects of oxygen availability in bottom sediments on a cyst assemblage from the ungraded Madeira Abyssal Plain f-turbidite of which only the upper layer has been oxidized. Based on differences in species composition between the oxidized and underlying, unoxidized layers of this turbidite, the influence of oxygen availability on the preservation of individual species has been estimated. Cyst species have been classified in ascending order of resistance to oxygen availability in sediments as: (1) highly sensitive (cysts formed by Protoperidinium species), (2) moderately sensitive (e.g. Spiniferites species), (3) moderately resistant (e.g. Impagidinium paradoxum and Nematosphaeropsis labyrinthus) and (4) resistant (e.g. Impagidinium aculeatum).     

Cyst and radionuclide evidence demonstrate historic Gymnodinium catenatum dinoflagellate populations in Manukau and Hokianga Harbours, New Zealand

Between May 2000 and February 2001, a major bloom of the toxic dinoflagellate Gymnodinium catenatum (a causative organism of Paralytic Shellfish Poisoning, PSP) affected over 1500 km of coastline of New Zealand’s North Island. As this was the first record of this species in New Zealand, we aimed to resolve whether this represented a recent introduction/spreading event or perhaps an indigenous cryptic species stimulated by environmental/climatic change. To answer this question, we analysed for G. catenatum resting cysts in ²¹⁰Pb dated sediment cores (18–34 cm long; sedimentation rates 0.34–0.69 cm per year) collected by SCUBA divers from Manukau Harbour, where the species was first detected, and from Hokianga Harbour, where the highest shellfish toxicity was recorded, while using Wellington Harbour as a well-monitored control site. The results of this study conclusively demonstrate that abundant G. catenatum has been in northern New Zealand at least since the early 1980s, increasing up to 1200 cysts/g around the year 2000, but with low cyst concentrations possibly present since at least 1937. In contrast, Wellington Harbour cores contained only very sparse G. catenatum cysts (8 cysts/g), present only to a depth of 7 cm (surface mixed layer depth), reflecting an apparent recent range expansion of this dinoflagellate in New Zealand, possibly stimulated by unusual climatic conditions associated with the 2000 La Nina event. The significant increases since the early 1980s also of Protoperidinium cysts at Hokianga Harbour and of Gonyaulax, Protoperidinium and Protoceratium cysts at Manukau Harbour suggest a broad scale environmental change has occurred in Northland, New Zealand.     

Dinoflagellate resting cysts as factors in phytoplankton ecology of the North Sea

The occurrence and distribution of dinoflagellate resting cysts were investigated at 11 locations in the south-eastern part of the North Sea. Twenty-six known cyst species and 7 unknown cyst types, which may act as seed population for planktonic dinoflagellate blooms, have been recorded for the first time in the area. The most common cysts in recent sediments were those ofScrippsiella trochoidea, Zygabikodinium lenticulatum, Peridinium dalei, Scrippsiella lachrymosa, Protoceratium reticulatum, Protoperidinium denticulatum, andP. conicum. At all stations,S. trochoidea dominated the cyst assemblages with a maximal abundance of 1303 living cysts/cm³ in the uppermost half centimetre. Cysts of the potentially toxic dinoflagellatesAlexandrium cf.excavatum andA. cf.tamarense were scarce. In the upper 2-cm layer of sediment, dinoflagellate cysts were found in concentrations of 1.8 up to 682 living cysts/cm³. Empty cysts constituted 22–56% of total cyst abundance. The comparative distribution of the cysts showed a general increase in abundance from inshore sites to the offshore area, whereby sandy stations exhibited the lowest cyst abundance and diversity. The wide distribution of living and empty cysts ofScrippsiella lachrymosa suggests that its motile form, which has not been officially recorded in the area until now, is a common plankton organism in German coastal waters. The relatively high abundance of cysts in recent sediments demonstrates the potential importance of benthic resting stages for the initiation of dinoflagellate blooms in the study area.     

Cyst formation,sedimentation, and preservation: Factors affecting dinoflagellate assemblages in recent sediments from trondheimsfjord,Norway

Plankton records and 25 samples of Recent sediment from Trondheimsfjord and the adjoining shelf were studied to investigate production, sedimentation, and preservation of cysts, as factors which influence the eventual composition of dinoflagellate cyst assembleges. All sediment samples were examined for dinoflagellate cysts using routine semiquantitative palynological procedures. In addition, fjord sediments were subjected to a limited sediment analysis, and, for three samples, results from preparations both with and without acid treatments were compared. For the first time, cyst assembleges from Recent sediments were directly compared with extensive plankton records from overlying waters. Results indicate that approximately 20% of the 55 locally recorded dinoflagellate species contribute cysts to bottom sediments. Once formed, cysts behave as fine silt particles in the sedimentary regime, increasing in abundance as the percentage abundance of finer sediment increases, usually with increased water depth. Cyst-forming species are almost entirely restricted to a few genera, particularly Gonyaulax and Peridinium, within the order Peridiniales. For some groups, reasonably good correspondence was found between percentage abundances of dinoflagellates in plankton and their cysts in sediment, though plankton records covering at least five years were required to establish this. Gonyaulax grindleyi Reinecke (Von Stosch 1969) appeared to be consistently overrepresented by cysts in sediment relative to available plankton evidence; possible explanations are suggested. At least 30% of the cyst species present, including most Peridinium species, were eliminated, or rendered unreliable for semiquantitative palynology, by application of routine palynological preparation treatments. Such cysts may provide useful, non-quantitative, palynological information from Recent and possibly Quaternary sediments, but their persistence would seem unlikely. Thus, factors of preservation probably further restrict the dinoflagellate fossil record. Cyst assemblages from Trondheimsfjord are comparable with those previously recorded from the northeastern coast of U.S.A., and from Scotland and northeastern England. Fjord assemblages are dominated by small, simple, spinose cysts which would be regarded as acritarchs if culture experiments had not proved that they are dinoflagellate cysts. Much potential biogeographic and palaeoenvironmental information was included within the less abundant species.Attention is drawn to the role which future culture experiments may be expected to play in helping to resolve taxonomic difficulties currently affecting dinoflagellate studies. Palynological significance of results from the present study is discussed especially with reference to recent work by Von Stosch which strongly suggests that cysts may be hypnozygotes formed routinely in sexual cycles of dinoflagellates.     

Comparison of radiolarian and sedimentologic paleoproductivity proxies in the latest Miocene–Recent Benguela Upwelling System

Estimating past ocean productivity from ocean sediments often gives different results depending on the measurement used. We have examined a suite of paleoproductivity proxies in latest Miocene–Recent sediments from DSDP Site 532 and ODP Site 1084, two deep-sea sections underlying the Benguela Upwelling System off the Atlantic coast of southern Africa. The productivity history of this system has been previously established via organic carbon concentration, diatom floras and alkenone based estimates of surface water temperature, and shows a change from low productivity in the early Pliocene to sustain high productivity in the late Pliocene–Recent. Each of our samples was split and simultaneously analysed for several proxies of ocean productivity, including organic carbon (TOC%), carbonate, abundance of opaline radiolarians, accumulation rate of benthic foraminifera (BFAR); the radiolarian faunal composition indices Upwelling Radiolarian Index (URI) and the Water Depth Ecology index (WADE); other proxies for opal and carbonate dissolution, plus stable isotopes of benthic foraminifera. Comparisons between proxies in the same measured samples, between sites in downcore plots and to the published productivity record for this region suggest that TOC and radiolarian faunal composition, particularly the WADE index, are good indicators of past productivity, albeit with different sensitivities (log–linear correlation WADE–TOC% r = 0.78, n = 65, p < 0.01). In contrast, carbonate, and carbonate-based proxies such as BFAR primarily reflect changes in dissolution. Radiolarian faunal composition indices do not appear to be affected by bulk opal accumulation or changes in opal preservation. WADE analysis of radiolarian faunas and TOC% measurements appear to be useful proxies for productivity in late Neogene sediments, particularly for sections where opal or carbonate dissolution is significant.     

Dinoflagellate cysts as indicators of water quality and productivity in British Columbia estuarine environments

Palynological analyses of 60 surface sediment samples from estuarine environments near Vancouver Island, including the Georgia Strait (GS), the Effingham (EFF) and the Seymour–Belize (SB) Inlets were performed in order to document the distribution of dinoflagellate cyst assemblages and their relationship to hydrographic conditions, productivity and nutrient concentrations. We tested transfer functions using the analogue method, and suggest that dinoflagellate cyst assemblages can be used to reconstruct primary productivity, temperature and salinity. The EFF and SB Inlets are characterized by a dominance of autotrophic taxa, particularly Operculodinium centrocarpum, whereas the Protoperidinioid and gymnodinial heterotrophic taxa such as Quinquecuspis concreta and Brigantedinium spp. dominate the assemblages of the GS. Multivariate analysis shows that this distribution is closely linked to primary productivity, sea-surface temperature (SST) and spring silica concentration. The abundance of autotrophic taxa in the EFF and SB Inlets is associated with high primary productivity and low summer SST, indicating summer upwelling of coastal British Columbia, whereas the heterotrophic taxa that characterizes the GS assemblages are related to low productivity, high summer SST and high silica concentration during spring. Multivariate analysis shows that the most important environmental parameters related to dinocyst distribution in the restricted embayment of the GS, are distance to the shore, distance to Vancouver Harbor, spring sea surface salinity (SSS), spring phosphate concentration and spring productivity. The autotrophic taxa are generally more common in coastal and shallow waters, but Spiniferites ramosus and Pentapharsodinium dalei show an opposite correlation to spring productivity and salinity. P. dalei is particularly abundant around Vancouver Harbor, near highly urbanized shores and within the Fraser River plume, where salinity is low and spring productivity and continental runoff are high. S. ramosus shows its highest abundance on the western coast of GS. Protoperidinioid and gymnodinial cysts characterize distal zones within the central and southern GS that are associated with a mixture of brackish waters coming from the Fraser River and deep upwelling waters entering the GS via Juan de Fuca Strait. The relationship between dinoflagellate cyst assemblages and primary productivity in these estuarine systems differs from that in oceanic and outer neritic zones, where the abundance of heterotrophic taxa is commonly associated with upwelling and high productivity.     

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.     

Recent Dinoflagellate cyst distribution in the North Canary Basin,NW Africa

The North Canary Basin (NW Africa) falls within a major eastern boundary upwelling system. This part of the coastal upwelling system is seasonal and is characterised by the development of large filaments migrating seawards. Hence, 16 samples from this location were selected to identify an “upwelling signal” in the composition of the dinoflagellate cyst assemblages.

Samples closest to the most intense upwelling cells are dominated by L. machaerophorum and G. catenatum and Protoperidinium spp. These make up the “upwelling signal” characteristic for the system. Moreover, the “upwelling signal” can be advected offshore, with filaments that may extend as far as 300 km. Finally, the finding of cysts from G. catenatum, a toxic dinoflagellate, raises the need for a better understanding of the relationship between its presence and distribution in the region, and the coastal upwelling system.     

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.     

Dinoflagellate cysts from sediment traps deployed in the Bellingshausen,Weddell and Scotia seas,Antarctica

Dinoflagellate cysts have been recovered from six long-term (1–2 yr) sediment trap moorings deployed in the Bellingshausen, Weddell and Scotia seas, Antarctica. These traps, mostly moored near the sea bed to sample the nepheloid layer, were located both within and to the north of the maximum sea-ice limit. The numbers of cysts, together with the composition of the assemblages, reinforce the importance of the maximum sea-ice limit as a modern biogeographic boundary for the distribution of dinoflagellate cysts. Cysts derived from heterotrophic dinoflagellates make up the highest proportions within the assemblages recovered from the traps. One trap sampled the export production, revealing little difference in cyst flux over those sampling the nepheloid layer. Cyst flux appears to be highest in areas closest to the Antarctic Convergence, north of the maximum sea-ice limit, and to high nutrient availability. There are, however, differences between the sediment trap assemblages and those recovered from core-top samples at the same or nearby sites. These differences, in the greater number of cysts, and in the higher numbers of round, brown Protoperidinium cysts in the traps, may reflect annual differences in the primary productivity and/or cyst production in the area. In some areas the sediment record may preserve little information about local surface water productivity because of the activity of bottom water currents, for example those arising from the Antarctic Circumpolar Current.     

Late Holocene variability in pelagic fish scales and dinoflagellate cysts along the west coast of Vancouver Island, NE Pacific Ocean

Fish stocks and dinoflagellates are essential components of the marine food chain. Sediment cores from a predominantly anoxic basin in Effingham Inlet, Vancouver Island, British Columbia, archive a late Holocene (500–5300 years BP) record of paleoproductivity in the North American Coastal Upwelling Domain (CUD). We present evidence that late Holocene changes in the dinoflagellate cyst assemblages, sedimentary record, and fish stocks in the northeastern Pacific Ocean fluctuated, at least partially, in accordance with regional and global climate cycles.Principal components analysis (PCA), and trend, wavelet and spectral analyses were used to identify relationships, cycles and trends in sediment grey-scale values, and the abundances of fish scales and dinoflagellate cysts on centennial to millennial time scales. Most observed cycles fluctuated in intensity over time, particularly following transition of the regional climate to a higher rainfall phase that impacted coastal oceanic dynamics 3400 ± 150 years ago. Correlation of the marine paleoproductivity records observed in Effingham Inlet with solar influenced climate proxy cycles observed in the North Atlantic region indicates that solar forcing at different scales might have influenced the climate in the northeast Pacific as well. In particular an 1100- to 1400-year cycle in regional climate is well represented in the fish productivity proxy and sedimentological record. It was also observed that colder water, high-productivity, Selenopemphix nephroides and anchovy-dominated “Anchovy Regime” ecosystems alternate with warmer water, herring-dominated “Herring Regime” ecosystems at millennial time scales. The fish scale record preserved in Effingham Inlet indicates that the NE Pacific is now in transition from an ‘anchovy-’ to a ‘herring’-dominated regime.     

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.     

Biogeography of dinoflagellate cysts in northwest Atlantic estuaries

Few biogeographic studies of dinoflagellate cysts include the near‐shore estuarine environment. We determine the effect of estuary type, biogeography, and water quality on the spatial distribution of organic‐walled dinoflagellate cysts from the Northeast USA (Maine to Delaware) and Canada (Prince Edward Island). A total of 69 surface sediment samples were collected from 27 estuaries, from sites with surface salinities >20. Dinoflagellate cysts were examined microscopically and compared to environmental parameters using multivariate ordination techniques. The spatial distribution of cyst taxa reflects biogeographic provinces established by other marine organisms, with Cape Cod separating the northern Acadian Province from the southern Virginian Province. Species such as Lingulodinium machaerophorum and Polysphaeridinium zoharyi were found almost exclusively in the Virginian Province, while others such as Dubridinium spp. and Islandinium? cezare were more abundant in the Acadian Province. Tidal range, sea surface temperature (SST), and sea surface salinity (SSS) are statistically significant parameters influencing cyst assemblages. Samples from the same type of estuary cluster together in canonical correspondence analysis when the estuaries are within the same biogeographic province. The large geographic extent of this study, encompassing four main estuary types (riverine, lagoon, coastal embayment, and fjord), allowed us to determine that the type of estuary has an important influence on cyst assemblages. Due to greater seasonal variations in SSTs and SSSs in estuaries compared to the open ocean, cyst assemblages show distinct latitudinal trends. The estuarine context is important for understanding present‐day species distribution, the factors controlling them, and to better predict how they may change in the future.     

九龙江西陂库区沉积物甲藻孢囊的分布

孢囊在甲藻的生活史中发挥重要作用,福建省九龙江从2009年起,暴发多次拟多甲藻水华事件。采用显微镜观察和单细胞PCR技术,对九龙江西陂库区2012—2013年不同月份沉积物中的甲藻孢囊进行种属判定,并对甲藻孢囊的分布及其影响因素进行分析。结果表明,西陂库区沉积物中的甲藻孢囊主要为拟多甲藻属,约占80%,其次为裸甲藻属,发现了2009年水华的优势种佩式拟多甲藻(Peridiniopsis penardii)孢囊。库区沉积物中甲藻孢囊的丰度在(13.7±1.2)—(105.2±8.3)个/g干重之间。多元相关分析结果显示甲藻孢囊的丰度与含水率呈现显著正相关性(P0.05),反映了甲藻孢囊沿水流方向逐渐积累。本研究结果填补了国内水库甲藻孢囊鉴定和萌发的研究空白,为九龙江甲藻水华的防治提供科学参考。     

Horizontal dinoflage11ate cyst distribution, sediment characteristics and benthic flux in Manila Bay, Philippines

The lateral variation of sediment properties and associated cyst content of sediment in Manila Bay were determined and their possible role/s in the occurrences of Pyrodinium bahamense Plate var. compressum (Bohm) Steidinger, Tester et Taylor toxic blooms were assessed. Manila Bay's surface sediment was determined to be silt dominated. Clay generally increased towards the coast, probably as a result of flocculation and rapid deposition upon entry of sediments from the rivers. High sand content characterized the southeastern part of the bay attributed to the greater sand inputs and relatively strong currents in this area. Bulk densities were lower in the eastern side of the bay from dilution by high organic load from sewage and urban areas. Benthic flux calculations, particularly NH3, suggest more than 50% nutrient contribution comes from sediments. In general, dinoflagellate cyst density increased from the center of the bay towards the coast, except in Pampanga Bay where it deceased near the coasts. A maximum of 23 dinoflagellate species were identified: 5 were autotrophic (Lingulodinium polyedrum (Stein) Dodge, Gonyaulax spp., Pyrophacus steinii (Schiller) Wall et Dale, Protoceratium reticulatum (Claparede et Lachmann) Butschli, and Pyrodinium bahamense var. compressum), and the rest were predominantly composed of Protoperidinium spp. and Diplopsalis spp. Heterotrophs comprised about 70% of the total cyst counts. Pyrodinium counts increased towards the northwestern part of the bay where it was the dominant autotroph species. Negative correlations were observed for live Pyrodinium cyst density and N flux, P flux, ratio of N to P and total organic carbon (TOC) content. However, areas with high N:P ratio contain abundant Pyrodinium live cysts.     

Control of modern dinoflagellate cyst distribution in the Irish and Celtic seas by seasonal stratification dynamics

Surface sediments from seven stations located in the seasonally stratified, frontal and mixed water regions in the Celtic and Irish seas have been analysed for their dinoflagellate cyst assemblages and dinosterol content. A total of 45 dinoflagellate cyst taxa have been identified and the assemblages related to surface and sediment conditions. Sediments from the mixed water region, at 30 m water depth, are characterised by a relatively low cyst concentration (∼2300 cysts/g dry weight) and high relative abundances of Lingulodinium machaerophorum accompanied by Spiniferites membranaceus, Brigantedinium spp. and Dubridinium caperatum. Assemblages from stratified and frontal water stations are dominated by Spiniferites ramosus associated with Operculodinium centrocarpum, Brigantedinium spp., cysts of Polykrikos schwartzii and Selenopemphix quanta. Ordination techniques performed on a restricted number of 35 taxa from the assemblages differentiated the stratified and frontal assemblages based on the abundance of the less abundant species Bitectatodinium tepikiense and Spiniferites elongatus. Among the environmental parameters (sea-surface temperature and salinity, stratification index, chlorophyll concentration and sediment grain-size classes), the seasonal stratification and sedimentological context, itself a function of tidal dynamics, explain most of the variance in the environmental conditions. These results indicate that dinoflagellate cyst analyses of shelf sediment records can be used to document the planktonic signal of seasonal stratification dynamics.     

EVIDENCE FOR ASEXUAL RESTING CYSTS IN THE LIFE CYCLE OF THE MARINE PERIDINOID DINOFLAGELLATE,SCRIPPSIELLA HANGOEI1

Scrippsiella hangoei (Schiller) Larsen is a peridinoid dinoflagellate that grows during winter and spring in the Baltic Sea. In culture this species formed round, smooth cysts when strains were mixed, indicating heterothallic sexuality and hypnozygote production. However, cysts of the same morphology were also formed in clonal strains exposed to slightly elevated temperature. To better understand the role of cysts in the life cycle of S. hangoei, cyst formation and dormancy were examined in culture experiments and the cellular DNA content of flagellate cells and cysts was compared in clonal and mixed strains using flow cytometry. S. hangoei exhibited a high rate of cyst formation in culture. Cysts produced in both clonal and mixed strain cultures were thick‐walled and underwent a dormancy period of 4 months before germinating. The S. hangoei flagellate cell population DNA distributions consisted of 1C, intermediate, and 2C DNA, indicative of respective eukaryotic cell cycle phases G1, S, and G2M. The majority (>95%) of cysts had a measured DNA content equivalent to the lower 1C DNA value, indicating a haploid nuclear phase and an asexual mode of cyst formation. A small percentage (<5%) of cysts produced in the mixed strain culture had 2C DNA, and thus could have been diploid zygotes. These findings represent the first measurements of dinoflagellate resting cyst DNA content, and provide the first quantitative evidence for dinoflagellate asexual resting cysts. Asexual resting cysts may be a more common feature of dinoflagellate life cycles than previously thought.     

HIGH ENCYSTMENT SUCCESS OF THE DINOFLAGELLATE SCRIPPSIELLA CF. LACHRYMOSA IN CULTURE EXPERIMENTS 1

Close to 100% encystment efficiency and a yield above 10⁵ cysts·mL ^{? 1} were routinely achieved in full strength f/2 medium‐based batch cultures (883 μM NO₃ ^? and 36 μM PO₄ ^{? 3}) of the marine dinoflagellate Scrippsiella cf. lachrymosa Lewis. Increases in cell density led to nutrient depletion in this enriched medium, which was the most likely cause for initiation of cyst formation. Lowering the concentration of either nutrient to 1/10 the initial levels decreased the encystment efficiency, whereas use of ammonium as the N source resulted in both low cell yield and low encystment efficiency. The mandatory dormancy period was ca. 60 days and was not affected by cold dark storage of the cysts. Cysts produced in the initial phase of sexual reproduction were relatively large (length 47 μm, width 31 μm) with a heavy calcareous cover. Cysts produced thereafter lacked apparent calcareous cover and were smaller (length 29 μm, width 19 μm). The decrease of cyst volume (by a factor of 0.24–0.4) suggested strong resource limitation during the course of encystment. However, after the mandatory dormancy period, germination success of the smaller cysts was higher (80%), compared with the larger cysts that had been produced initially (50%). Germling survival (74%) was independent of cyst type but was enhanced by higher nutrient concentration during incubation. The ratio of initial nutrient concentration in the medium to the cyst yield was used as a proxy to estimate the cellular nutrient quota. The conservative estimates of 9 pmol N·cyst ^{? 1} and 0.4 pmol P·cyst ^{? 1} obtained in this manner are at the low end of the range of previous published estimates for other dinoflagellate cysts. Given the high encystment observed in laboratory experiments, we have no reason to assume an inherently lower encystment success in dinoflagellate field populations. Our results do not challenge the low nutrient paradigm for dinoflagellate sexuality. We believe that the high encystment success and cyst yield of this particular species is at least partly due to its ability to achieve very high cell densities in cultures, which evidently leads to nutrient depletion even in f/2 medium.     

Encystment of Peridinium gatunense: occurrence, favourable environmental conditions and its role in the dinoflagellate life cycle in a subtropical lake

1. The abundance of cysts of the bloom‐forming dinoflagellate Peridinium gatunense in the sediments of Lake Kinneret and the effects of environmental conditions on encystment were studied in relation to bloom dynamics. Peak cyst formation coincided with the highest growth rate of the population, prior to bloom peak. 2. Peridinium cysts were counted in water and sediment corer samples from 2000 to 2003 and in archived sediment trap samples collected during 1993–94. The cyst data were examined in relation to ambient temperature and nutrient records, and revealed no direct correlation. 3. In laboratory encystment experiments with Peridinium cells collected from the lake, 0.2–3% of the vegetative cells encysted. Temperature, light and cell density had no significant effect on the percentage of encystment. 4. Cysts were always present in the lake sediments but their abundance in ‘non Peridinium’ years was much lower than after a massive bloom. Vegetative cells were always present in the water column after the collapse of the annual dinoflagellate bloom, potentially serving as the inoculum for the next bloom. We propose that the hardy cysts serve as an emergency ‘gene bank’ to initiate population build up following catastrophic die outs.