Morpho-molecular diversity and phylogeny of Bysmatrum (Dinophyceae) from the South China Sea and France

The dinoflagellate genus Bysmatrum encompasses five epibenthic or tide-pool species and has been characterized by separated anterior intercalary plates. In the present study, we obtained six strains of Bysmatrum from the South China Sea and French Atlantic coast by isolating single cells/cysts from plankton and sediment samples. All strains were examined with light microscopy and scanning electron microscopy. Based on morphological observations, three strains were identified as Bysmatrum subsalsum, characterized by the elongated and rectangular first and a hexagonal second anterior intercalary plate. They differ from each other in the number of sulcal lists and the configuration of the first anterior intercalary plate. One strain was identified as Bysmatrum gregarium and the other two as Bysmatrum granulosum. The cyst-theca relationship of B. subsalsum from the French Atlantic was established by incubation of the cyst, and the geochemical composition of the cyst wall was measured through micro-Fourier transform infrared spectroscopy. Bysmatrum subsalsum from Malaysia shows a bright red stigma in the sulcal area under light microscopy, which was confirmed with transmission electron microscopy: it was identified as a type B eyespot. Small subunit ribosomal DNA (SSU rDNA), partial large subunit ribosomal DNA (LSU rDNA) and internal transcribed spacer (ITS) sequences were obtained from all six strains. The maximum likelihood and Bayesian inference analysis based on concatenated SSU, ITS and LSU sequences revealed that Bysmatrum is monophyletic and nested within Peridiniales. Our strains of B. subsalsum form a new ribotype in the molecular phylogeny (designated as ribotype B). The genetic distance based on ITS sequences among Bysmatrum species ranged from 0.34 to 0.47 and those genetic distances at the intraspecific level of B. subsalsum could reach 0.41, supporting the possibility of hidden crypticity within B. subsalsum.     

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.     

Impact of grazing and bioturbation of marine benthic deposit feeders on dinoflagellate cysts

The impact of benthic deposit feeders on marine dinoflagellate cysts was studied by adding a concentrated natural Swedish cyst assemblage to sediment with different deposit feeders in replicate 4-l aquaria. The deposit feeders used were the bivalve Abra nitida, the echinoderm Amphiura filiformis, and the polychaetes Melinna cristata and Nereis diversicolor. These species occur naturally near the Swedish west coast and were selected to represent different ways of feeding. The results showed a significant relative decrease of unfossilizable cyst species; whereas, the common fossilizable species Lingulodinium polyedrum significantly increased in the cyst assemblage after grazing. This work suggests that differences in dinoflagellate cyst compositions can in part be caused by different animal grazing behaviors.     

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。     

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).     

ISLANDINIUM BREVISPINOSUM SP. NOV. (DINOFLAGELLATA), A NEW ORGANIC‐WALLED DINOFLAGELLATE CYST FROM MODERN ESTUARINE SEDIMENTS OF NEW ENGLAND (USA)1

Modern estuarine environments remain underexplored for dinoflagellate cysts, despite a rapidly increasing knowledge of cyst distributions in open marine sediments. A study of modern estuarine sediments in New England has revealed the presence of Islandinium brevispinosum sp. nov., a new organic‐walled dinoflagellate cyst that is locally common and probably of heterotrophic affinity. Resistance of this cyst to standard palynological processing indicates its geological preservability, although fossils are not yet known. Previously assigned species of the genus Islandinium are characteristic of polar and subpolar environments today and cold paleoenvironments in the Quaternary. The present record of I. brevispinosum extends the ecological and geographical range of this genus into the warm temperate zone, where I. brevispinosum occupies specific environments with reduced salinities and elevated nutrient levels.     

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 cyst biostratigraphy of the Opalinuston Formation (Middle Jurassic) in the Aalenian type area in southwest Germany and north Switzerland

Feist‐Burkhardt, S. & Pross, J. 2010: Dinoflagellate cyst biostratigraphy of the Opalinuston Formation (Middle Jurassic) in the Aalenian type area in southwest Germany and north Switzerland. Lethaia, Vol. 43, pp. 10–31. In order to provide a detailed dinoflagellate cyst stratigraphy of the Lower Aalenian Opalinuston Formation from the Aalenian type area, 68 samples from four boreholes and one outcrop section were analysed. The sample localities are Hausen an der Fils and Wittnau in southwest Germany, Weiach in north Switzerland and Mont Russelin in the Swiss Jura Mountains. Dinoflagellate cyst assemblages were recovered from the Late Toarcian Aalensis Zone to the Late Aalenian Murchisonae Zone. The samples yielded rich, well‐preserved and diverse assemblages with 51 dinoflagellate cyst taxa identified in total. The dinoflagellate cyst distribution data obtained from this study allow a high‐resolution biostratigraphical subdivision of the lowermost Middle Jurassic Opalinuston Formation into four palynostratigraphical units. First and last occurrences, acmes and consistent presence of the species Batiacasphaera sp. A, Evansia cf. granochagrinata, Kallosphaeridium praussii, Nannoceratopsis triangulata, Phallocysta? frommernensis and Wallodinium laganum were selected as the criteria for defining these units. The obtained high‐resolution palynostratigraphical scheme provides a basis for establishing and further refining early Middle Jurassic biostratigraphy in the Boreal and Tethyan realms. □Aalenian, biostratigraphy, dinoflagellate cysts, Germany, Jurassic, Switzerland, Toarcian.     

Dinoflagellate cyst records in recent sediments from Daya Bay, South China Sea

Nine sediment cores of 8–26 cm in length were collected from two basins of Daya Bay, the South China Sea, by Tokyo University Fisheries Oceanography Laboratory core sampler in August 2001 to investigate the distribution of dinoflagellate resting cysts. In the present study, 51 different cyst morphotypes representing 22 genera were identified from 65 sediment samples. Among them, there were 21 autotrophic species and 30 heterotrophic ones. Cyst species richness in each sample varied from 12 to 29, while the values of Shannon‐Weaver diversity index (H′) were between 0.15 and 4.13. There were an obvious increase in both species richness and values of H′in 2–6 cm sediments. Cyst concentrations varied from 154 to 113 483 cysts per gram dry weight sediment, and were much higher in upper sediments. Scrippsiella trochoidea was the most dominant cyst type, which took up over 90% of cyst assemblages in the upper sediments. The abrupt increase of S. trochoidea cysts in the surface sediments reflected the bloom of this species in Daya Bay in 2000. The results from cyst assemblages showed some trend of changes in water quality in this area, and indicated a typical type of pollution caused by cultural eutrophication, which started in the 1980s and greatly accelerated in the middle of 1990s. Cysts of Alexandrium, mainly those of Alexandrium catenella and Alexandrium tamarense complex, occurred frequently and abundantly in this area, with the highest concentration and relative frequency of 503 cysts per gram dry weight sediment and 22.3%, respectively. The high abundance of Alexandrium cysts provided rich ‘seed bed’ for Alexandrium blooms and was also an important source of paralytic shellfish poisoning toxins, especially in winter.     

Benthic flattened cells of the phylogenetically related marine dinoflagellates Protoceratium reticulatum and Ceratocorys mariaovidiorum (Gonyaulacales): a new type of cyst?

A planktonic‐benthic relationship has been described for many dinoflagellate species as part of their ecological strategy to overcome highly variable aquatic environments. Here, the phylogenetically and morphologically related marine dinoflagellates Protoceratium reticulatum and Ceratocorys mariaovidiorum were studied in relation to an unknown benthic life form. In vivo and fixed samples from cultures were analyzed in detail by light and scanning electron microscopy. In both species, a cell type with a morphology different from that of vegetative cells was observed in cultures grown until stationary phase. This cell type was always benthic, swimming sporadically only when it was disturbed. Its main feature included a strong dorsoventral compression. These cells originated from vegetative cells whose protoplasm underwent a progressive flattening, resulting in a gradual detachment of the reticulate and thick thecal plates and the formation of very thin non‐reticulated new plates with pores. When returned to fresh full‐strength medium, the cells recovered their spherical vegetative‐like morphology, including new reticulated thick plates and subsequent cell divisions. The kinetics of flattened cell formation showed that in both species, this cell type increased exponentially until the onset of the culture stationary phase and then decreased. The results of this study are discussed in the context of the planktonic–benthic coupling in dinoflagellate life cycles, including those newly appreciated to be well adapted to the benthic environment.     

Seasonal distribution of dinoflagellate resting cysts in surface sediments from Changjiang River Estuary

In order to understand the distribution of dinoflagellate cysts, surface sediments were collected from 15 stations in Changjiang River Estuary from 122°E to 123.5°E and from 29°N to 32°N in four cruises from May 2002 to February 2003. In the present study, 38 different cyst morphotypes representing 21 genera and 6 groups were identified, while 1 type was not identified into genus level. Species number and cell density of dinoflagellate cysts ranged from 10 to 25 species and from 12 to 587 per gram of dry weight, respectively. There were no obvious differences in cyst composition and density among seasons. However, the highest cyst species number and density were recorded in summer and winter, respectively. Cysts of heterotrophic dinoflagellates, which held 55.7% of the overall cyst density averagely, dominated cyst assemblages. Cyst density and species number increased from the west to the east, from the north to the south within the study area. Cysts of toxic dinoflagellates Alexandrium cat‐enella and Alexandrium tamarense complex distributed widely and were observed in almost all stations, with the maximum cell density of 81 per gram of dry weight.     

Species diversity and abundance of dinoflagellate resting cysts seven months after a bloom of Alexandrium catenella in two contrasting coastal systems of the Chilean Inland Sea

In Chile, 90% of the fish farms and major natural shellfish beds are located in the region surrounding the Inland Sea, where over the last few decades harmful phytoplankton blooms have often been observed. The onset and recurrence of bloom events are often related to the resuspension and germination of resting cysts that have accumulated in the sediments. The degree of cyst settling, accumulation and germination is highly variable between areas and depends on physical and environmental factors. To learn how differences in oceanographic exposure, amount of river runoff and bathymetry affect dinoflagellate cyst deposition, we examined the diversity and abundance of dinoflagellate resting cysts from two hydrographically contrasting coastal areas (oceanic Guaitecas Archipelago and estuarine Pitipalena Fjord) of the Chilean Inland Sea in September 2006, seven months after a bloom of Alexandrium catenella, a producer of paralytic shellfish toxin. Cyst species diversity consisted of 18 taxa, including A. catenella and the noxious species Protoceratium reticulatum, both of which have caused blooms in the study area. Our results revealed significant differences between the two study sites in terms of the abundance and diversity of resting cysts, suggesting that in the specific case of A. catenella, only Guaitecas stations have potential for cyst accumulation and successful growth of cells. However, there was no evidence of long-term resting cyst beds of A. catenella at either study site.     

Viability of phytoplankton resting stages in the sediments of a coastal Swedish fjord

Viable diatom and dinoflagellate resting stages were recovered from sediments in Koljö Fjord on the west coast of Sweden. To determine the maximum survival time of buried resting stages, samples from sediment depths down to 50?cm were incubated at temperatures of 3, 10 and 18?°C. Sediment cores were dated by ²¹⁰Pb and the age of samples containing viable resting stages was determined using the constant rate of supply model. Dilution cultures of surface sediments allowed semiquantitative estimates of the potential seed bank. Dinoflagellate cysts from species such as Diplopsalis sp., Gymnodinium nolleri, Oblea rotunda and Protoceratium reticulatum were viable down to 15?cm depth, or 37 years old. Spores and resting cells of the diatoms Chaetoceros spp., Detonula confervacea and Skeletonema costatum were viable to over 40?cm depth, and may have been buried for many decades. The seed bank of living resting stages in surficial sediments was found to be rich (c. 57000 diatom resting stages g^?1 wet weight and c. 200 dinoflagellate cysts g^?1 wet weight), and the percentage of viable resting stages was higher for spore- and cyst-forming species. The oxygen-deficient sediments in Koljö Fjord appear to be a natural conservator of cell viability, a condition not easily simulated in laboratory studies. These results are ecologically important since spores and cysts are a repository of genetic material able to repopulate waters if resuspended and exposed to suitable light, temperature and nutrients.     

中国渤海海域甲藻孢囊的种类多样性和生态地理分布

在渤海湾24个站位采集0–10 cm表层沉积物144份样品, 进行甲藻孢囊的种类多样性和生态地理分布研究, 并分析渤海海域水体的富营养化趋势。在渤海海域共采集鉴定出6类32种甲藻孢囊, 包括原多甲藻类孢囊11种、裸甲藻类孢囊6种、膝沟藻类孢囊7种、钙质类孢囊5种、翼藻类孢囊2种及Tuberculodinioid类孢囊1种。其中, 自养型甲藻孢囊17种, 优势种为锥状斯氏藻(Scrippsiella trochoidea)孢囊, 丰富的孢囊库为锥状斯氏藻赤潮提供种源, 可作为该海域发生锥状斯氏藻赤潮的参考依据; 异养型甲藻孢囊15种, 优势种为无纹多沟藻(Polykrikos schwartzii)和锥形原多甲藻(Protoperidinium conicum)孢囊; 发现1种产麻痹性贝类毒素的孢囊--塔玛亚历山大藻(Alexandrium tamarense)孢囊, 但是仅在S6、S8和S14三个站位发现, 且孢囊数量较少。渤海甲藻孢囊组成与世界其它海域相似, 异养型甲藻孢囊种类丰富, 提示渤海海域富营养化程度较高。每个站位沉积物样品分为3层(每层约3 cm), 春夏两季表层、中层和底层之间孢囊组成差异不大, 平均种类数为5.58–7.29, 孢囊的平均密度范围在58.82–103.57 cysts·g^–1 DW之间; 孢囊种类丰富的站点, 孢囊密度也较高, 如S9、S14站位。对渤海海区表层沉积物中甲藻孢囊的生物多样性进行了统计分析, 发现表、中、底3层甲藻孢囊的平均种类多样性指数在春季(5月)分别为1.69、1.61和1.52; 略低于夏季(8月)的1.83、1.89和1.60。     

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.     

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.     

Protoperidinium tricingulatum sp. nov. (Dinophyceae), a new motile form of a round,brown, and spiny dinoflagellate cyst

A small, broadly ovoidal and heterotrophic dinoflagellate containing round, brownish, and spiny cyst was found in the water column of Huibertsplaat in the Wadden Sea off the coast of the Netherlands. This dinoflagellate had these conspicuous morphological characters: a five‐sided first apical plate (1′), only three cingular plates, and an extremely small first antapical plate. Based on these morphological features, Protoperidinium tricingulatum Kawami, vanWezel, Koeman et Matsuoka is described as a new species. The flagellar pore of P. tricingulatum is covered with a small fin, which rises from the left side of the right sulcal plate to the large V‐shaped posterior sulcal plate. This feature suggests that P. tricingulatum is assigned to the Abé's Monovela Group. The cyst stage of P. tricingulatum was positively linked to the vegetative stage by comparison of the ribosomal 5.8S rDNA, internal transcribed spacers (ITS1 and ITS2). Living cysts of P. tricingulatum are round, brownish, and covered with many slender spines bearing capitate or cauliforate distal ends. The cyst also possesses a theropylic archeopyle formed by a slit corresponding to parasutures between three apical and two apical intercaraly plates. These morphological characters indicate that this species is morphologically related to two dinoflagellate cyst‐genera Islandinium and Echinidinium.     

Dinoflagellate cyst distribution along a shelf to slope transect of an oligotrophic tropical sea (Sunda Shelf, South China Sea)

From 51 surface samples collected along a shelf to slope transect of the Sunda Shelf, South China Sea, 36 taxa of organic‐walled dinoflagellate cysts are identified. Oligotrophic tropical shelf assemblages on the Sunda Shelf are dominated by gonyaulacoids such as Spiniferites species, Operculodinium centmcarpum and Operculodinium israelianum. Concentrations of dinoflagellate cysts in the shelf sediments are generally low and correlate well with the content of fine‐grained (clay and silt fraction) sediments. Detailed comparisons of sediment grain‐size distributions to concentrations of dominant dinoflagellate taxa (Spiniferites species, 0. centmcarpum and 0. israelianum) in the shelf sediments indicate that these taxa behave in water like sediment particles with size range φ 5.75–6.25 (13–18 μm). In contrast, slope assemblages in fine‐grained sediments are dominated by protoperidinioids. This may reflect higher nutrient availability as a result of weak winter upwelling. The concentrations of dinoflagellate cysts in the shelf sediments are mainly controlled by transport and winnowing processes and are probably not representative of surface water conditions.     

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.