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.     

Spatial distribution of calcareous dinoflagellate cysts in surface sediments of the Atlantic Ocean between 13 degrees N and 36 degrees S

To enhance the limited information available about the palaeo-ecological significance of calcareous dinoflagellates, we have studied their lateral distribution in surface sediments of the equatorial and south Atlantic between 13 degrees N and 36 degrees S. Calcareous dinoflagellate cysts appear to be widely distributed throughout the studied area. In the surface sediments, concentrations (cyst per gram dry sediment) of the vegetative stage Thoracosphaera heimii are generally higher than that of the (presumably) calcareous resting cysts. Distribution patterns in surface sediments of Orthopithonella granifera (Fütterer) Keupp and Versteegh, Rhabdothorax spp. Kamptner., Sphaerodinella albatrosiana (Kamptner) Keupp and Versteegh S. albatrosiana praratabulated, Sphaerodinella tuberosa var. 1 (Kamptner) Keupp and Versteegh and S. tuberosa var. 2 and the ratios between these species have been compared with temperature, salinity, density and stratification gradients in the upper water column. Rhabdothorax spp. is characteristically present in sediments of more temperate regions characterized by high seasonality. Dinoflagellates producing these cysts are able to tolerate high nutrient concentrations, and mixing of the water column. S. albatrosiana is abundant in regions characterized by high sea surface temperatures and oligotrophic surface water conditions. In contrast, the distribution of S. tuberosa var. 2 is negatively related to temperature. The other cyst species did not show a characteristic pattern in relation to the studied environmental gradients.The ratio of Sphaerodinella tuberosa var. 2 to Orthopithonella granifera can be used for reconstructing the presence of stratification in the upper 50m of the water column, whereas the ratios of S. tuberosa var. 2 to Sphaerodinella albatrosiana and of O. granifera to Rhabdothorax spp. might be used for palaeotemperature reconstructions. Calcareous dinoflagellate cysts are abundant in oligotrophic areas and may be useful for the reconstruction of palaeoenvironmental 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.     

Calcareous dinoflagellate cyst distributions in surface sediments from upwelling areas off NW Africa, and their relationships with environmental parameters of the upper water column

Only very few studies have focussed on the spatial distribution and ecology of calcareous dinoflagellate cysts (dinocysts) in upwelling areas. Here, distributions of individual calcareous dinocyst species in surface sediment samples from the coastal upwelling zone off NW Africa and their relationships with known environmental parameters of the (sub-)surface waters have been analysed in order to enhance our knowledge on their modern distribution patterns and to determine the ecological significance and palaeoenvironmental reconstruction potential of each calcareous dinocyst species within this exemplary high-productivity region. In addition to calculating relative cyst abundances, well-constrained sedimentation rates have allowed the calculation of dinocyst accumulation rates (cysts cm^− 2 ka^− 1) for most of the surface sample positions, thus providing a much more accurate estimation of actual calcareous dinocyst production in the upper water column than could be warranted in similar studies so far. Distinct differences in the geographic distribution patterns of nine species were observed. In general, high accumulation rates of calcareous dinocysts were found around and south of 29°N and offshore the westernmost Canary Island La Palma, with slightly lower accumulation rates in the upwelling-influenced areas off Cape Yubi and in the upwelling filament area off Cape Ghir. Multivariate ordination techniques were applied in order to compare the cyst accumulation rates of individual species with physical parameters (temperature, salinity, density, mixed layer depth) and the trophic state (nitrate, phosphate and chlorophyll concentrations) of the overlying (sub-)surface waters. All species were found to relate significantly to one or more of the environmental parameters, partly confirming previous results on cyst ecology but also providing new information which will be useful for future palaeoenvironmental reconstructions within upwelling areas.     

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.     

Organic-walled dinoflagellate cysts in western equatorial Atlantic surface sediments: distributions and their relation to environment

In contrast to the wide range of studies carried out in temperate and high-latitude oceanic regions, only a few studies have focused on recent and Holocene organic-walled dinoflagellate cyst assemblages from the tropics. This information is, however, essential for fully understanding the ability of species to adapt to different oceanographic regimes, and ultimately their potential application to local and regional palaeoenvironmental and palaeoceanographic reconstructions. Surface sediment samples of the western equatorial Atlantic Ocean north of Brazil, an area greatly influenced by Amazon River discharge waters, were therefore analysed in detail for their organic-walled dinoflagellate cyst content. A diverse association of 43 taxa was identified, and large differences in cyst distribution were observed. The cyst thanatocoenosis in bottom sediments reflects the seasonal advection of Amazon River discharge water through the Guyana Current and the North Equatorial Countercurrent well into the North Atlantic. To establish potential links between cyst distribution and the environmental conditions of the upper water column, distribution patterns were compared with mean temperature, salinity, density and stratification gradients within the upper water column (0-100m) over different times of the year, using correspondence analysis and canonical correspondence analysis. The analyses show that differences in these parameters only play a subordinate role in determining species distribution. Instead, nutrient availability, or related factors, dominates the distribution pattern. The only possible indicators of slightly reduced salinities are Trinovantedinium applanatum and Lingulodinium machaerophorum. Four assemblage groups of cyst taxa with similar environmental affinities related to specific water masses/currents can be distinguished and have potential for palaeoenvironmental reconstruction.     

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.     

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。     

Pithonelloid wall-type of the Late Cretaceous calcareous dinoflagellate cyst genus Tetratropis

The late Cretaceous calcareous dinoflagellate genus Tetratropis features both a pithonelloid wall-type (evenly inclined wall-components, proven here by a polarisation optical revision) and a peridinialean paratabulation strongly suggesting a dinoflagellate origin of at least part of the Pithonelloideae. This affinity with dinoflagellates sheds more light on the palaeoecology of the Pithonelloideae (commonly termed “calcispheres”), which are characteristic of the middle to Late Cretaceous. The very short-term stratigraphic occurrence of all Tetratropis species is comparable to the distribution pattern of other calcareous dinoflagellate cyst species with a distinctive paratabulation and is thought to reflect a narrow palaeoecological niche. Tetratropis species can be interpreted either as paratabulated morphotypes of otherwise atabulate Pithonelloideae formed under exceptional palaeoenvironmental conditions or as invaders from a highly specific palaeoecological niche during short-term palaeoceanographic events probably related to the initiation of the Late Cretaceous global cooling.     

广东大亚湾甲藻孢囊及其与锥状斯氏藻赤潮的关系

1999年12月至2001年1月,在大亚湾澳头海域用沉积物捕捉器(Sediment trap)及TFO重力采泥器对甲藻孢囊进行每月一次的周年监测,并同时研究了浮游植物的季节变化.结果显示,晚秋孢囊形成率最高(3.48105 cysts/m2d),冬季形成率较低,年平均为1.28105 cysts/m2d.锥状斯氏藻(Scrippsiella trochoidea)是大亚湾沉积物孢囊中的绝对优势种,除个别季节外,其形成率一般占孢囊总形成率的50%以上.2000年8月至9月,该海域发生了一次较大规模的锥状斯氏藻赤潮,最高细胞密度达4.0104 cells/mL.赤潮中后期,锥状斯氏藻孢囊包括暂时性孢囊和休眠孢囊大量形成,孢囊的形成减少了水体中营养细胞数量,是赤潮消退原因之一.       

Modern dinoflagellate cysts in hypertrophic coastal waters of Tokyo Bay, Japan

A survey of dinoflagellate resting cysts in surface sedimentsamples was carried out in Tokyo Bay, Japan, to document theirhorizontal distribution. At least 21 different cyst types werefound. Dominant cyst types allowed the recognition of assemblageswhich form three different dinoflagellate cyst communities:the innermost part of the Bay, the central area and the moutharea. In all stations in Tokyo Bay, heterotrophic dinoflagellatecysts always occupied more than half of the cyst populations.Cysts of Polykrikos schwartzii/kofoidii are the most abundantheterotrophic species. These assemblages may reflect highlynutrient-enriched (hypertrophic) and turbulent water conditions.Among the cyst types found were probable ellipsoidal cysts ofAlexandrium tamarense. This is the first record of toxic Alexandriumspecies cysts in Tokyo Bay sediments.     

The (palaeo) environmental significance of modern calcareous dinoflagellate cysts: a review

Increased interest in the environmental applicability of calcareous dinoflagellate cysts for palaeo-environmental studies arose in the last two decades, when it was discovered that they constitute a widespread and abundant group within the calcareous phytoplankton. As primary producers they are directly influenced by environmental Parameters of the surrounding water masses, and therefore extremely suitable for detailed palaeo-environmental and -oceanographical reconstructions. During the last years, detailed investigations on their distribution in surface sediments of the South Atlantic Ocean, the Mediterranean Sea and the Arabian Sea were carried out, in order to reveal which environmental parameters influence their distribution. This, in turn, formed the base for palaeo-environmental and -oceanographical reconstructions. The compilation of the available data on calcareous dinoflagellate ecology, morphology, taxonomy and palaeo-environmental application presented here demonstrates their ample application potential as palaeo-environmental tools.      

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.     

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.     

Coupling planktonic and benthic shifts during a bloom of Alexandrium catenella in southern Chile: Implications for bloom dynamics and recurrence

Cell abundances and distributions of Alexandrium catenella resting cysts in recent sediments were studied along time at two locations in the Chilean Inland Sea exposed to different oceanographic conditions: Low Bay, which is much more open to the ocean than the more interior and protected Ovalada Island. The bloom began in interior areas but maximum cyst concentrations were recorded in locations more open to the ocean, at the end of the Moraleda channel. Our results showed a time lapse of around 3 months from the bloom peak (planktonic population) until the number of resting cysts in the sediments reached a maximum. Three months later, less than 10% of the A. catenella cysts remained in the sediments. Maximum cyst numbers in the water column occurred one month after the planktonic peak, when no cells were present. The dinoflagellate assemblage at both study sites was dominated by heterotrophic cysts, except during the A. catenella bloom. CCA analyses of species composition and environmental factors indicated that the frequency of A. catenella blooms was associated with low temperatures, but not with salinity, chlorophyll a concentration, and predator presence (measured as clam biomass). However, resting cyst distribution was only related to cell abundance and location. The occurrence of A. catenella cysts was also associated with that of cysts from the toxic species Protoceratium reticulatum. By shedding light on the ecological requirements of A. catenella blooms, our observations support the relevance of encystment as a mechanism of bloom termination and show a very fast depletion of cysts from the sediments (<3 months), which suggest a small role for resting cyst deposits in the recurrence of A. catenella blooms in this area.     

First report of fossilized cysts produced by the benthic Bysmatrum subsalsum (Dinophyceae) from a shallow Mexican lagoon in the Gulf of Mexico

Cysts belonging to the benthic dinoflagellate Bysmatrum subsalsum were recovered from palynologically treated sediments collected in the Alvarado Lagoon (southwestern Gulf of Mexico). The cysts are proximate, reflecting the features of the parent thecal stage, and their autofluorescence implies a dinosporin composition similar to the cyst walls of phototrophic species. This finding is important for our understanding of B. subsalsum life cycle transitions and ecology. Encystment may play an important role in the bloom dynamics of this species as it can enable the formation of a sediment cyst bank that allows reinoculation of the water column when conditions become favorable. This is the first report of a fossilized cyst produced by a benthic dinoflagellate recovered from sub‐recent sediments.     

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.     

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).     

CONTROL OF GERMINATION OF ALEXANDRIUM TAMARENSE (DINOPHYCEAE) CYSTS FROM THE LOWER ST. LAWRENCE ESTUARY (CANADA)

Cysts of the toxic dinoflagellate Alexandrium tamarense (Lebour) Balech 1992 from the lower St. Lawrence estuary were used in a test of the following hypotheses: (1) cyst germination is triggered by a change in temperature, and (2) germination rate varies throughout the year and is controlled by a circannual internal biological clock. Results show that cyst germination was not affected significantly by temperature of incubation over the range 1°–16° C, and light showed no significant stimulation of germination. This is supported by the lack of effect of cyst incubation conditions during evaluation of the seasonal changes in germination rate (two temperatures: 4° and 15° C, and two light conditions: darkness and 150 μmol photons·m^?2·s^?1). Thus, direct environmental control through short-term increases in temperature and exposure to light has no effect on the germination of the cysts tested. The rate of germination, observed monthly over a 16-month period, showed low germination (<20%) over most of the period tested, except for a maximum reaching more than 50% germination in August to October of the second year of the experiment. This pattern was observed for cysts both from monthly field collections and from laboratory-stored cysts kept under constant environmental conditions (4° C, in the dark). The peak in germination observed under constant environmental conditions (in the laboratory), the almost coincidental increase in cyst germination observed for the field-collected cysts, and the absence of effects of temperature and light during incubation could be explained either by a temperature-controlled cyst maturation period (the time-temperature hypothesis of Huber and Nipkow 1923) or by the presence of an internal biological clock. However, the large decline in the rate of germination 2 months after the maximum provides strong support for the biological clock hypothesis. The ca. 12-month maturation (dormancy) period observed for the laboratory-stored cysts is the longest reported for this species to our knowledge; this might be related to the low storage temperature (4° C), which is close to bottom temperatures generally encountered in this environment (0° to 6° C). Similar field and laboratory storage temperatures could explain the coincidental increase in germination rate in the fall of the second year if cyst maturation is controlled by temperature. A fraction of the laboratory-stored cysts did not follow a rhythmic pattern: A rather constant germination rate of about 20% was observed throughout the year. This continuous germination of likely mature cysts may supplement the local blooms of this toxic dinoflagellate, as these often occur earlier than peak germination observed in late summer. It seems that two cyst germination strategies are present in the St. Lawrence: continuous germination after cyst maturation, with temperature controlling the length of the maturation period, and germination controlled by a circannual internal rhythm.