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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.

How important are refugia for plankton biogeography? Here for the first time we report living cysts of the fossil dinoflagellate Dapsilidinium pastielsii from Southeast Asia: Shioya Bay (Okinawa, Japan), Koror (Palau), Ambon (Indonesia), East Vietnam Sea (Vietnam), and Masinloc (the Philippines). This species, thought to have become extinct in the early Pleistocene, is the last survivor of a major early Cenozoic lineage. Its disappearance from the Atlantic following the early Pleistocene implies cooling, and the discovery of living D. pastielsii in the Indo-Pacific warm pool suggests that this unique environment with stable temperatures served as an important refuge for thermophilic dinoflagellates with a >50 m.y. lineage. This is the first record of a refugium plankton species within the Indo-Pacific warm pool.

Recent studies have correlated dinoflagellate resting cyst morphology to salinity and density variations in the water column, suggesting that morphology can be used for paleoceanographic reconstructions. However, the univariate statistics used by these studies are appropriate only where morphology is related to a single variable. Density is a function of salinity and temperature, so more advanced statistical methods are needed to understand which parameters affect morphology. In this study based on surface (coretop) sediments, a set of environmental variables (sea-surface salinity, temperature, nitrate, and phosphate) was simultaneously correlated to morphological variations seen in resting cysts produced by the dinoflagellate Protoceratium reticulatum (= Operculodinium centrocarpum sensu Wall and Dale). Approximately 3200 measurements were obtained from the North Atlantic Ocean and used to generate a working model based on the Akaike information criterion. Hierarchical partitioning was then applied to establish the independent and joint effects for each predictor variable. Results from these analyses showed that while salinity constitutes the dominant variable affecting process length in the cysts of P. reticulatum in the North Atlantic, it is not the sole explanatory variable and that multicollinearity exists. Temperature and nutrients also showed a significant relationship to the morphology, requiring multiple regression to construct a representative model. The applicability of the North Atlantic working model was finally evaluated by comparing the results to data from the Mediterranean, Marmara, and Black seas, and Baltic–Kattegat–Skagerrak estuarine system. This comparison showed regional differences in morphological–environmental correlation. While salinity constitutes the most important explanatory factor in both the North Atlantic and Baltic–Skagerrak system, this is not so for the Mediterranean–Black Sea region where temperature is the dominant variable. It is concluded that a predictive salinity model based on P. reticulatum cyst morphology has at best a regional application.

The number of cingular plates has been used to differentiate Protoperidinium from Peridinium and related genera. Protoperidinium is characterized by the presence of three cingular plates plus a transitional plate (3C+t). However, many Protoperidinium species have been described that exhibit different cingular plate tabulations. How these species should be classified within the genus remains unclear. To address this question, the phylogenetic relationship of four Protoperidinium species, with three or four cingular plates and lacking a transitional plate, were examined in relationship to other Protoperidinium species. These four species were germinated from cysts deposited in surface sediments collected from the East China Sea, the Bohai Sea and the Yellow Sea. Three of the isolated species, P.?tricingulatum, P.?americanum and P.?parthenopes, were described previously. The fourth is here described as P.?haizhouense sp. nov. with the plate formula Po, X, 4', 3a, 7'', 3C, 6S, 5''', 2''''. Differences in the cyst stages of these four species, which can be taxonomically informative, were compared. Partial large subunit ribosomal DNA sequences were obtained by single-cell polymerase chain reaction. Maximum-likelihood and Bayesian inference showed that these four species, P.?fukuyoi and Islandinium minutum form a monophyletic clade with maximal support. The genus as a whole, however, appeared polyphyletic. Our results suggest that the presence/absence of a transitional plate is significant in the phylogeny of Protoperidinium.

Recent studies have correlated dinoflagellate resting cyst morphology to salinity and density variations in the water column, suggesting that morphology can be used for paleoceanographic reconstructions. However, the univariate statistics used by these studies are appropriate only where morphology is related to a single variable. Density is a function of salinity and temperature, so more advanced statistical methods are needed to understand which parameters affect morphology. In this study based on surface (coretop) sediments, a set of environmental variables (sea-surface salinity, temperature, nitrate, and phosphate) was simultaneously correlated to morphological variations seen in resting cysts produced by the dinoflagellate Protoceratium reticulatum (=Operculodinium centrocarpum sensu Wall and Dale). Approximately 3200 measurements were obtained from the North Atlantic Ocean and used to generate a working model based on the Akaike information criterion. Hierarchical partitioning was then applied to establish the independent and joint effects for each predictor variable. Results from these analyses showed that while salinity constitutes the dominant variable affecting process length in the cysts of P. reticulatum in the North Atlantic, it is not the sole explanatory variable and that multicollinearity exists. Temperature and nutrients also showed a significant relationship to the morphology, requiring multiple regression to construct a representative model. The applicability of the North Atlantic working model was finally evaluated by comparing the results to data from the Mediterranean, Marmara, and Black seas, and Baltic-Kattegat-Skagerrak estuarine system. This comparison showed regional differences in morphological-environmental correlation. While salinity constitutes the most important explanatory factor in both the North Atlantic and Baltic-Skagerrak system, this is not so for the Mediterranean-Black Sea region where temperature is the dominant variable. It is concluded that a predictive salinity model based on P. reticulatum cyst morphology has at best a regional application.

The study investigates the morphological variability of the dinoflagellate cyst Operculodinium centrocarpum (resting cyst of Protoceratium reticulatum) in core-top samples distributed over the Southern Hemisphere and the tropics in relation to sea-surface temperature (SST) and sea-surface salinity (SSS) at the corresponding sites. The process lengths show a moderate inverse relationship to summer SST (sSST) (R²=0.44) and sSSS/sSST (R²=0.4), however, lateral transport of cysts probably produced noise in the plots. After excluding tropical and Southern Hemisphere sites considered to have been affected by long distance lateral transport, the relationship between process length and density follows the equation sD=0.8422x+1016.9 (R²=0.55) with a Root Mean Square Error=0.63 kg m-3, while the negative correlation with sSST increases up to R²=0.79. Next to salinity, this study thus highlights the importance of a second factor, temperature, affecting process length in the topics and the Southern Hemisphere oceans.

Reconstruction of salinity in the Holocene Black Sea has been an ongoing debate over the past four decades. Here we calibrate summer surface water salinity in the Black Sea, Sea of Azov and Caspian Sea with the process length of the dinoflagellate cyst Lingulodinium machaerophorum. We then apply this calibration to make a regional reconstruction of paleosalinity in the Black Sea, calculated by averaging out process length variation observed at four core sites from the Black Sea with high sedimentation rates and dated by multiple mollusk shell ages. Results show a very gradual change of salinity from ~14 ± 0.91 psu around 9.9 cal ka BP to a minimum ~12.3 ± 0.91 psu around 8.5 cal ka BP, reaching current salinities of ~17.1 ± 0.91 psu around 4.1 cal ka BP. The resolution of our sampling is about 250 years, and it fails to reveal a catastrophic salinization event at ~9.14 cal ka BP advocated by other researchers. The dinoflagellate cyst salinity-proxy does not record large Holocene salinity fluctuations, and after early Holocene freshening, it shows correspondence to the regional sea-level curve of Brückner et al. (2010) derived from Balabanov (2007).

Lycopodium clavatum tablets are commonly added as a spike to determine dinoflagellate cyst concentrations in sediments. In this study we investigate the effects of different processing techniques on dinoflagellate cyst concentrations using well-mixed sediment samples from Saanich Inlet, British Columbia, Canada. At the onset of any dinoflagellate cyst investigation, we suggest following the recommendations of Maher (1981) to experimentally adjust the sample size to obtain a ratio close to ~ 2 of dinoflagellate cysts counted to Lycopodium spores counted, in order to obtain reproducible concentrations. Results further show that both oven-drying at ~ 45 °C and freeze-drying are viable, non-destructive techniques yielding reproducible results. Use of warm HCl (40–60 °C) for a short time (30 min) is harmless, whereas treatment with warm HF (40–60 °C) affects the reproducibility of the concentrations. Pre-sieving can result in loss of cysts and/or spike but this can be easily monitored by checking the residue. Perforated metal sieves show more consistent results than the Nitex nylon meshes. The use of 30 second sonication does not affect the reproducibility, and is advised to remove amorphous organic matter. Adding the Lycopodium spike at the end of preparation yields consistently lower concentrations, which were usually not reproducible, suggesting noticeable losses of Lycopodium spores during processing if the Lycopodium spores are added at the beginning. This method can be considered a viable alternative, but the discrepancy should be taken into account.

Results are presented from a regional comparison of average process length variation in cysts of Protoceratium reticulatum and Lingulodinium polyedrum, extracted from surface sediments in the Skagerrak–Kattegat–Baltic estuarine system, with the environmental variables of seawater temperature and salinity. Although too few cysts of Lingulodinium polyedrum were recovered from the sediments to make reliable correlations, cysts of Protoceratium reticulatum were well represented, and average process length was correlated significantly with both salinity and temperature. Owing to dominant summer surface production, and regional covariation between salinity and density, we propose the use of the significant correlation with summer sea surface salinity (SSSsummer) by the equation SSSsummer=3.16 × average process length -0.84 (R2=0.8). Application of this equation down-core in Limfjord (northern Denmark) shows its usefulness as a regional palaeosalinity proxy.

Detailed palynological studies in the northeast (NE) Pacific, Strait of Georgia (BC, Canada), southeast (SE) Pacific and northwest Pacific (Dongdo Bay, South Korea) resulted in the recognition of the new dinoflagellate cyst species Selenopemphix undulata sp. nov. This species is restricted to cool temperate to sub-polar climate zones, where it is found in highest relative abundances in highly productive non-to reduced upwelling regions with an annual mean sea-surface temperature (aSST) below 16°C and an annual mean sea-surface salinity (aSSS) between 20 and 35 psu. Those observations are in agreement with the late Quaternary fossil records from Santa Barbara Basin (ODP 893: 34°N) and offshore Chile (ODP 1233: 41°S), where this species thrived during the last glacial. This period was characterised by high nutrient availability and the absence of species favouring upwelling conditions. The indirect dependence of S. undulata sp. nov. abundances on nutrient availability during reduced or non-upwelling periods is expressed by the synchronous fluctuations with diatom abundances, since the distribution and growth rates of the latter are directly related with the availability of macronutrients in the surface waters.

Absolute abundances (concentrations) of dinoflagellate cysts are often determined through the addition of Lycopodium clavatum marker-grains as a spike to a sample before palynological processing. An inter-laboratory calibration exercise was set up in order to test the comparability of results obtained in different laboratories, each using its own preparation method. Each of the 23 laboratories received the same amount of homogenized splits of four Quaternary sediment samples. The samples originate from different localities and consisted of a variety of lithologies. Dinoflagellate cysts were extracted and counted, and relative and absolute abundances were calculated. The relative abundances proved to be fairly reproducible, notwithstanding a need for taxonomic calibration. By contrast, excessive loss of Lycopodium spores during sample preparation resulted in non-reproducibility of absolute abundances. Use of oxidation, KOH, warm acids, acetolysis, mesh sizes larger than 15 µm and long ultrasonication (> 1 min) must be avoided to determine reproducible absolute abundances. The results of this work therefore indicate that the dinoflagellate cyst worker should make a choice between using the proposed standard method which circumvents critical steps, adding Lycopodium tablets at the end of the preparation and using an alternative method.

A biometrical analysis of the dinoflagellate cyst Lingulodinium machaerophorum [Deflandre, G., Cookson, I.C., 1955. Fossil microplankton from Australia late Mesozoic and Tertiary sediments. Australian journal of Marine and Freshwater Research 6: 242–313.] Wall, 1967 in 144 globally distributed surface sediment samples revealed that the average process length is related to summer salinity and temperature at a water depth of 30 m by the equation (salinity/temperature) = (0.078⁎average process length + 0.534) with R2 = 0.69. This relationship can be used to reconstruct palaeosalinities, albeit with caution. The particular ecological window can be associated with known distributions of the corresponding motile stage Lingulodinium polyedrum (Stein) Dodge, 1989. Confocal laser microscopy showed that the average process length is positively related to the average distance between process bases (R2 = 0.78), and negatively related to the number of processes (R2 = 0.65). These results document the existence of two end members in cyst formation: one with many short, densely distributed processes and one with a few, long, widely spaced processes, which can be respectively related to low and high salinity/temperature ratios. Obstruction during formation of the cysts causes anomalous distributions of the processes. From a biological perspective, processes function to facilitate sinking of the cysts through clustering.

Dinoflagellate cysts and other palynomorphs were studied from ODP Hole 1002C in the Cariaco Basin over the past 30 000 years. The assemblage shifts between a dominance of heterotrophic dinoflagellate cysts (mainly Brigantedinium spp., Lejeunecysta spp., Selenopemphix nephroides and Stelladinium reidii) and autotrophic dinoflagellate cysts (mainly Spiniferites ramosus, Lingulodinium machaerophorum and Operculodinium centrocarpum). These assemblage shifts are associated with stronger upwelling during stadials and stronger river influx during interstadials. Increases in productivity caused by enhanced upwelling are coupled to improved preservation and vice versa. More stratified water is indicated by higher abundances of L. machaerophorum and succeeds Heinrich events. The average process length of L. machaerophorum can be used to track changes in salinity, since this shows a similar pattern as the d18OSW (paired Mg/Ca -d18O) reconstruction. During the last glacial, conditions were more saline than during the current interglacial. On a millennial scale, changes in salinity are opposite to open ocean salinities and the hydrological proxies, which can be explained by a modulation of the signal by stratification, isolation of the Basin or advection of freshwater masses. These results highlight both generalities and particularities of the palaeoecological record of this tropical semi-enclosed basin.

A high-resolution palynological analysis and a detailed palaeomagnetic study of a marine sequence recovered during IODP Expedition Leg 307 in the Porcupine Basin southwest of Ireland provide new insights into the regional depositional history and palaeoenvironmental evolution during Early Neogene times. The Hole 1318B studied was drilled on the upper slope of the continental margin in a water depth of 409 m, upslope from a province of carbonate mounds (the Belgica mound province). The diverse and well-preserved dinoflagellate cyst associations consist typically of deep neritic and oceanic species, mixed with a neritic component transported from the shelf, reflecting the deep depositional setting at the continental margin. The palaeomagnetic record together with the ranges of key dinoflagellate cyst species constrain the age of the studied sequence between 16.7 Ma and 12.01 Ma, that is, between the late Burdigalian and late Serravallian. The distinct unconformity terminating the Miocene sequence correlates to the global sequence boundary Ser4/Tor1 dated at 10.5 Ma, and represents, according to previous extensive seismic studies, a basin-wide erosional event. The overlying sediments are of Middle Pleistocene or younger age. Downslope from IODP Site 1318, carbonate mounds root on the erosional surface. The dinoflagellate cyst associations from the Porcupine Basin distinctly mirror the global cooling phase following the Middle Miocene Climatic Optimum. Cooling phase Mi3, a short-lived glaciation, is particularly well expressed and here dated at 13.6 Ma. The palynomorph record furthermore indicates a reduction of the productivity and an increase of oceanic oligotrophic species after 14 Ma, suggesting a reduction or perhaps even a shutdown of the upwelling.