Dinocyst Species Research Articles

Late Pleistocene–Holocene palynology and paleoceanography of İmralı Basin, Marmara Sea: Pollen-spore, dinoflagellate cyst and other NPP zonations

The Marmara Sea is the oceanographic gateway that links the hypersaline Eastern Mediterranean Sea with the hyposaline Black Sea via Straits of Dardanelles and Bosphorus, respectively, and also occupies a transitional vegetation and climatic regime between the Mediterranean and Euxinic phytogeographic ecosystems. Recent palynological studies of long cores from the shallow İmralı Basin perched on the southeastern slope of the Marmara Sea reveal that major paleoecological changes accompanied the isolation and reconnection of the Marmara Sea during the last two Pleistocene glacial–interglacial cycles. Here, we use data primarily from two İmralı Basin cores to determine new quantitative zonations for the pollen-spore, dinoflagellate cyst (dinocyst) and other non-pollen palynomorph (NPP) assemblages in sediments representing the last ∼30 cal ka. A gap in one core is closed using data from three short gravity cores. Twenty-four radiocarbon ages and 20 correlation-ties for sapropel and ash layers from these five cores provide an essentially complete record for the past 30 cal ka. Use of a full suite of palynomorph data (pollen and spores, fungal spores, dinocysts and other phytoplankton, aquatic parasites/saprophytes, zoo-planktonic and zoo-benthic NPPs) shows for the first time that fluctuations in aquatic palynoassemblages during the glacial periods are more complex than simple salinity or temperature responses. The changes involve food web-level turnovers in trait- and size-based food web components in which potential parasites/saprophytes appear critically important.Pollen stratigraphies reveal a significant loss of Euxinic tree taxa by MIS 3c, with only partial recovery in MIS 1. The Euxinic forest elements are replaced by cultivars and other indicators of anthropogenic influences during the MIS 1 that is also marked by a dry interval and corresponding in part with the 4.2 cal ka Eastern Mediterranean Drought Event. The dinocyst zones for the last ∼30 cal ka show a clear turnover from assemblages dominated by brackish Ponto–Caspian taxa to marine assemblages at ∼11.7 cal ka. This phytoplanktonic signal lags slightly behind (a) the vegetation and shift in the other NPP commencing ∼13.2 cal ka, the initial marine incursion at 13.8 cal ka, and the arrival of coccolithophores at ∼13 cal ka. Comparison to published MIS 6 and 5 data reveals higher MIS 2–1 species diversity, reappearance of the MIS 6 marker Impagidinium inaequalis, and the emergences of Pyxidinopsis psilata f. quadrata, cf. Boreadinium breve, and Peridinium spp. of Brenner 2005. The pre-reconnection microbiota is dominated by the unicellular alga Sigmopollis, spikes of colonial Pediastrum algae, the saprophytes/parasites cf. Multiplicisphaeridium and Chytridiomycota. This microbiota is displaced by the post-reconnection assemblage dominated by Botryococcus coenobia, various Platyhelminth egg capsules and ostracod lining-types that persist through the Holocene. A distinctive transitional microbiota with Discocotyle-type turbellarian egg capsules marks the Bølling–Allerød warming, followed at ∼13.3 ka by a marine assemblage with micro-foraminiferal linings and crustacean eggs. The interglacial marine phase of MIS 1 in İmralı Basin has much higher dinocyst species diversity than MIS 5e, reflecting Neolithic impacts (agriculture, land clearance) and expanded shipping trade. Initial marine flooding in the Marmara Sea during both interglacials are marked by sapropel deposition but variations of ravinement surface erosion, and marine deep water residence time in a stratified Marmara Sea are potentially important factors underlying hypoxia/anoxia and sapropel deposition.

Palynostratigraphy and palaeoenvironment of the Danian sediments from the Nsukka Formation of the Anambra Basin at the vicinity of Ikpankwu, southeastern Nigeria.

Detailed investigations of biostratigraphy of the Nsukka Formation in the Ikpankwu area are paramount due to the fluctuation of depositional environment, fossil flora abundance and diversity interchange, and lateral facies change dynamics from the southern to northern sections of this formation. The microflora assemblages, chronostratigraphic age and depositional paleoenvironment of index palynomorphs suggest a Danian age for the studied strata, based on global dinoflagellate cyst events. The persistent abundance of monocolpate assemblages and the biostratigraphic resolution attained, suggest that the studied sediment interval is continuous. The marker species of terrestrial microflora (Constructipollenites ineffectus, Echitriporites trianguliformis, Foveotriletes margaritae, Longapertites group, Proxapertites group, Retidiporites magdalenensis, Scabratriporites annellus, Spinizonocolpites group, Syncolporites marginatus) and dinocysts (Carpatella cornuta, Danea californica, Fibrocysta licia, Kenleyia pachycerata, Spiniferella cornuta subsp. kasira), compared with several reference sections from the tropical to subtropical province, engineered a refined geologic age and paleoenvironment of deposition of the sediments. These dinocysts taxa are global Danian index fossils which are valuable in the identification of earliest Palaeocene sequence of the formation. The depositional paleoenvironment alternately fluctuates from continental setting to marginal marine inner neritic and outer neritic conditions, based on the presence of land-derived miospores and dinocysts. These macroenvironments are the products of effective vacillating depositional environment systems between the lower and upper deltaic plains, oscillating from tidal flat, lagoon, tidal bar to nearshore open marine influenced settings. Lycopodiaceae-Zlivisporis blanensis fern spore with fragile perispore substantiate the autochtonous vegetation and least transport and palynomorph hydrodynamic system inherent in the predominant pollen and spore species and close proximity to fresh water settings visible in medium to coarse grained sandstone units of the middle section. These microenvironment syntheses of the Nsukka Formation revealed a confirmed and pronounced transgressive systems tract (TST), articulated in the shale sample from the Ikpankwu section of the Nsukka Formation, Anambra Basin. The palaeoenvironments and palynostratigraphic dynamics illustrate that overall progradation was trailed by sporadic retrogradation of the delta with a high biostratigraphic resolution system indicative of continuous biological imprints of the terrestrial and marine microflora of the Nsukka Formation during the Danian.

Dinoflagellate cyst and pollen assemblages as tracers for marine productivity and river input in the northern Gulf of Mexico

Abstract. Both marine dinoflagellate cysts and terrestrially derived pollen and spores are abundant in coastal sediments close to river mouths, making sediment records from such settings ideal to simultaneously study land–ocean climate interactions, marine productivity patterns and freshwater input over time. However, few studies consider the combined calibration of these palynological proxies in modern coastal sediments offshore from rivers, which is needed to strengthen the interpretation of paleoreconstructions. Here, we analyze the palynological content of marine surface sediments along land–sea transects off the Mississippi and Atchafalaya river mouths in the northern Gulf of Mexico (GoM) and test three palynological indices which are often employed in the paleo-domain: (i) the abundance of cysts of heterotrophic and autotrophic dinoflagellates (dinocysts) as a tracer for primary productivity (H/A ratio) and (ii) the ratio between non-bisaccate pollen and bisaccates (P/B) as well as (iii) the ratio between pollen (excluding bisaccates) and dinocysts (P/D), which are both tracers for river input and distance to the coast. Our results show that dinoflagellate cysts are most abundant on the shelf, where heterotrophic dinocyst species dominate coastal assemblages in reach of the river plume, while autotrophic taxa are more present in the oligotrophic open ocean. This is clearly reflected in decreasing H/A values further offshore. Individual dinocyst taxa also seem to inhabit specific niches along an onshore–offshore transect, linked to nutrient availability and proximity to the turbid river plume. The highest pollen concentrations are found close to the Mississippi river mouth and mostly represent a mixture of local coastal and upstream vegetation, whereas bisaccate pollen was most abundant further offshore of the Mississippi river. Multivariate redundancy analysis (RDA) performed on both pollen and dinocyst assemblages, a set of environmental parameters, and the three palynological ratios showed that net primary productivity was the most important variable influencing the dinocyst assemblages, likely as the result of nutrient input. Additionally, the RDA confirmed that the H/A ratio indeed seems to track primary productivity, while the P/B ratio results in a robust indicator for distance to the coast, and the P/D ratio better reflects river input. Together, our data confirm and further specify the suitability of these three palynological ratios in river-dominated coastal margins as proxies for (past) marine productivity and distance to the coast and river.

Single-species dinoflagellate cyst carbon isotope fractionation in core-top sediments: environmental controls, CO2 dependency and proxy potential

Abstract. Sedimentary bulk organic matter and various molecular organic components exhibit strong CO2-dependent carbon isotope fractionation relative to dissolved inorganic carbon sources. This fractionation (εp) has been employed as a proxy for paleo-pCO2. Yet, culture experiments indicate that CO2-dependent εp is highly specific at genus and even species level, potentially hampering the use of bulk organic matter and non-species-specific organic compounds. In recent years, significant progress has been made towards a CO2 proxy using controlled growth experiments with dinoflagellate species, also showing highly species-specific εp values. These values were, however, based on motile specimens, and it remains unknown whether these relations also hold for the organic-walled resting cysts (dinocysts) produced by these dinoflagellate species in their natural environment. We here analyze dinocysts isolated from core tops from the Atlantic Ocean and Mediterranean Sea, representing several species (Spiniferites elongatus, S. (cf.) ramosus, S. mirabilis, Operculodinium centrocarpum sensu Wall and Dale (1966) (hereafter referred to as O. centrocarpum) and Impagidinium aculeatum) using laser ablation–nano-combustion–gas-chromatography–isotope ratio mass spectrometry (LA/nC/GC-IRMS). We find that the dinocysts produced in the natural environment are all appreciably more 13C-depleted compared to the cultured motile dinoflagellate cells, implying higher overall εp values, and, moreover, exhibit large isotope variability. Where several species could be analyzed from a single location, we often record significant differences in isotopic variance and offsets in mean δ13C values between species, highlighting the importance of single-species carbon isotope analyses. The most geographically expanded dataset, based on O. centrocarpum, shows that εp correlates significantly with various environmental parameters. Importantly, O. centrocarpum shows a CO2-dependent εp above ∼ 240 µatm pCO2. Similar to other marine autotrophs, relative insensitivity at low pCO2 is in line with active carbon-concentrating mechanisms at low pCO2, although we here cannot fully exclude that we partly underestimated εp sensitivity at low pCO2 values due to the relatively sparse sampling in that range. Finally, we use the relation between εp and pCO2 in O. centrocarpum to propose a first pCO2 proxy based on a single dinocyst species.

Dinoflagellate cysts and benthic foraminifera from surface sediments of Svalbard fjords and shelves as paleoenvironmental indicators

Due to the Arctic amplification effect, the Svalbard archipelago is an important area for studying ongoing environmental changes. However, its marine ecosystem is extremely complex. In this study, we analyze modern assemblages of dinoflagellate cysts (dinocysts) and benthic foraminifera from surface sediment samples around Svalbard. We use multivariate statistical analyses to examine relationships between environmental conditions (summer and winter sea surface temperature and salinity, sea-ice cover, etc.) and both microfossil groups to evaluate their use as proxies for reconstructions of the marine environment in the region. Our results show that the most important factor controlling the environment around Svalbard is the Atlantic Water which mostly impacts the western coast, but its influence reaches as far as the eastern coast of Nordaustlandet. However, on a local scale, such factors as the sea-ice cover, the presence of tidewater glaciers, or even the morphology and hydrology of fjords become increasingly important. We found that two dinocyst species, cysts of Polarella glacialis and Echinidinium karaense, can be considered regional winter drift ice indicators. The relationships between environmental parameters and benthic foraminiferal assemblages are much more difficult to interpret. Although statistical analysis shows a correlation of benthic foraminiferal species with various environmental parameters, this correlation might be somewhat coincidental and caused by other factors not analyzed in this study. Nevertheless, the use of two complementary microfossil groups as (paleo)environmental indicators can provide a more comprehensive picture of the environmental conditions.

An expanded database of Southern Hemisphere surface sediment dinoflagellate cyst assemblages and their oceanographic affinities

Abstract. Dinoflagellate cyst assemblages present a valuable proxy to infer paleoceanographic conditions, yet factors influencing geographic distributions of species remain largely unknown, especially in the Southern Ocean. Strong lateral transport, sea-ice dynamics, and a sparse and uneven geographic distribution of surface sediment samples have limited the use of dinocyst assemblages as a quantitative proxy for paleo-environmental conditions such as sea surface temperature (SST), nutrient concentrations, salinity, and sea ice (presence). In this study we present a new set of surface sediment samples (n=66) from around Antarctica, doubling the number of Antarctic-proximal samples to 100 (dataset wsi_100) and increasing the total number of Southern Hemisphere samples to 655 (dataset sh_655). Additionally, we use modelled ocean conditions and apply Lagrangian techniques to all Southern Hemisphere sample stations to quantify and evaluate the influence of lateral transport on the sinking trajectory of microplankton and, with that, to the inferred ocean conditions. k-means cluster analysis on the wsi_100 dataset demonstrates the strong affinity of Selenopemphix antarctica with sea-ice presence and of Islandinium spp. with low-salinity conditions. For the entire Southern Hemisphere, the k-means cluster analysis identifies nine clusters with a characteristic assemblage. In most clusters a single dinocyst species dominates the assemblage. These clusters correspond to well-defined oceanic conditions in specific Southern Ocean zones or along the ocean fronts. We find that, when lateral transport is predominantly zonal, the environmental parameters inferred from the sea floor assemblages mostly correspond to those of the overlying ocean surface. In this case, the transport factor can thus be neglected and will not represent a bias in the reconstructions. Yet, for some individual sites, e.g. deep-water sites or sites under strong-current regimes, lateral transport can play a large role. The results of our study further constrain environmental conditions represented by dinocyst assemblages and the location of Southern Ocean frontal systems.

Taxonomical Diversity of Cenomanian-Turonian Dinocyst in the Northern Hemisphere: Some Aspects of Paleobiogeography and Paleoclimatology

Taxonomic typification of dinocyst assemblages was carried out on the basis of a qualitative and quantitative assessment of their generic composition for the Northern Hemisphere in the Cenomanian-Turonian time in order to identify correlation taxa. Three types of dinocyst assemblages were identified in the Cenomanian. Cenomanian dinocyst assemblages are poorly differentiated; a large number of cosmopolitan genera was identified, which is in good agreement with paleobotanical data on a very warm, humid climate. Three types of dinocyst assemblages were established for the Turonian age. The number of cosmopolitan genera of dinocysts decreases, and species endemism sharply increases in individual Turonian basins. The differentiation of assemblages increases and their correlation potential decreases. However, the identified common genera and species of dinocysts in different types of assemblages provide the possibility of interregional correlation of Upper Cretaceous deposits at the stage, substage, and, at some sections, even more detailed levels in different climatic zones.

Cenomanian-Turonian dinocysts of North Hemisphere: some aspects of biogeography and paleogeography

Taxonomic typization of dinocyst assemblages was carried out on the basis of a qualitative and quantitative assessment of their generic composition for the Northern Hemisphere in the Cenomanian-Turonian time in order to identify correlation taxa. Typization was carried out by the method of cluster analysis according to the Jacquard calculation model (BioDiversity Professional program, 1997). To combine clusters into a hierarchical structure, the method of group average connection was used. Three types of dinocyst assemblages have been identified in the Cenomanian. Cenomanian dinocyst assemblages are poorly differentiated; a number of cosmopolitan genera have been identified, which is in good agreement with paleobotanical data on a very warm, humid climate. Three types of dinocyst assemblages have been established for the Turonian time. The number of cosmopolitan genera of dinocysts decreases, and species endemism sharply increases in some Turonian basins. The differentiation of assemblages increases and the correlation potential decreases. However, the identified common genera and species of dinocysts in various types of assemblages provide the possibility of interregional correlation of Upper Cretaceous deposits at the stage, substage, and, at some sections, at a more detailed level in different climatic zones.

Environmental inferences based on benthic foraminifera in surface sediments from the Beagle Channel (Argentina): A multiproxy approach

Micropaleontological analysis was performed on 22 samples of recent marine sediments from the Beagle Channel (Tierra del Fuego, Argentina) between 10 and 100 m of depth. The most important foraminifera found in the channel were Cibicidoides dispars (d'Orbigny), Buccella peruviana (d'Orbigny), Cibicides aknerianus (d'Orbigny), Cibicides fletcheri Galloway and Wissler, and Neoeponides bradyi (Le Calvez), which inhabit in clayey–sandy marine sediments. The observed variations in diversity and abundance revealed shallow marine conditions, low-energy, and low to moderate salinity, in the west sector and a deeper environment with normal marine conditions, high-energy, highly‑oxygenated and cold seawater in the east sector of the channel. The study area is a complex scenario with numerous landforms that lead to the formation of micro-environments in which patchy distribution of foraminifera was recognized. This study based on foraminifera and the sedimentological analysis supports the previous palynological information from the channel, which also shows an increase in number and diversity of dinocyst species reflecting the eastward changes of the marine environmental conditions due to the influence of the Atlantic.

Palaeoenvironmental changes based on foraminifera during the late Holocene at the Beagle Channel, Argentina.

A foraminiferal faunal study was carried out in a Holocene marine section from Arroyo Baliza, located on the northwest coast of the Beagle Channel, to contribute to the knowledge of the palaeoenvironmental conditions during the marine Holocene event. Foraminiferal assemblage was represented by 32 species distributed among 21 genera. The assemblage was dominated by Elphidium macellum (Fichtel & Moll) Elphidium alvarezianum (d'Orbigny), Cribroelphidium excavatum (Terquem) and Buccella peruviana (d'Orbigny), accompanied by Cibicides fletcheri Galloway & Wissler and Cibicidoides dispars (d'Orbigny) in low proportion. The predominance of Peridiniales dinocysts in the marine palynomorphs assemblage, suggested inner neritic conditions with cooler and more nutrient-rich waters. The distribution of the foraminiferal species was variable throughout the section indicating palaeoenvironmental changes in Arroyo Baliza between 3499-2595 cal yr BP. A gradual passage from high energy, cold and well-oxygenated marine waters towards a shallower environment with low energy and low to moderate salinity of the waters were linked to the regressive phase. This study complements and supports the previous palynological data from this section, which reflects an increase in number and diversity of dinocyst species indicating marine environmental conditions during the late Holocene as it exists today in the Beagle Channel.

Cadosinopsis rehakovii sp. nov., a new calcareous dinocyst from the Jurassic-Cretaceous transitional interval of the Western Tethys.

Variegated limestones, a transitional series between red, Upper Jurassic radiolarite and whitish, Lower Cretaceous Maiolica limestone in the Pieniny Klippen Belt deposits in the Polish part of the Western Carpathians, have yielded rich microfossil assemblages with common calcareous dinoflagellate resting stages, hereafter, dinocysts. We found an undescribed dinocyst species in red-greenish limestone of a deep water, pelagic habitat in the Branisko succession of the Pieniny Klippen Basin and named it Cadosinopsis rehakovii sp. nov. The new species has a spherical to oval calcareous test ranging from 34 to 59 μm in length and 30 to 50 μm in width, with two layered wall. The inner layer is built of coarse–thick, plate-shaped calcite crystals and is white in transmitted light. The outer layer is built of fibrous crystals, vitreous (transparent) in transmitted light. The aperture is wide and seen only in the inner layer. We compared the detailed morphological characteristics of the new species with another species from the same genus in the Jurassic and Cretaceous Tethyan deposits. Specimens have been measured, grouped and interpreted using cluster analysis, principal component analysis (PCA) and canonical variate analysis (CVA). Among them, the new species shows a resemblance in cross-section to other species of Cadosinopsis, C. nowaki Borza, 1984, and C. andrusovi Scheibner 1967, previously described in literature. However, the two species are easily distinguishable by some features. Cadosinopsis. nowaki is bigger in size (length from 50 to76 μm and width from 43 to 67 μm), its inner layer is thicker and consists of vitreous-sparite calcite, and it has less centrically located chamber. C. andrusovi is much bigger in size as its length ranges from 68 to 108 μm and width ranges from 60 to 80 μm, and the cyst is more oval and its chamber less spherical. The new species is the third Cadosinopsis species described in the Tethyan realm and about two hundred and sixty-first fossil species (morphotype) described in the world so far.

Dinoflagellate cyst distribution in surface sediments of Ambon Bay (eastern Indonesia): Environmental conditions and harmful blooms

The present study aimed to document dinocyst ecological preferences in Ambon Bay, Eastern Indonesia, and to investigate if the bay sediments serve as a seedbank for toxic bloom events. To this end, dinocyst and geochemical analyses of surface sediment samples were performed, along with physicochemical water column parameters. Twentythree dinocyst species were identified, and high dinocyst concentrations (up to ~12,000 cysts g−1 dry sediment) were found in the inner bay. Environmental factors such as surface water temperature and salinity generally played an important role in dinocyst distribution. The concentration of Polysphaeridium zoharyi cysts showed a strong positive correlation with phosphorus. A statistically significant correlation was also found with the concentration of other autotrophic dinocysts in the sediments, and an inverse correlation was observed with the sediment C/N ratio. Cysts may serve as seedbanks for Pyrodinium bahamense blooms in the area.

A multi‐proxy based high‐resolution stratigraphical analysis of the possible Palaeocene–Eocene boundary interval, Salt Range, Pakistan

The Palaeocene–Eocene boundary (PEB) interval has been recognized in the Patala Formation, Nammal Gorge Section, Pakistan using lithostratigraphy, stable carbon isotope stratigraphy and integrated biostratigraphy. Four distinct lithological units are identified in the Patala Formation in stratigraphic order from bottom to top; unit‐1 to unit‐4. The top of unit‐3 hosts three closely spaced unconformities demarcated as U1, U2, and U3. The PEB interval is represented by a negative shift of 1.61‰ in δ13CV‐PDB representing the carbon isotopic excursion (CIE) in unit‐2. Based on the encountered foraminifera, dinocysts, and calcareous nannoplanktons, the following biozones are identified across the PEB; two larger benthonic foraminiferal (LBF) zones (SBZ4 and SBZ5), the zonal boundary slightly postdating the CIE, two smaller benthonic foraminiferal zones (BB1 and BB2), three planktonic foraminiferal zones (P4/P5 + E1), four dinocysts zones (Pak‐DV, Pak‐DVI, Pak‐DVII, Pak‐DVIII), and two calcareous nannoplankton subzones (NP9a and NP9b). The following protist, dinocyst and calcareous nannoplanktons encountered in the studied interval advocates for the placement of the possible PEB at the level between samples NAM‐SH‐07 and 08; 1—presence of the smaller benthonic foraminifera Angulogavelinella avnimelechi diagnostic of the base of the CIE and the benthonic extinction event (BEE) Tethys wide, 2—presence of the cosmopolitan and the PEB diagnostic dinocyst species ‘Axiodinium augustum’, 3—presence of the PEB diagnostic Rhomboaster‐Discoaster calcareous nannoplankton assemblage. The LBF species Alveolina vredenburgi [a marker species of the Larger Foraminiferal Turnover (LFT) associated with the Palaeocene Eocene Thermal Maximum (PETM) and PEB in western Tethys] occurs slightly above the onset of the CIE. Previously reported compositional change in the LBF assemblage from the Salt Range [i.e. the replacement of the Palaeocene orbitoidiform LBF (Setia and Orbitosiphon) by the Eocene LBF (Orthophragminids, Alveolina, Nummulites and Assilina)] also postdates the PEB interval here. The late appearance of Alveolina vredenburgi and the compositional change in the LBF after the onset of the CIE are attributed to the facies variation across the PEB reflecting fluctuating depositional environment due to the active tectonic across this interval. This fluctuation is evident from the smaller benthonic and planktonic foraminiferal dominating assemblages in unit‐2 representing bathyal settings followed by LBF dominating unit‐3 representing shallow carbonate platform settings. The change in LBF assemblage closely resembles the well‐known LFT associated with the PETM as well as the carbonate platform stage III and the PEB reported from other Tethyan sections.

Distribution of dinoflagellate cyst assemblages in recent sediments from a southern Mediterranean lagoon (Mellah, Algeria) with emphasis on toxic species.

This is the first study on the dinoflagellate cysts in Algerian waters and in Mellah Lagoon (South Western Mediterranean), located within a protected reserve. In total, 42 species of dinocysts belonging to 7 orders, 12 families and 23 genera, were identified in the 26 superficial sediment samples from Mellah Lagoon. The distribution of dinocysts in the sediment of this lagoon is heterogeneous. Indeed, their abundance oscillates between 1 and 315cystsg-1 dry sediment (DS). Cyst morphotype assemblages were dominated by a few numbers of species: Alexandrium minutum (15.87%), Gonyaulax verior (9.81%), Protoperidinium spp. (7.74%), Alexandrium affine (7.05%), Scrippsiella trochoidea (6.67%), and Alexandrium pseudogonyaulax (6.19%). There is a positive correlation between the density of cysts and the depth (r = 0.61; p < 0.05), organic matter (r = 0.70; p < 0.05), water content (r = 0.71; p < 0.05), and the fine fraction of sediment (r = 0.74; p < 0.05). Surprisingly, although the Mellah Lagoon is almost semi-closed, it holds an important specific richness in dinocysts (42 species) higher than others observed in Mediterranean lagoons. However, cyst abundances are low compared to other lagoons in the Mediterranean Sea. Finally, the presence of dinocysts of Alexandrium catenella/tamarense, A. minutum, and Gymnodinium catenatum associated to paralytic shellfish toxins, A. pseudogonyaulax which produces goniodomin A, also Protoceratium reticulatum and Gonyaulax spinifera complex which produce yessotoxins, needs to implement a monitoring program to prevent a potential human intoxication due to the consumption of contaminated sea products by these potent neurotoxins.

Campanian to Danian dinoflagellate cyst assemblages from the southwestern Tethyan margin (Tattofte section, western External Rif, Morocco): Biostratigraphic and paleobiogeographic interpretations

A palynological study of the Upper Cretaceous–lower Paleocene outcrop section at Tattofte (westernmost External Rif, Morocco) has produced a well-preserved palynological assemblage rich in dinoflagellate cysts (dinocysts). For the first time, these assemblages have allowed us to determine the age of this section, to highlight a new Cretaceous–Paleogene boundary, and to make paleobiogeographic interpretations. The samples analyzed yielded 209 dinocyst species and 81 biostratigraphic dinocyst events (first occurrence, or FO, and last occurrence, or LO). We recognized the upper Campanian based on the FOs of Areoligera coronata, Areoligera reticulata, Areoligera senonensis, Cerodinium diebelii and Andalusiella mauthei subsp. mauthei, as well as the LOs of Batiacasphaera solida, Odontochitina costata, Odontochitina porifera, Sentusidinium eisenackii, Trichodinium castanea and Trithyrodinium suspectum. The lower Maastrichtian is determined by the FOs of Isabelidinium bujackii, Palaeocystodinium golzowense and Trithyrodinium evittii, and by the LO of Alterbidinium acutulum, and the upper Maastrichtian is recognized primarily by the FOs of Cerodinium speciosum subsp. speciosum, Disphaerogena carposphaeropsis, Glaphyrocysta perforata and Manumiella seelandica and the LO of Dinogymnium spp. The Danian is recognized on the basis of the FOs of several taxa, including Carpatella cornuta, Danea californica and Senoniasphaera inornata. The succession of the dinocyst events in the uppermost part of the studied interval has revealed a relatively complete record across the Cretaceous–Paleogene boundary. The Cretaceous–Paleogene boundary coincides with a 8 cm-thick reddish layer, which resembles those observed previously in the Cretaceous–Paleogene boundary reference sections in the Mediterranean area, most notably the Cretaceous–Paleogene boundary stratotype at El Kef (Tunisia). The upper Campanian dinocyst assemblage indicates a subtropical to temperate setting, while the Maastrichtian and lower Danian assemblages indicate a subtropical to warm temperate setting. The incursion of a few higher latitude cold-water dinocyst species in these provinces during the latest Campanian and around the Cretaceous–Paleogene boundary may suggest a southward migration of these species, probably related to short cooling episodes and ocean circulation.

Middle Eocene dinoflagellate cysts from the Tsoul section, eastern External Rif, Morocco: biostratigraphy and paleoenvironmental interpretations

This palynological study is carried out on the Eocene sediments in the Tsoul section, which is located in the Taza region, Tsoul Unit, eastern External Rif, northeastern Morocco. It aims their age assessment and paleoenvironmental interpretations, based on palynological data. The palynological content is diverse, well preserved and rich in organic-walled dinoflagellate cysts (dinocysts), whereas spores and pollen are rare. We identified 117 dinocyst species (including some reworked species), among which 6 species and 1 acme are good biostratigraphic markers of the Bartonian (middle Eocene). The main dinocyst bioevents that are used to assign the studied section to the Bartonian include the lowest occurrences (LOs) of Glaphyrocysta semitecta, Impagidinium dispertitum, Lentinia serrata, Membranophoridium aspinatum and Operculodinium divergens, the highest occurrence (HO) of Castellodinium compactum and the acme of Polysphaeridium spp. Regarding the paleoenvironment, the studied section was deposited in a marine environment, characterized by a low productivity inferred from the scarcity of peridinioid cysts and by important environmental changes ranging from restricted inner neritic (lagoon) conditions, inferred from the dominance of Homotryblium and Polysphaeridium, to outer neritic open marine conditions inferred from the dominance of Spiniferites.

Dinocysts from the mid–upper Eocene Aktumsuk section, Ustyurt, Uzbekistan: biostratigraphy and paleoenvironments

The results of palynological study of the MidUpper Eocene sediments of the Aktumsuk key-section (Ustyurt Plateau, Uzbekistan), calibrated with nannoplankton data, are presented. Dinocyst stratigraphical distribution permitted to recognize five Dinoflagellate zones from the Peri-Tethys zonation: Costacysta bucina, Enneadocysta pectiniformis, Rhombodinium draco, Rhombodinium perforatum, and Thalassiphora reticulata. First-order calibration with nannoplankton zones permitted to precise the lowermost occurrences of the key dinocyst species. Based on the dinocyst and nannoplankton data, two important stratigraphic hiatuses, corresponding to the upper Ypresian and Upper BartonianLower Priabonian, were recognized. Analysis of quantitative fluctuations of different groups of palynomorphs through the section indicates paleoenvironmental changes in this part of the Peri-Tethys during the LutetianPriabonian: the beginning of significant transgressive phase in the early Lutetian, relative deepening in the Mid Lutetian; after the break in sedimentation at the end of Bartianianbeginning of Priabonian the sedimentation continues again in the open-marine (outer neritic) environment.

Middle–Late Eocene Dinocysts from the Aktumsuk Section (Ustyurt Plateau, Uzbekistan): Biostratigraphy and Paleoenvironments

The results of palynological study of the mid-upper Eocene sediments of the Aktumsuk key-section (Ustyurt Plateau, Uzbekistan), calibrated with nannoplankton data, are presented. Dinocyst stratigraphical distribution permitted to recognize five Dinoflagellate Zones from the Peri-Tethys zonation: Costacysta bucina, Enneadocysta pectiniformis, Rhombodinium draco, Rhombodinium perforatum, and Thalassiphora reticulata. First-order calibration with nannoplankton zones permitted to precise the lowermost occurrences of the key dinocyst species. Based on the dinocyst and nannoplankton data, two important stratigraphic hiatuses, corresponding to the upper Ypresian and upper Bartonian-lower Priabonian, were recognized. Analysis of quantitative fluctuations of different groups of palynomorphs through the section indicates paleoenvironmental changes in this part of the Peri-Tethys during the Lutetian-Priabonian: the beginning of significant transgressive phase in the early Lutetian, relative deepening in the mid Lutetian; after the break in sedimentation at the end of Bartianian-beginning of Priabonian the sedimentation continues again in the open-marine (outer neritic) environement.

Transport Bias by Ocean Currents in Sedimentary Microplankton Assemblages: Implications for Paleoceanographic Reconstructions

AbstractMicrofossils from plankton are used for paleoceanographic reconstructions. An often‐made assumption in quantitative microplankton‐based paleoceanographic reconstructions is that sedimentary assemblages represent conditions of the directly overlying surface water. However, any immobile particle sinking down the water column is subjected to transport by three‐dimensional currents, which results in a lateral relocation along transport. We model dinoflagellate cyst (dinocyst) transport in a high‐resolution (0.1° horizontally) global model of the present‐day ocean and compare ocean conditions in the simulated origin of sedimentary particles to that in the directly overlying water. We find that the assumption that sedimentary particles represent the overlying surface waters is in most regions not valid. The bias induced by dinocyst transport depends on ocean current strength and direction, aggregation of particles which could increase the sinking speed, and the sediment sample depth. By using realistic sinking speeds of dinocysts and aggregates, extreme biases up to approximately ±16 °C warmer or ±4 PSU saltier are found, while other regions show lower bias from particle transport. Our model results provide a way to mechanistically and statistically explain the unexpected occurrences of some dinocyst species outside of their “normal” occurrence region, such as the northerly occurrence of the allegedly sea‐ice‐affiliated dinocyst Selenopemphix antarctica. Exclusion of such outlier occurrences will yield better constrained ecological affinites for dinocyst species, which has implications for microfossil‐based quantitative and qualitative proxies for paleoceanographic conditions. We recommend paleoceanographers to a priori evaluate the (paleo)water depth, oceanographic setting, current strength, and particle aggregation probability for their sedimentary microplankton assemblages.

Lutetian to Priabonian dinocyst assemblages from the northwestern Tethyan margin (Adelholzen section, Eastern Alps, Germany)

This paper is the first systematic documentation of Eocene organic-walled dinoflagellate cyst (dinocyst) assemblages from the northwestern shelf of the Tethys. A taxonomic list of the rich and well-preserved dinocyst assemblages found within 28 samples collected at the Adelholzen section in southeastern Germany is presented herein. The sediments accumulated during the Lutetian and Priabonian stages at a ramp-type margin. One-hundred organic-walled dinocyst species were found at the section, which is riddled with unconformities. The composition of the successive dinocyst assemblages reflects the sea-level changes within the depositional area. Areoligera coronata and Cordosphaeridium gracile dominate the assemblage within the glauconite-rich siliciclastic sandstone (lower Adelholzen beds) of the lower transgressive system tract, which spans the calcareous nannoplankton sub-Zones NP15a and NP15b. A hiatus comprising at least c. 1.5 million years truncates these deposits and indicates an erosional episode. Rapid sea-level rise in Biochron NP16 caused a new transgression and the deposition of calcareous marlstone and limestone rich in nummulitids. In this part of the section (middle Adelholzen beds), Homotryblium tenuispinosum is by far the most abundant dinocyst species, and rare but consistent occurrences of Impagidinium spp. suggest a more open marine environment. A lithological shift from limestone to marlstone, a change from nummulitids to discocyclinids, and finally the complete disappearance of larger benthic foraminifera indicates continuous deepening of the environment. An increase in Spiniferites spp., Operculodinium spp., and occurrences of Homotryblium floripes characterize the dinocyst assemblage of this marlstone (upper Adelholzen beds). A 0.5 m thick condensed layer, which consists essentially of iron-coated glauconite grains and scattered phosphate nodules, represents the maximum flooding surface of the transgressive sequence, for which a paleodepth of c. 300 m is assumed. The overlying marlstone (“Stockletten”) of the highstand system tract is punctuated by a hiatus of c. 4.3 million years spanning the entire Bartonian. The stratigraphically important species Rhombodinium longimanum, Rhombodinium perforatum, Distatodinium ellipticum, Nematosphaeropsis labyrinthus and Selenopemphix nephroides have their lowest occurrences in the Priabonian (Zones NP18 and NP19) of the section.