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Schematic dinoflagellate life cycle. A) Haploid phase where individual cells divide through mitosis. This is the vegetative stage, which leads to the formation of bloom events. Following an internal and/or external stimuli, gametes are formed and will fuse to produce a diploid zygote; B) Planozygote (pelagic motile zygote) phase where the diploid cell will eventually lose motility, and in which the cyst will develop and eventually be released in the water column, then down to the sediment surface; C) Hypnozygote stage where the cyst will undergo a mandatory dormancy period controlled by an endogenous clock before excystment takes place. Modified after Evitt [31]. 

Schematic dinoflagellate life cycle. A) Haploid phase where individual cells divide through mitosis. This is the vegetative stage, which leads to the formation of bloom events. Following an internal and/or external stimuli, gametes are formed and will fuse to produce a diploid zygote; B) Planozygote (pelagic motile zygote) phase where the diploid cell will eventually lose motility, and in which the cyst will develop and eventually be released in the water column, then down to the sediment surface; C) Hypnozygote stage where the cyst will undergo a mandatory dormancy period controlled by an endogenous clock before excystment takes place. Modified after Evitt [31]. 

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Dinoflagellates are eukaryotic organisms and constitute an important group of marine primary producers. Approximately 10-15% of living dinoflagellates produce a highly resistant dormant cyst that is fossilisable, and which constitute an excellent proxy indicator of the upper water column conditions and productivity. Relatively little is known on th...

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... are single-celled eukaryotic organisms that inhabit both freshwater and marine environments from equatorial to polar latitudes. In marine environments, they constitute together with diatoms and coccolithophores an important group of primary producers. About half of the extant dinoflagellates are autotrophic, or mixotrophic (combination of phototrophy and heterotrophy), while the other half are heterotrophic, and some are parasitic. Approximately 10-15% of dinoflagellates form a cyst composed of a highly resistant polymer, as part of their life cycle [1]. The cyst stage, or hypnozygote, is believed to be a benthic phase in the life cycle of the dinoflagellates [2] and can remain dormant in the sediments from several days to years. Quaternary dinoflagellate cysts, known as dinocysts, assemblages are widely used by marine palynologists as a proxy indicator of conditions in the upper part of the water column [3, 4, 5], eutrophication [6, 7, 8] and productivity [9, 10]. Over the last few decades, the distribution of dinocyst assemblages in the surface sediments of the ocean basins, especially the Arctic basins, have been the subject of numerous studies in an effort to develop their potential for paleoceanographical reconstructions [11, 12, 13, 14, 15, 16, 17]. So far, approximately 31 dinocyst taxa have been observed in surface sediments of the Canadian Arctic [3, 14, 41, 45]. Several studies reported the distribution of the vegetative motile forms of dinoflagellates in the Arctic [18, 19, 20, 21, 22, 23, 24, 25, 25, 26, 27] and ~192 taxa belonging to 42 genera have been recorded from the water column of the Canadian High Arctic [18, 23, 24, 24, 27, 27, 28, 29]. However, our knowledge of the relationships between the plankton motile forms and their cyst counterpart in the sediments is rather limited. For example, the cyst stage of several motile forms collected from the phytoplankton have not been recorded in surface sediments yet, while several motile forms related to the known Arctic cyst taxa have not been observed in Arctic phytoplankton samples or described yet. The knowledge of the biological affinity or cyst-theca relationships for Arctic dinoflagellates is important because it provides useful information on dinoflagellate life cycle. In addition, the increasing understanding of the motile form will prove extremely useful for paleoceanographical reconstructions and interpretation. A better knowledge of the distribution of the dinoflagellate motile forms will also provide hints on the transport mechanisms affecting the dinocysts after their formation and subsequent deposition in the surface sediments. Dinoflagellates have a relatively complex life cycle that includes a motile plankton or vegetative stage, and some include a resting or cyst stage (Figure 1). The motile forms are characterized by the presence of two flagella, one longitudinal and the other located in a depression or cingulum, around the cell. The cell is comprised within a membrane, the amphiesma, which can include a series of cellulosic plates (referred to as “armored” or thecate), while others lack those cellulosic plates and are referred to as “naked” or “athecate”. Dinocysts are either composed of dinosporin, a complex biomacromolecular substance composed of phenolic, alcoholic and/or carboxylic hydroxides, and fatty acids ranging from C14 to C16, accompanied by tocopherols and sterols (mostly cholesterol and dinosterol) [30], or calcium carbonate. The morphology of most motile and cyst species are often very different, which makes it difficult to understand the complete life cycle. Also, the motile and cyst forms were studied separately by biologists and marine palynologists respectively. This gave rise to two distinct nomenclature systems, which further complicates the understanding of their life cycle. The distribution of dinoflagellates is not only dependent on the physico-chemical variables (currents, temperature, salinity, irradiance, nutrients) of the water column, but also on their feeding strategies, which may include mixotrophy and phagotrophy (ingestion of large food particles or preys) as well as the distribution of their prey. Over the last decade, several research programs aimed at studying the changing Arctic environment were conducted in the northern Baffin Bay (International North Water Polynya Study), the Beaufort Sea (Canadian Arctic Shelf Exchange Study) and the Canadian Arctic Archipelago (ArcticNet). The ecology of plankton and ice diatoms and other protists have been extensively studied over long-term periods as part of these Arctic research programs, but very little has been done on the ecology of dinoflagellate bloom events. The main reasons are that dinoflagellates are usually less abundant in the water column and they are adapted to growth under low turbulence and low nutrients concentrations [56]. Dinoflagellate blooms often occur after the one of diatoms and usually are restricted in time and space, therefore easily missed. For example, in highly productive Arctic basins, such as the North Water Polynya in northern Baffin Bay, diatoms usually dominate the phytoplankton and may account up to 70% of the total phytoplankton carbon biomass, while dinoflagellates and ciliates comprise approximately 24% of the total phytoplankton carbon biomass [24, 24, 32]. However, dinoflagellates may account for the majority of the phytoplankton carbon biomass during certain periods in relation to the depth of the surface mixed layer. 2.1. Cyst-theca relationships of Arctic dinoflagellates Dinoflagellates have been studied by biologists since the mid-late eighteenth century, while dinoflagellate cysts were studied by marine palynologists since the early nineteenth century [31], which lead to a double nomenclature system for the motile and cyst stages. The first cyst incubation experiments in the 1960s [33] paved the way to a number of studies aiming at establishing the cyst- theca relationship of the cyst stage [34, 35, 36, 37, 38, 39]. Several dinoflagellate species closely resemble each other and, in several cases, only detailed examination in scanning electron microscopy resulted in accurate identification. The best example is the Gonyaulax spinifera (Claparède et Lachmann 1859) Diesing 1866 complex, in which ~13 cyst species were supposedly derived from one single motile species [39]. Therefore, species identification can sometimes be erroneous when observing plankton samples because the minute details may have been missed. Six studies provide checklists of motile dinoflagellate species present in the Canadian High Arctic [18, 23, 24, 27, 28, 29] and they accounted for most of the phytoplankton surveys in the area since the 1970s. A total of ~192 taxa belonging to 42 genera have been identified, among which 10 species are known to produce resistant resting cysts as part of their life cycle ( Brigantedinium cariacoense = Protoperidinium avellana (Meunier 1919) Balech 1974; B. simplex = Protoperidinium conicoides (Paulsen 1905) Balech 1973; Pentapharsodinium dalei Indelicato et Loeblich 1986 = theca-based name; Polykrikos kofoidii Chatton 1914 = theca-based name; Polykrikos schwartzii Bütschli 18, 1873 = theca-based name; Protoperidinium americanum (Gran et Braarud 1935) Balech 1974 = theca-based name; Protoperidinium nudum = theca-based name; Spiniferites elongatus = Gonyaulax elongata (Reid 1974, 74) Ellegaard, Daugbjerg, Rochon, J. Lewis et Harding 2003; Trinovantedinium applanatum = Protoperidinium pentagonum (Gran 1902) Balech 1974). In some cases, the cyst has been observed while forming inside the theca within plankton samples, while in other cases, the relationship has been confirmed through laboratory cyst incubation experiments. Several studies have ...

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... The complete phytoplankton community analysis revealed a high abundance of diatoms at the time this port was visited (Howland and Simard, unpublished data). Thus, it appears that dinoflagellate sampling in this port was done during a diatom bloom shortly after ice break-up, which usually occurs before the dinoflagellate bloom (Rochon, 2009;Simo-Matchim et al., 2017). Diatoms are the first to bloom after the melting of the sea ice due to their capacity to thrive in areas characterized by low light availability and high nutrient concentrations (Sarthou et al., 2005). ...
... Diatoms are the first to bloom after the melting of the sea ice due to their capacity to thrive in areas characterized by low light availability and high nutrient concentrations (Sarthou et al., 2005). The diatom bloom eventually ends when the upper mixed layer is depleted of nitrate and silicate (Tremblay et al., 2002) and is followed by the dinoflagellate bloom, which do not require silicate or high nutrient concentrations to continue developing (Margalef, 1978;Tremblay et al., 2002;Rochon, 2009). ...
... The low richness in Milne Inlet and Iqaluit in 2015 could be result from early sampling relative to the sea ice melt (Fig. 7). The distribution of dinoflagellate communities depends on currents, temperature, salinity, nutrients, feeding strategies and the distribution of their prey (Rochon, 2009). In fact, species of this group have different trophic modes and their biomass and distribution depend on autotrophic and heterotrophic nutrition (Golubkov et al., 2019). ...
Article
The expected increase of shipping activities in the Canadian Arctic is predicted to enhance potential introductions of non-indigenous species (NIS), including dinoflagellate taxa, which may have important ecological and economic impacts once released in a new environment. The lack of information about native species represents an obstacle in detecting the arrival of NIS. In this context, the present study characterizes dinoflagellate communities in high-risk Canadian Arctic ports to provide baseline data and to verify the presence of potential NIS and harmful taxa. In total, we identified 49 dinoflagellate taxa from 9 families in the ports of Churchill, Deception Bay, Iqaluit and Milne Inlet, including 7 taxa known to be potential toxin producers. Dinoflagellate communities differed significantly between ports and among time periods in the heavily used ports, those of Churchill (between 2007 and 2015) and Iqaluit (between 2015 and 2019). Comparisons between dinoflagellate communities in the ports and those in ballast water showed that 12 taxa found in ballast water of vessels discharging in Churchill and Deception Bay are potential NIS, confirming the introduction of new species by shipping activities. This may be exacerbated in the near future as a result of extended/prolonged ice-free conditions due to global warming.
... One of the key challenges in the application of sediment dinoflagellate cysts as tracers of past environments, particularly in polar and subpolar seas, is the limited knowledge of the life-cycle transitions of individual dinoflagellate species (Rochon, 2009;Kremp, 2013): i.e., the seasonal patterns and environmental cues of cyst production and the relationships of sedimentary cysts to their planktonic counterparts are not known. This challenge is not always faced outright since spatial distribution of dinoflagellate cysts in recently deposited surface sediments can be directly linked to modern average surface water conditions via statistical modeling, i.e., "transfer functions", without the need to consider planktonic motile stages or the transitions in the life cycle. ...
Article
We present continuous bi-weekly to bi-monthly dinoflagellate cyst, tintinnid loricae and tintinnid cyst fluxes at two mooring sites in Hudson Bay (subarctic Canada) from October 2005 to September 2006. The total dinoflagellate cyst fluxes at the site on the western side of the bay ranged from 4600 to 53,600 cysts m− 2 day− 1 (average 20,000 cysts m− 2 day− 1), while on average three times higher fluxes (average 62,300 cysts m− 2 day− 1) were recorded at the site on the eastern side of the bay with a range from 2700 to 394,800 cysts m− 2 day− 1. These values are equivalent to the average fluxes calculated from the top 1-cm sediment layer of 210Pb-dated box cores at corresponding locations, and hence lend support to the use of sediment dinoflagellate cysts in palaeoceanography. Tintinnid fluxes ranged from 1200 to 80,000 specimens m− 2 day− 1 (average 32,100 tintinnids m− 2 day− 1) in the west, and 1600 to 1,240,800 specimens m− 2 day− 1 (average 106,800 tintinnids m− 2 day− 1) in the east, with the highest Salpingella sp. fluxes recorded during the sea-ice cover season.
... Some of the dinoflagellate cyst taxa in our sediment samples have known motile counterparts, e.g., motile stages of O. centrocarpum, S. elongatus and Selenopemphix quanta and are mentioned in the published species lists (Bursa, 1961;Harvey et al., 1997). However, most of the theca equivalents of arctic and sub-arctic dinoflagellate cysts are not yet known (Rochon, 2009). Dinoflagellate cyst assemblages show distinct spatial patterns revealing three compositional domains in the HBS: eastern, western-central and Hudson Strait (Figs. 3,4,5). ...
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Surface sediment samples from the Hudson Bay system were analysed in order to examine the role of key regulators of arctic marine productivity — light and nutrients as affected by freshwater stratification and sea-ice cover — on the spatial distribution and production of dinoflagellate cysts. Total cyst fluxes vary from 0.2 × 106 to 30.6 × 106 cysts m− 2 a− 1, with the highest values observed in eastern Hudson Bay. A total of 24 cyst taxa, representing 11 genera of five orders, were identified and distribution maps of the most common taxa have been produced. This is the first record of Echinidinium aculeatum, Echinidinium karaense, cf. Echinidinium delicatum, Islandinium brevispinosum, Selenopemphix quanta, cysts of Protoperidinium americanum, cysts of cf. Biecheleria sp. and Polarella glacialis in the Hudson Bay system. Dinoflagellate cyst assemblages show distinct spatial patterns revealing three compositional domains: eastern Hudson Bay, western-central Hudson Bay and Hudson Strait. The eastern domain is characterised by a dominance of autotrophic cysts of Pentapharsodinium dalei whereas the western-central domain is characterised by autotrophic Operculodinium centrocarpum with some contribution by heterotrophic Polykrikos sp. var. arctic morphotype and Polykrikos spp. Sites from Hudson Strait are distinguished by an overwhelming prevalence of heterotrophic Protoperidiniaceae cysts, mainly Islandinium minutum, and have the highest values of sedimentary biogenic silica, used as a proxy for diatom productivity.
... Several studies have recorded dinocyst taxa throughout the Arctic and subarctic regions (Harland et al. 1980, Rochon et al. 1999, Solomon et al. 2000, Hamel et al. 2002, Richerol et al. 2008. The relationships between the cyst and its motile counterpart in these regions have been confirmed for few species (Head 1996), representing less than one third of the recorded cyst species (Rochon 2009). The biological affinities of round brown spiny cysts occurring in these regions are currently unknown. ...
Article
Round brown spiny cysts constitute a morphological group common in high latitude dinoflagellate cyst assemblages. The dinoflagellate cyst Islandinium minutum (Harland et Reid) Head, Harland et Matthiessen is the main paleoecological indicator of seasonal sea‐ice cover in the Arctic. Despite the importance of this cyst in paleoceanographical studies, its biological affinity has so far been unknown. The biological affinity of the species I. minutum and its phylogenetic position based on the small subunit ribosomal RNA gene (SSU rDNA) and the large subunit ribosomal RNA gene (LSU rDNA) were established from cyst incubation experiments in controlled conditions, optical and scanning electron microscopy, and single‐cell PCR. The thecal motile cell obtained was undescribed. Although the motile cell was similar to Archaeperidinium minutum (Kofoid) Jörgensen, the motile cell of I. minutum lacked a transitional plate in the cingular series, which is present in Archaeperidinium spp. Islandinium minutum and Archaeperidinium spp. were paraphyletic in all phylogenetic analyses. Furthermore, Protoperidinium tricingulatum, which also lacks a transitional plate, was closely related to I. minutum and transfered to the genus Islandinium. Based on available data, it is clear that Islandinium is distinct from Archaeperidinium. Therefore, we considered Islandinium Head, Harland et Matthiessen as a non‐fossil genus and emend its description, as well as the species I. minutum. This is the first description of a cyst–theca relationship and the first study that reports molecular data based on SSU rDNA and LSU rDNA on a species assigned to the genus Islandinium.
... Many Arctic dinoflagellate taxa belong to Peridiniales, Gymnodiniales, Gonyaulacales and Dinophysiales (Okolodkov, 1998). However, only a few produce cysts that preserve in sediments (e.g., de Vernal et al., 2001;Matthiessen et al., 2005;Rochon, 2009). The cyst taxa recovered in sediment of sea ice environments are mostly Brigantedinium spp. ...
... The knowledge about the biologyeecology of dinoflagellates and their related cysts is incomplete, notably in the Arctic and subarctic seas (Rochon, 2009). To date, no dinoflagellate yielding fossilisable cyst has been demonstrated to live in sea ice, except I. minutum for which !99% similar SSU rRNA sequences have been retrieved (Potvin et al., 2013) during a study using high throughput amplicon sequencing on sea ice samples collected in the Beaufort Sea (Comeau et al., 2013). ...
... Dinocysts are thus excellent paleoenvironmental proxies especially for high latitude environments (e.g., Rochon, 2009), with respect to calcareous (e.g., foraminifers, coccolithophores) and siliceous (e.g., diatoms) microfossils which undergo severe dissolution in cold environments de Vernal et al., 2005b). The high sedimentation rates and excellent preservation of the oxidation-sensitive taxa, such as Brigantedinium cysts (produced by the Protoperidinium species) allow us to assume that little or no selective degradation occurred at station 803. ...
Article
Late Holocene paleoceanography and climate variability of the Southeastern Beaufort Sea (Canadian Arctic) have been investigated on the basis of sedimentary cores collected over the Mackenzie Slope. Piston, trigger and box cores were sampled at station 803 in 2004 aboard the CCGS Amundsen at 218 m water depth. The chronology of the piston core is constrained by 4 AMS-14C dates, as the sedimentation rate in the box core is assessed from 210Pb data. We obtain a continuous composite sequence covering the last 4600 years, with a sedimentation rate of ~ 140 cm.kyr− 1. Transfer functions (modern analogue technique) based on dinoflagellate cyst (dinocyst) assemblages were used to reconstruct the evolution of sea-surface conditions over the time period covered by the cores.
... Dinocysts are thus excellent paleoenvironmental proxies especially for high latitude environments (e.g., Rochon, 2009), with respect to calcareous (e.g., foraminifers, coccolithophores) and siliceous (e.g., diatoms) microfossils which undergo severe dissolution in cold environments de Vernal et al., 2005b). The high sedimentation rates and excellent preservation of the oxidation-sensitive taxa, such as Brigantedinium cysts (produced by the Protoperidinium species) allow us to assume that little or no selective degradation occurred at station 803. ...
Article
This study aims at reconstructing late Holocene sea-surface parameters in the Beaufort Sea area (Western Canadian Arctic) on the basis of sedimentary cores collected over the Mackenzie Slope. Piston, trigger and box cores were sampled at station 803 in 2004 aboard the CCGS Amundsen (CASES) at 218 m water depth. Sedimentation at this particular location is influenced by both the Beaufort gyre and the Mackenzie River, whose sedimentary discharge is by far the largest among all other Arctic rivers. The chronology of the piston core is constrained by 4 AMS-14C dates, as the sedimentation rate in the box core is assessed from 210Pb data. We obtain a continuous composite sequence covering the last 4600 years, with a sedimentation rate of ˜140 cm/kyr. Palynological data reveal that dinocyst assemblages are dominated by Operculodinium centrocarpum (mean of 43.3%), with the accompanying taxa Brigantedinium spp. (19.6%), Islandinium minutum (15.6%) and cysts of Pentapharsodinium dalei (13.7%). Four zones have been established on the basis of dinocyst relative abundances, with the following key taxa. Zone I (4600-4000 cal BP): high relative abundances of cysts of P. dalei (mean of 14.9%) and I. minutum var. cezare (up to 8.9%); zone II (4000-2600 cal BP): high abundances of Spiniferites frigidus/elongatus (up to 7.3%); zone III (2600-1600 cal BP): well- represented heterotrophic taxa Brigantedinium spp. (30.8%) and cysts of Polykrikos sp. arctic/quadratus (up to 4.2%); zone IV (1600 cal BP - present): high relative abundances of I. minutum (mean of 19.8%) and cysts of P. dalei (17.3%). Quantitative reconstructions of sea-surface parameters indicate relatively stable conditions from 4600 to 1600 cal BP. Conversely, a trend of increasing summer (August) sea-surface temperature (from ˜5° C to ˜6° C; actual value = 6° C), increasing salinity (from ˜20 to ˜26 psu; modern value = 19 psu) and decreasing sea-ice cover (from ˜9 to 8 month/yr; actual value = 10) is observed over the last 1600 yrs. These data are consistent with similar studies held in adjacent areas, describing the warming of Western Canadian Arctic, in comparison with a cooling Eastern Arctic. Prevailing hydroclimatic cycles in the Beaufort Sea area are also investigated on the basis of freshwater input indicators concentrations, using spectral and wavelet analysis. Potential links with the Arctic Oscillation and the Pacific Decadal Oscillation are discussed.