The dinoflagellates (Greek δῖνος dinos "whirling" and Latin flagellum "whip, scourge") are a classification subgroup of protista.They are a large group of flagellate eukaryotes that constitute the phylum Dinoflagellata. Most are marine plankton, but they also are common in freshwater habitats. Their populations are distributed depending on sea surface temperature, salinity, or depth. Many dinoflagellates are known to be photosynthetic, but a large fraction of these are in fact mixotrophic, combining photosynthesis with ingestion of prey (phagotrophy). In terms of number of species, dinoflagellates are one of the largest groups of marine eukaryotes, although this group is substantially smaller than diatoms. Some species are endosymbionts of marine animals and play an important part in the biology of coral reefs. Other dinoflagellates are unpigmented predators on other protozoa, and a few forms are parasitic (for example, Oodinium and Pfiesteria). Some dinoflagellates produce resting stages, called dinoflagellate cysts or dinocysts, as part of their lifecycles.
About 1,555 species of free-living marine dinoflagellates are currently described. Another estimate suggests about 2,000 living species, of which more than 1,700 are marine (free-living, as well as benthic) and about 220 are from fresh water. The latest estimates suggest a total of 2,294 living dinoflagellate species, which includes marine, freshwater, and parasitic dinoflagellates.
A bloom of certain dinoflagellates can result in a visible coloration of the water, colloquially known as red tide, which can cause shellfish poisoning if humans eat contaminated shellfish. Some dinoflagellates also exhibit bioluminescence—primarily emitting blue-green light.
In 1753, the first modern dinoflagellates were described by Henry Baker as "Animalcules which cause the Sparkling Light in Sea Water", and named by Otto Friedrich Müller in 1773. The term derives from the Greek word δῖνος (dinos), meaning whirling, and Latin flagellum, a diminutive term for a whip or scourge.
In the 1830s, the German microscopist Christian Gottfried Ehrenberg examined many water and plankton samples and proposed several dinoflagellate genera that are still used today including Peridinium, Prorocentrum, and Dinophysis.
These same dinoflagellates were first defined by Otto Bütschli in 1885 as the flagellate order Dinoflagellida. Botanists treated them as a division of algae, named Pyrrophyta or Pyrrhophyta ("fire algae"; Greek pyrr(h)os, fire) after the bioluminescent forms, or Dinophyta. At various times, the cryptomonads, ebriids, and ellobiopsids have been included here, but only the last are now considered close relatives. Dinoflagellates have a known ability to transform from noncyst to cyst-forming strategies, which makes recreating their evolutionary history extremely difficult.
Dinoflagellates are unicellular and possess two dissimilar flagella arising from the ventral cell side (dinokont flagellation). They have a ribbon-like transverse flagellum with multiple waves that beats to the cell's left, and a more conventional one, the longitudinal flagellum, that beats posteriorly. The transverse flagellum is a wavy ribbon in which only the outer edge undulates from base to tip, due to the action of the axoneme which runs along it. The axonemal edge has simple hairs that can be of varying lengths. The flagellar movement produces forward propulsion and also a turning force. The longitudinal flagellum is relatively conventional in appearance, with few or no hairs. It beats with only one or two periods to its wave. The flagella lie in surface grooves: the transverse one in the cingulum and the longitudinal one in the sulcus, although its distal portion projects freely behind the cell. In dinoflagellate species with desmokont flagellation (e.g., Prorocentrum), the two flagella are differentiated as in dinokonts, but they are not associated with grooves.
Dinoflagellates have a complex cell covering called an amphiesma or cortex, composed of a series of membranes, flattened vesicles called alveolae (= amphiesmal vesicles) and related structures. In armoured dinoflagellates, these support overlapping cellulose plates to create a sort of armor called the theca or lorica, as opposed to athecate dinoflagellates. These occur in various shapes and arrangements, depending on the species and sometimes on the stage of the dinoflagellate. Conventionally, the term tabulation has been used to refer to this arrangement of thecal plates. The plate configuration can be denoted with the plate formula or tabulation formula. Fibrous extrusomes are also found in many forms. Together with various other structural and genetic details, this organization indicates a close relationship between the dinoflagellates, the Apicomplexa, and ciliates, collectively referred to as the alveolates.
Dinoflagellate tabulations can be grouped into six "tabulation types": gymnodinoid, suessoid, gonyaulacoid–peridinioid, nannoceratopsioid, dinophysioid, and prorocentroid.
The chloroplasts in most photosynthetic dinoflagellates are bound by three membranes, suggesting they were probably derived from some ingested algae. Most photosynthetic species contain chlorophylls a and c2, the carotenoid beta-carotene, and a group of xanthophylls that appears to be unique to dinoflagellates, typically peridinin, dinoxanthin, and diadinoxanthin. These pigments give many dinoflagellates their typical golden brown color. However, the dinoflagellates Karenia brevis, Karenia mikimotoi, and Karlodinium micrum have acquired other pigments through endosymbiosis, including fucoxanthin. This suggests their chloroplasts were incorporated by several endosymbiotic events involving already colored or secondarily colorless forms. The discovery of plastids in the Apicomplexa has led some to suggest they were inherited from an ancestor common to the two groups, but none of the more basal lines has them. All the same, the dinoflagellate cell consists of the more common organelles such as rough and smooth endoplasmic reticulum, Golgi apparatus, mitochondria, lipid and starch grains, and food vacuoles. Some have even been found with a light-sensitive organelle, the eyespot or stigma, or a larger nucleus containing a prominent nucleolus. The dinoflagellate Erythropsidium has the smallest known eye.
Some athecate species have an internal skeleton consisting of two star-like siliceous elements that has an unknown function, and can be found as microfossils. Tappan gave a survey of dinoflagellates with internal skeletons. This included the first detailed description of the pentasters in Actiniscus pentasterias, based on scanning electron microscopy. They are placed within the order Gymnodiniales, suborder Actiniscineae.
The formation of thecal plates has been studied in detail through ultrastructural studies.
'Core dinoflagellates' (dinokaryotes) have a peculiar form of nucleus, called a dinokaryon, in which the chromosomes are attached to the nuclear membrane. These carry reduced number of histones. In place of histones, dinoflagellate nuclei contain a novel, dominant family of nuclear proteins that appear to be of viral origin, thus are called dinoflagellate/ viral nucleoproteins (DVNPs) which are highly basic, bind DNA with similar affinity to histones, and occur in multiple posttranslationally modified forms. Dinoflagellate nuclei remain condensed throughout interphase rather than just during mitosis, which is closed and involves a uniquely extranuclear mitotic spindle. This sort of nucleus was once considered to be an intermediate between the nucleoid region of prokaryotes and the true nuclei of eukaryotes, so were termed mesokaryotic, but now are considered advanced rather than primitive traits. In addition to dinokaryotes, DVNPs can be found in a group of basal dinoflagellates (known as Marine Alveolates, "MALVs") that branch as sister to dinokaryotes (Syndiniales).
Dinoflagellates are protists which have been classified using both the International Code of Botanical Nomenclature (ICBN, now renamed as ICN) and the International Code of Zoological Nomenclature (ICZN). About half of living dinoflagellate species are autotrophs possessing chloroplasts and half are nonphotosynthesising heterotrophs.
The peridinin dinoflagellates, named after their peridinin plastids, appears to be ancestral for the dinoflaggelate lineage. Almost half of all known species have chloroplasts, which are either the original peridinin plastids or new plastids acquired from other lineages of unicellular algae through endosymbiosis. The remaining species have lost their photosynthetic abilities and have adapted to a heterotrophic, parasitic or kleptoplastic lifestyle.
Most (but not all) dinoflagellates have a dinokaryon, described below (see: Life cycle, below). Dinoflagellates with a dinokaryon are classified under Dinokaryota, while dinoflagellates without a dinokaryon are classified under Syndiniales.
Although classified as eukaryotes, the dinoflagellate nuclei are not characteristically eukaryotic, as some of them lack histones and nucleosomes, and maintain continually condensed chromosomes during mitosis. The dinoflagellate nucleus was termed ‘mesokaryotic’ by Dodge (1966), due to its possession of intermediate characteristics between the coiled DNA areas of prokaryotic bacteria and the well-defined eukaryotic nucleus. This group, however, does contain typically eukaryotic organelles, such as Golgi bodies, mitochondria, and chloroplasts.
Jakob Schiller (1931–1937) provided a description of all the species, both marine and freshwater, known at that time. Later, Alain Sournia (1973, 1978, 1982, 1990, 1993) listed the new taxonomic entries published after Schiller (1931–1937). Sournia (1986) gave descriptions and illustrations of the marine genera of dinoflagellates, excluding information at the species level. The latest index is written by Gómez.
English-language taxonomic monographs covering large numbers of species are published for the Gulf of Mexico, the Indian Ocean, the British Isles, the Mediterranean and the North Sea.
The main source for identification of freshwater dinoflagellates is the Süsswasser Flora.
Dinoflagellates can occur in all aquatic environments: marine, brackish, and fresh water, including in snow or ice. They are also common in benthic environments and sea ice.
All Zooxanthellae are dinoflagellates and most of them are members within the genus Symbiodinium. The association between Symbiodinium and reef-building corals is widely known. However, endosymbiontic Zooxanthellae inhabit a great number of other invertebrates and protists, for example many sea anemones, jellyfish, nudibranchs, the giant clam Tridacna, and several species of radiolarians and foraminiferans. Many extant dinoflagellates are parasites (here defined as organisms that eat their prey from the inside, i.e. endoparasites, or that remain attached to their prey for longer periods of time, i.e. ectoparasites). They can parasitize animal or protist hosts. Protoodinium, Crepidoodinium, Piscinoodinium, and Blastodinium retain their plastids while feeding on their zooplanktonic or fish hosts. In most parasitic dinoflagellates, the infective stage resembles a typical motile dinoflagellate cell.
Three nutritional strategies are seen in dinoflagellates: phototrophy, mixotrophy, and heterotrophy. Phototrophs can be photoautotrophs or auxotrophs. Mixotrophic dinoflagellates are photosynthetically active, but are also heterotrophic. Facultative mixotrophs, in which autotrophy or heterotrophy is sufficient for nutrition, are classified as amphitrophic. If both forms are required, the organisms are mixotrophic sensu stricto. Some free-living dinoflagellates do not have chloroplasts, but host a phototrophic endosymbiont. A few dinoflagellates may use alien chloroplasts (cleptochloroplasts), obtained from food (kleptoplasty). Some dinoflagellates may feed on other organisms as predators or parasites.
Mechanisms of capture and ingestion in dinoflagellates are quite diverse. Several dinoflagellates, both thecate (e.g. Ceratium hirundinella, Peridinium globulus) and nonthecate (e.g. Oxyrrhis marina, Gymnodinium sp. and Kofoidinium spp.), draw prey to the sulcal region of the cell (either via water currents set up by the flagella or via pseudopodial extensions) and ingest the prey through the sulcus. In several Protoperidinium spp., e.g. P. conicum, a large feeding veil — a pseudopod called the pallium — is extruded to capture prey which is subsequently digested extracellularly (= pallium-feeding). Oblea, Zygabikodinium, and Diplopsalis are the only other dinoflagellate genera known to use this particular feeding mechanism. Katodinium (Gymnodinium) fungiforme, commonly found as a contaminant in algal or ciliate cultures, feeds by attaching to its prey and ingesting prey cytoplasm through an extensible peduncle. Two related species, polykrikos kofoidii and neatodinium, shoots out a harpoon-like organelle to capture prey. The feeding mechanisms of the oceanic dinoflagellates remain unknown, although pseudopodial extensions were observed in Podolampas bipes.
Dinoflagellate blooms are generally unpredictable, short, with low species diversity, and with little species succession. The low species diversity can be due to multiple factors. One way a lack of diversity may occur in a bloom is through a reduction in predation and a decreased competition. The first may be achieved by having predators reject the dinoflagellate, by, for example, decreasing the amount of food it can eat. This additionally helps prevent a future increase in predation pressure by cause predators that reject it to lack the energy to breed. A species can then inhibit the growth of its competitors, thus achieving dominance.
Dinoflagellates sometimes bloom in concentrations of more than a million cells per millilitre. Under such circumstances, they can produce toxins (generally called dinotoxins) in quantities capable of killing fish and accumulating in filter feeders such as shellfish, which in turn may be passed on to people who eat them. This phenomenon is called a red tide, from the color the bloom imparts to the water. Some colorless dinoflagellates may also form toxic blooms, such as Pfiesteria. Some dinoflagellate blooms are not dangerous. Bluish flickers visible in ocean water at night often come from blooms of bioluminescent dinoflagellates, which emit short flashes of light when disturbed.
The same red tide mentioned above is more specifically produced when dinoflagellates are able to reproduce rapidly and copiously on account of the abundant nutrients in the water. Although the resulting red waves are an unusual sight, they contain toxins that not only affect all marine life in the ocean, but the people who consume them, as well. A specific carrier is shellfish. This can introduce both nonfatal and fatal illnesses. One such poison is saxitoxin, a powerful paralytic neurotoxin. Human inputs of phosphate further encourage these red tides, so strong interest exists in learning more about dinoflagellates, from both medical and economic perspectives. The ecology of harmful algal blooms is extensively studied.
At night, water can have an appearance of sparkling light due to the bioluminescence of dinoflagellates. More than 18 genera of dinoflagellates are bioluminescent, and the majority of them emit a blue-green light. These species contain scintillons, individual cytoplasmic bodies (about 0.5 µm in diameter) distributed mainly in the cortical region of the cell, outpockets of the main cell vacuole. They contain dinoflagellate luciferase, the main enzyme involved in dinoflagellate bioluminescence, and luciferin, a chlorophyll-derived tetrapyrrole ring that acts as the substrate to the light-producing reaction. The luminescence occurs as a brief (0.1 sec) blue flash (max 476 nm) when stimulated, usually by mechanical disturbance. Therefore, when mechanically stimulated—by boat, swimming, or waves, for example—a blue sparkling light can be seen emanating from the sea surface.
Dinoflagellate bioluminescence is controlled by a circadian clock and only occurs at night. Luminescent and nonluminescent strains can occur in the same species. The number of scintillons is higher during night than during day, and breaks down during the end of the night, at the time of maximal bioluminescence.
The luciferin-luciferase reaction responsible for the bioluminescence is pH sensitive. When the pH drops, luciferase changes its shape, allowing luciferin, more specifically tetrapyrrole, to bind. Dinoflagellates can use bioluminescence as a defense mechanism. They can startle their predators by their flashing light or they can ward off potential predators by an indirect effect such as the "burglar alarm". The bioluminescence attracts attention to the dinoflagellate and its attacker, making the predator more vulnerable to predation from higher trophic levels.
Bioluminescent dinoflagellate ecosystem bays are among the rarest and most fragile, with the most famous ones being the Bioluminescent Bay in La Parguera, Lajas, Puerto Rico; Mosquito Bay in Vieques, Puerto Rico; and Las Cabezas de San Juan Reserva Natural Fajardo, Puerto Rico. Also, a bioluminescent lagoon is near Montego Bay, Jamaica, and bioluminescent harbors surround Castine, Maine. Within the United States, Central Florida is home to the Indian River Lagoon which is abundant with dinoflagellates in the summer and bioluminescent ctenophore in the winter. There are a total of 5 bioluminescence bays in the world and 3 of them are in Puerto Rico .
Dinoflagellates have a haplontic life cycle, with the possible exception of Noctiluca and its relatives. The life cycle usually involves asexual reproduction by means of mitosis, either through desmoschisis or eleuteroschisis. More complex life cycles occur, more particularly with parasitic dinoflagellates. Sexual reproduction also occurs, though this mode of reproduction is only known in a small percentage of dinoflagellates. This takes place by fusion of two individuals to form a zygote, which may remain mobile in typical dinoflagellate fashion and is then called a planozygote. This zygote may later form a resting stage or hypnozygote, which is called a dinoflagellate cyst or dinocyst. After (or before) germination of the cyst, the hatchling undergoes meiosis to produce new haploid cells.
One of their most striking features is the large amount of cellular DNA that dinoflagellates contain. Most eukaryotic algae contain on average about 0.54 pg DNA/cell, whereas estimates of dinoflagellate DNA content range from 3–250 pg/cell, corresponding to roughly 3000–215 000 Mb (in comparison, the haploid human genome is 3180 Mb and hexaploid Triticum wheat is 16 000 Mb). Polyploidy or polyteny may account for this large cellular DNA content, but studies of DNA reassociation kinetics do not support this hypothesis.
In addition to their disproportionately large genomes, dinoflagellate nuclei are unique in their morphology, regulation, and composition. Their DNA is so tightly packed it is still uncertain exactly how many chromosomes they have.
The dinoflagellates share an unusual mitochondrial genome organisation with their relatives, the Apicomplexa. Both groups have very reduced mitochondrial genomes (around 6 kilobases (kb) in the Apicomplexa vs ~16kb for human mitochondria). One species, Amoebophrya ceratii, has lost its mitochondrial genome completely, yet still has functional mitochondria. The genes on the dinoflagellate genomes have undergone a number of reorganisations, including massive genome amplification and recombination which have resulted in multiple copies of each gene and gene fragments linked in numerous combinations. Loss of the standard stop codons, trans-splicing of mRNAs for the mRNA of cox3, and extensive RNA editing recoding of most genes has occurred. The reasons for this transformation are unknown. In a small group of dinoflagellates, called ‘dinotoms’ (Durinskia and Kryptoperidinium), the endosymbionts (diatoms) still have mitochondria, making them the only organisms with two evolutionarily distinct mitochondria.
In most of the species, the plastid genome consist of just 14 genes.
The DNA of the plastid in the peridinin-containing dinoflagellates is contained in a series of small circles. Each circle contains one or two polypeptide genes. The genes for these polypeptides are chloroplast-specific because their homologs from other photosynthetic eukaryotes are exclusively encoded in the chloroplast genome. Within each circle is a distinguishable 'core' region. Genes are always in the same orientation with respect to this core region.
In terms of DNA barcoding, ITS sequences can be used to identify species, where a genetic distance of p≥0.04 can be used to delimit species. A recent study revealed a substantial proportion of dinoflagellate genes encode for unknown functions, and that these genes could be conserved and lineage-specific.
Dinoflagellates are mainly represented as fossils by fossil dinocysts, which have a long geological record with lowest occurrences during the mid-Triassic, whilst geochemical markers suggest a presence to the Early Cambrian.
Molecular phylogenetics show that dinoflagellates are grouped with ciliates and apicomplexans (=Sporozoa) in a well-supported clade, the alveolates. The closest relatives to dinokaryotic dinoflagellates appear to be apicomplexans, Perkinsus, Parvilucifera, syndinians, and Oxyrrhis. Molecular phylogenies are similar to phylogenies based on morphology.
All dinoflagellates contain red algal plastids or remnant (nonphotosynthetic) organelles of red algal origin. The parasitic dinoflagellate Hematodinium however lacks a plastid entirely. Some groups that have lost the photosynthetic properties of their original red algae plastids has obtained new photosynthetic plastids (chloroplasts) through so-called serial endosymbiosis, both secondary and tertiary. Like their original plastids, the new chloroplasts in these groups can be traced back to red algae, except from those in the members of the genus Lepidodinium, which possess plastids derived from green algae, possibly Trebouxiophyceae or Ulvophyceae. Lineages with tertiary endosymbiosis are Dinophysis, with plastids from a cryptomonad, the Karenia, Karlodinium, and Takayama, which possess plastids of haptophyte origin, and the Peridiniaceae, Durinskia and Kryptoperidinium, which has plastids derived from diatoms Some species also perform kleptoplasty.
Dinoflagellate evolution has been summarized into five principal organizational types: prorocentroid, dinophysoid, gonyaulacoid, peridinioid, and gymnodinoid. The transitions of marine species into fresh water have been infrequent events during the diversification of dinoflagellates and in most cases have not occurred recently, possibly as late as the Cretaceous.
Dinodnavirus is a genus of viruses that infect dinoflagellates. This genus belongs to the clade of nucleocytoplasmic large DNA viruses. The name is derived from 'dino' (dinoflagellate) and DNA (from its genome).
The type species is Heterocapsa circularisquama DNA virus 01.Dinokaryota
Dinokaryota is a main grouping of dinoflagellates. They include all species where the nucleus remains a dinokaryon throughout the entire cell cycle, which is typically dominated by the haploid stage. All the "typical" dinoflagellates, such as Peridinium and Gymnodinium, belong here. Others are more unusual, including some that are colonial, amoeboid, or parasitic. Symbiodinium contains the symbiotic zooxanthellae.
The non-photosynthetic members are believed to derive from photosynthetic ancestors.Duboscquella
Duboscquella is a genus of dinoflagellates.Gonyaulacales
The Gonyaulacales are an order of dinoflagellates found in marine environments.Gonyaulax
Gonyaulax is a genus of dinoflagellates with the type species Gonyaulax spinifera (Claparède et Lachmann) Diesing. Gonyaulax belongs to red dinoflagellates and commonly causes red tides.Gymnodiniales
The Gymnodiniales are an order of dinoflagellates, of the class Dinophyceae. Members of the order are known as gymnodinioid or gymnodinoid (terms that can also refer to any organism of similar morphology). They are athecate, or lacking an armored exterior, and as a result are relatively difficult to study because specimens are easily damaged. Many species are part of the marine plankton and are of interest primarily due to being found in algal blooms. As a group the gymnodinioids have been described as "likely one of the least known groups of the open ocean phytoplankton."Of the families in the order, the Polykrikaceae and Warnowiaceae are well known for possessing exceptionally complex assemblies of organelles, such as nematocysts, trichocysts, and pistons. The Warnowiaceae uniquely possess an ocelloid, an extremely complex light-sensitive subcellular structure composed of mitochondria and plastids.Gymnodinium
Gymnodinium is a genus of dinoflagellates, a type of marine plankton. It is one of the few naked dinoflagellates, or species lacking armor (cellulosic plates). Since 2000, the species which had been considered to be part of Gymnodinium have been divided into several genera, based on the nature of the apical groove and partial LSU rDNA sequence data. Amphidinium was redefined later. Gymnodinium belong to red dinoflagellates that, in concentration, can cause red tides.
Gymnodinium sensu stricto
Histioneis is a genus of dinoflagellates.Karenia (dinoflagellate)
Karenia is a genus that consists of unicellular, photosynthetic, planktonic organisms found in marine environments. The genus currently consists of 12 described species. They are best known for their dense toxic algal blooms and red tides that cause considerable ecological and economical damage; some Karenia species cause severe animal mortality. One species, Karenia brevis, is known to cause respiratory distress and neurotoxic shellfish poisoning (NSP) in humans.Karlodinium
Karlodinium is a genus of dinoflagellates.Noctilucales
The Noctilucales are an order of marine dinoflagellates. They differ from most others in that the mature cell is diploid and its nucleus does not show a dinokaryotic organization. They show gametic meiosis.Ornithocercus
Ornithocercus is a genus of planktonic dinoflagellate that is known for its complex morphology that features considerable lists growing from its thecal plates, giving an attractive appearance. Discovered in 1883, this genus has a small number of species currently categorized but is widespread in tropical and sub-tropical oceans. The genus is marked by exosymbiotic bacteria gardens under its lists, the inter-organismal dynamics of which are a current field of research. As they reside only in warm water, the genus has been used as a proxy for climate change and has potential to be an indicator species for environmental change if found in novel environments.Oxyphysis
Oxyphysis is a genus of dinoflagellates.
It includes the species Oxyphysis oxytoides.Oxyrrhis
Oxyrrhis is a genus of dinoflagellates. It includes the species Oxyrrhis marina.It is sometimes considered to be a monotypic genus. Some sources assign the species Oxyrrhis parasitica and O. phaeocysticola, as well.Peridiniales
Peridiniales is an order of dinoflagellates. Below is the list of the families in Peridinales.
Heterocapsaceae syn. Heterocapsineae
Peridinium is a genus of motile, marine and freshwater dinoflagellates. Their morphology is considered typical of the armoured dinoflagellates, and their form is commonly used in diagrams of a dinoflagellate's structure. Peridinium can range from 30-70 μm in diameter, and has very thick thecal plates.Polarella
Polarella is a genus of dinoflagellates.
It includes the species Polarella glacialis.Suessiales
Suessiales is an order of dinoflagellates.It includes Polarella and Symbiodinium.Syndiniales
The Syndiniales are an order of early branching dinoflagellates (also known as Marine Alveolates, "MALVs"), found as parasites of crustaceans, fish, algae, cnidarians, and protists (ciliates, radiolarians, other dinoflagellates). The trophic form is often multinucleate, and ultimately divides to form motile spores, which have two flagella in typical dinoflagellate arrangement. They lack a theca and chloroplasts, and unlike all other orders, the nucleus is never a dinokaryon. A well-studied example is Amoebophrya, which is a parasite of other dinoflagellates and may play a part in ending red tides. Several MALV groups have been assigned to Syndiniales; recent studies, however, show paraphyly of MALVs suggesting that only those groups that branch as sister to dinokaryotes ('core dinoflagellates') belong to Syndiniales.
Extant life phyla/divisions by domain