THREE NEW BENTHIC SPECIES OF PROROCENTRUM (DINOPHYCEAE) FROM BELIZE: P. NORRISIANUM SP. NOV., P. TROPICALIS SP. NOV., and P. RETICULATUM SP. NOV.1

Three new benthic, photosynthetic dinoflagellate species, Prorocentrum norrisianum, Prorocentrum tropicalis, and Prorocentrum reticulatum, from floating detritus and coral rubble of Central America are described from scanning electron micrographs. Species were identified based on shape, size, surface micromorphology, thecal plate ornamentation, and architecture of the periflagellar area and intercalary band. Cells of P. norrisianum are ovate with a cell size of 20–25 μm long and 13–16 μm wide. The theca is delicate, its surface smooth, pores species specific with 95 to 105 pores per valve. Pores are round with a diameter of about 0.1 μm. The periflagellar area is V-shaped, located on the right valve in a shallow depression. It has no ornamentation. The flagellar and auxiliary pores are unequal in size. The intercalary band is smooth. Prorocentrum tropicalis cells are ovoid, 50–55 μm long and 40–45 μm wide in valve view with maximum width behind the middle region, narrow at the anterior end. The periflagellar area, situated in the right valve, is a V-shaped wide triangle with a deeply indented depression; the left valve exhibits a flat ridge. The periflagellar area is unornamented, and the flagellar and auxiliary pores are unequal in size. The valve surface is rugose with evenly distributed valve poroids. Each poroid appears to have a small dome in the center. The intercalary band is rimlike around the cell margin, granulated, and horizontally striated. Prorocentrum reticulatum cells are oblong in valve view; cells are 55–60 μm long and 40–45 μm wide. Thecal surface is reticulated; it is composed of a labyrinth of ridges with alternating depressions that vary in size and shape. Each depression has a narrow, oblong-kidney-shaped opening about 0.6 μm long. The periflagellar area is a deep, V-shaped triangle. The right valve of P. reticulatum is excavated, and contains a large flagellar pore and a smaller auxiliary pore surrounded by a narrow apical collar. The left valve margin exhibits a curved flat ridge. The intercalary band is smooth.     

THREE NEW BENTHIC SPECIES OF PROROCENTRUM (DINOPHYCEAE) FROM CARRIE BOW CAY,BELIZE: P. SABULOSUM SP. NOV., P. SCULPTILE SP. NOV., AND P. ARENARIUM SP. NOV.1

Three new benthic, sand-dwelling dinqflagellate species, Prorocentrum sabulosum, Prorocentrum scuptile, and Prorocentrum arenarium, from coral rubble are described from scanning electron micrographs. Species were identified based on shape, size, surface micromorphology, ornamentation of thecal plates, and architecture of the periflagellar area and intercalary band. Cells of P. sabulosum are oval with a cell size of 48–50 μm long and 41–48 μm wide. The areolae are round to oval and numerous (332–450 per valve) and range from 1 to 1.6 μm in size. The periflagellar area of P. sabulosum bears a wide V-shaped depression with a flat ridge and lacks ornamentation; it accommodates six pores: one large flagellar pore, an adjacent smaller auxiliary pore, and four pores of unknown function. The flagellar and auxiliary pores are surrounded by a narrow apical collar. The intercalary band of P. sabulosum is smooth. Prorocentrum sculptile cells are broadly oval, 32–37 nm long, and 30–32 μm wide in valve view with a deep-sculptured apical area. The valves are smooth and are marked with shallow depressions (856–975 per valve). Some of these depressions have a small round opening (0.13 μm in diameter). The periflagellar area is V-shaped with a deeply indented depression; it accommodates the two flagella and a thin angled apical plate. The intercalary band is smooth. Prorocentrum arenarium cells are nearly round in valve view 30–32 μm in diameter. Thecal surface is smooth with scattered kidney-shaped valve poroids (65–73 per valve) and marginal poroids (50–57 per valve). Length and width of poroids are 0.62 μm and 0.36 μm, respectively. The periflagellar area is an unornamented, broad triangle into which a large flagellar pore and a smaller auxiliary pore are fitted. Both flagella, longitudinal and transverse, protrude from the flagellar pore. The intercalary band is smooth. The presence of a peduncle-like structure (2–3 μm long) in P. arenarium was observed situated in the flagellar pore.     

OBSERVATION OF SAND-DWELLING TOXIC DINOFLAGELLATES (DINOPHYCEAE) FROM WIDELY DIFFERING SITES, INCLUDING TWO NEW SPECIES

Dinoflagellate associations, including toxic and potentially toxic benthic species, were examined in sand from South Water Cay and Carrie Bow Cay, Belize. The inshore sand habitat in localized areas of warm shallow lagoonal waters supported blooms of toxic assemblages of dinoflagellates. In the sand, the dominant microalgae were dinoflagellates; cyanobacteria were a minor component and diatoms were absent. Ciliates and nematodes were present. Assemblages of microorganisms in colored sand were examined for 4 consecutive days after which a storm washed away the patch. The sand-dwelling dinoflagellate assemblage included 16 species where densities ranged from as low as 1.3% to 15% of total cell densities. The dominant species was Scrippsiella subsalsa, having 1.8 × 10⁵ to 2.6 × 10⁵ cells g^-1 sand. Toxic dinoflagellates identified in the sand were Gambierdiscus toxicus, Ostreopsis lenticularis, Prorocentrum lima, Prorocentrum mexicanum, and Amphidinium carteri. The potentially toxic Ostreopsis labens, Gambierdiscus belizeanussp. nov., and Coolia tropicalis sp. nov. were also identified. Toxic and potentially toxic species represented 36% to 60% of total microalgal cell assemblage. The morphology of a new sand-dwelling species, Gambierdiscus belizeanus sp. nov., was examined with the scanning electron microscope. The plate formula was P_o, 3′, 7″, 6c, s?, 5?, 1p, and 2″″.Dimensions of G. belizeanus cells were 53–67 pm long, 54–63 μm wide, and 92–98 μm in dorsoventral depth. Cells were deeply areolated, ellipsoid in apical view, and compressed anteroposteriorly. The cells of G. belizeanus were identified by the cell's long, narrow, pentagonal, posterior intercalary plate (1p) wedged between the wide postcingular plates 2″’and 4″; 1p occupied 20% of the width of the hypotheca. The plate formula for Coolia tropicalis sp. nov. was P_o, 3′, 7″, 7c, 8s?, 5″″, and 2″″, Cell size ranges were 23–40 μm long, 25–39 μm wide, and 35–65 μm in dorsoventral diameter. Cells were spherical, smooth, and covered with scattered round pores. The epitheca was smaller than the hypotheca. Precingular plates 1″ and 7″ were small and narrow, and the first apical plate 1″ and precingular plate 6″ were the largest plates on the epitheca. The apical pore was straight and 7 μm long, and was situated in the apical plate complex. Cells of C. tropicalis were distinguished from C. monotis by the wedge-shaped plate 1′, a four-sided 3’plate, and a short apical pore.     

PROROCENTRUM BELIZEANUM,PROROCENTRUM ELEGANS,AND PROROCENTRUM CARIBBAEUM,THREE NEW BENTHIC SPECIES (DINOPHYCEAE) FROM A MANGROVE ISLAND,TWIN CAYS,BELIZE1

Three new benthic dinoflagellate species, Prorocentrum belizeanum, Prorocentrum elegans, and Prorocentrum caribbaeum, from mangrove floating detritus are described from scanning electron micrographs. Species were identified based on shape, size, surface micromorphology, ornamentation of thecal plates, and architecture of the periflagellar area and intercalary band. Cells of P. belizeanum are round to slightly oval with a cell size of 55–60 μm long and 50–55 μm wide. Areolae are round and numerous (853–1024 per valve) and range from 0.66 to 0.83 μm in size. The periflagellar area of P. belizeanum is a broad V-shaped depression; it accommodates a flagellar and an auxiliary pore and a flared, curved apical collar. The intercalary band of P. belizeanum is horizontally striated. Prorocentrum elegans is a small species 15–20 μm long and 10–14 μm wide, with an ovate cell shape. The thecal surface is smooth. Two sizes of valve pores were recognized: large, round pores (20–22 per valve) arranged in a distinct pattern and smaller pores situated in an array along the intercalary band. The periflagellar area is V-shaped; it accommodates an uneven sized flagellar pore, an auxiliary pore, and an angled protuberant flagellar plate. The intercalary band is transversely striated. It is a bloom-forming species. Prorocentrum caribbaeum cells are heart-shaped with a rounded anterior end and a pointed posterior end. Cells range from 40 to 45 μm long and 30 to 35 μm wide. Thecal surface has two different-sized pores: large, round pores (145–203 per valve) arranged perpendicularly from the posterior margins, and small, round pores unevenly distributed on the thecal surface. The periflagellar area is ornate. It is V-shaped with a curved apical collar located next to the auxiliary pore; a smaller protuberant apical plate is adjacent to the flagellar pore. The intercalary band is transversely striated and sinuous. Cells are active swimmers.     

MORPHOLOGY AND MOLECULAR PHYLOGENY OF PROROCENTRUM CONSUTUM SP. NOV. (DINOPHYCEAE), A NEW BENTHIC DINOFLAGELLATE FROM SOUTH BRITTANY (NORTHWESTERN FRANCE)1

A new marine benthic Prorocentrum species from sandy habitats of South Brittany (northwestern France), P. consutum sp. nov., is described using LM and SEM and molecular phylogenetic analyses. Cells have a subcircular to broadly ovoid shape and are plainly flattened. They are 57–61 μm long and 52–55 μm wide. A central pyrenoid is present, and the kidney‐shaped nucleus is positioned in the posterior region. In right valve view, the periflagellar area is deeply excavated, and the left valve forms a prominent apical ridge. The periflagellar area consists of nine platelets, and a small narrow collar is present around the flagellar pore. The ornamentation of this new species is very peculiar and is characterized by a ring of round areolae located at the periphery of the valves, each areola containing three or four pores. Apart from this ring of areolae, the cell surface is smooth and with scattered pores. Pores are not present in the center of the right or left valve. The intercalary band is generally narrow and faintly striated horizontally. The molecular phylogenetic position of P. consutum sp. nov. was inferred using SSU and LSU rDNA. In both analyses, this species branched with high support in the clade comprising species with a symmetric shape and appeared to be a sister group to that formed by P. lima and other tropical benthic species, such as P. arenarium, P. belizeanum, P. hoffmannianum, and P. maculosum.     

MORPHOLOGY AND ECOLOGY OF THE MARINE DINOFLAGELLATE OSTREOPSIS LABENS SP. NOV. (DINOPHYCEAE)1

A new species, Ostreopsis labens Faust et Morton sp. nov., is described from three marine habitats: lagoonal water and lagoonal sand from the barrier reef of Belize, and associated with macroalgae from coral reef habitats of Oshigaki and Iriomote Islands, Japan. Dimensions of Ostreopsis labens cells are 60–86 μm long, 70–80 μm wide, and 81–110 μm in dorsoventral depth. Cells are broadly ovoid, anterioposteriorly compressed bearing a spherical nucleus and many chloroplasts. The epitheca is convex and composed of three apical plates, seven precingular plates, and an apical pore plate. The cingulum is composed of six plates. The hypotheca is constructed of five postcingular plates, one posterior intercalary, and two antapical plates. The sulcus is small, recessed, and hidden and exhibits a ventral pore and a ridged, curved plate. The thecal arrangement of O. labens is P_o, 3′, 7″ 6C, 6S(?), V_p, R_p, 5″, 1p, 2″. Only one sulcal list is present. The thecal plates have a smooth surface with distinct round pores. The intercalary band between the thecal plates is smooth. A row of marginal pores line the lipped cingulum. Ostreopsis species are anteroposteriorly flattened, photosynthetic, benthic dinoflagellates that are more diverse in ecology than previously known. Ostreopsis labens is capable of living in three marine habitats: in the water column, in sand, and on macroalgal surfaces. It was most numerous in sand and less in lagoonal waters, and only a few cells were associated with macroalgae. Light and scanning electron microscopy studies revealed engulfed cells within O. labens, which indicates mixotrophic/phagotrophic behavior. A ventral opening situated in the cingulum of O. labens exhibits size variability; it may serve as an opening for engulfiing food particles because it varies in size. We propose that ingestion of prey by O. labens occurs through the ventral opening, the proposed feeding apparatus of this species, which is similar to the function of the peduncle-like structure of mixotrophic dinoflagellates. The behavior of O. labens appears similar to that previously described for Dinophysis species.     

PROROCENTRUM BIMACULATUM SP. NOV. (DINOPHYCEAE,PROROCENTRALES), A NEW BENTHIC DINOFLAGELLATE SPECIES FROM KUWAIT (ARABIAN GULF)1

A new benthic dinoflagellate species, Prorocentrum bimaculatum sp. nov., is studied from Kuwait’s marine sediments, based on detailed morphological and molecular data. Cells are large, oblong oval in shape. They are 49.9–55.3 μm long and 38.4–43.2 μm wide. The ornamentation of this new species is peculiar, and characterized by smooth valves with large pores (0.32–0.50 μm) scattered on their surface, except in two circular patches of ～15 μm in diameter, devoid of ornamentation and located on both sides of the valve centers. The periflagellar area is widely triangular, located in a moderate excavation of the right valve, and comprises nine platelets. The intercalary band of P. bimaculatum is smooth. The molecular phylogenetic position of this new taxon was inferred from SSU and LSU rDNA genes. In both phylogenetic analyses, P. bimaculatum branched with high support with Prorocentrum consutum and formed a clade sister to the one including P. lima and related species such as P. arenarium, P. belizeanum, P. hoffmannianum, and P. maculosum. From the phylogenetic study, since most species related to P. bimaculatum are known for their toxic effects and production of okadaic acid, this new species can be considered as a potential toxin producer, but this has to be analyzed.     

PROTOPERIDINIUM BELIZEANUM SP. NOV. (DINOPHYCEAE) FROM MANATEE CAY,BELIZE, CENTRAL AMERICA1

A new marine heterotrophic dinoflagellate species, Protoperidinium belizeanum sp. nov., from a coral reef‐mangrove pond was identified from scanning electron micrographs. Recognition of this new species was based on unique features of the thecal morphology, which included cell size and shape, presence of short and wide postcingular plates, sulcal architecture, antapical spines, and intricate thecal plate patterns of ridged hexagonal depressions. The thecal plate formula is as follows: Po, X, 4′, 3a, 7″, 4C (3+t), 6S, 5?, 2″″. Species association of P. be‐lizeanum sp. nov. within the genus Protoperidinium, its habitat, and associated dinoflagellates species are discussed.     

MORPHOLOGY,MOLECULAR PHYLOGENY AND TAXONOMY OF TWO NEW SPECIES OF PLEODORINA (VOLVOCEAE,CHLOROPHYCEAE)1

The volvocacean genus Pleodorina has been morphologically characterized as having small somatic cells in spheroidal colonies and anisogamous sexual reproduction with sperm packets. In this study we examined two new species that can be assigned to the genus Pleodorina based on morphology: P. starrii H. Nozaki et al. sp. nov. and P. thompsonii F. D. Ott et al. sp. nov. P. starrii was collected from Japan and had 32‐ or 64‐celled colonies with anterior somatic cells and spheroidal individual cellular sheaths that were weakly attached to each other within the colonial envelope. P. thompsonii from Texas (USA) exhibited four or 12 somatic cells in the anterior pole of 16‐ or 32‐celled colonies, respectively, and had a single large pyrenoid in the chloroplast of mature reproductive cells. The chloroplast multigene phylogeny placed P. starrii and P. indica (Iyenger) H. Nozaki in a clade that was robustly separated from the type species P. californica Shaw and P. japonica H. Nozaki. Pleodorina thompsonii was resolved as a basal branch within a large monophyletic group (Eudorina group) composed of Eudorina, Pleodorina and Volvox (excluding section Volvox). Thus, Pleodorina was found among three separate lineages within the Eudorina group in which Eudorina and Volvox were also resolved as nonmonophyletic. The DNA sequences from additional species/strains as well as recognition of morphological attributes that characterize the monophyletic groups within the Eudorina group are needed to construct a natural generic classification within these members of the Volvocaceae.     

MORPHOLOGY AND MOLECULAR PHYLOGENY OF A NEW MARINE SAND‐DWELLING PROROCENTRUM SPECIES,P. TSAWWASSENENSE (DINOPHYCEAE,PROROCENTRALES), FROM BRITISH COLUMBIA,CANADA1

A new marine benthic, sand‐dwelling Prorocentrum species from the temperate region of the Pacific coast of British Columbia, Canada, is described using LM and EM and molecular phylogenetic analyses. The cells have a broad oval shape, 40.0–55.0 μm long and 30.0–47.5 μm wide, and a wide U‐shaped periflagellar area on the right thecal plate. The left thecal plate consists of a straighter apical outline in the form of a raised ridge. Five to six delicate apical spines in the center of the periflagellar area are present. The nucleus is located in the posterior region of the cell, and a conspicuous pusule is located in the anterior region of the cell. The cells have golden‐brown chloroplasts with a compound, intrachloroplast pyrenoid that lacks a starch sheath. The thecal plates are smooth with round pores of two different sizes. The larger pores are arranged in a specific pattern of radial rows that are evenly spaced around the plate periphery and of irregular rows (or double rows) that form an incomplete “V” at the apical end of the plates. Large pores are absent in the center of the left and right thecal plates. The intercalary band is striated transversely and also has faint horizontal striations. Trichocysts and two types of mucocysts are present. The molecular phylogenetic position of Prorocentrum tsawwassenense sp. nov. was inferred using SSU rDNA sequences. This new species branched with high support in a Prorocentrum clade containing both benthic and planktonic species.     

ALEXANDRIUM TAMUTUM SP. NOV. (DINOPHYCEAE): A NEW NONTOXIC SPECIES IN THE GENUS ALEXANDRIUM1

A new species of the dinoflagellate genus Alexandrium, A. tamutum sp. nov., is described based on the results of morphological and phylogenetic studies carried out on strains isolated from two sites in the Mediterranean Sea: the Gulf of Trieste (northern Adriatic Sea) and the Gulf of Naples (central Tyrrhenian Sea). Vegetative cells were examined in LM and SEM, and resting cysts were obtained by crossing strains of opposite mating type. Alexandrium tamutum is a small‐sized species, resembling A. minutum in its small size, the rounded‐elliptical shape and the morphology of its cyst. The main diagnostic character of the new species is a relatively wide and large sixth precingular plate (6″), whereas that of A. minutum is much narrower and smaller. Contrary to A. minutum, A. tamutum strains did not produce paralytic shellfish poisoning toxins. Phylogenies inferred from the nuclear small subunit rDNA and the D1/D2 domains of the large subunit nuclear rDNA of five strains of A. tamutum and numerous strains of other Alexandrium species showed that A. tamutum strains clustered in a well‐supported clade, distinct from A. minutum.     

MORPHOLOGY AND MOLECULAR PHYLOGENY OF ANKISTRODINIUM GEN. NOV. (DINOPHYCEAE), A NEW GENUS OF MARINE SAND‐DWELLING DINOFLAGELLATES FORMERLY CLASSIFIED WITHIN AMPHIDINIUM1

The classical athecate dinoflagellate genera (Amphidinium, Gymnodinium, Gyrodinium) have long been recognized to be polyphyletic. Amphidinium sensu lato is the most diverse of all marine benthic dinoflagellate genera; however, following the redefinition of this genus ～100 species remain now of uncertain or unknown generic affiliation. In an effort to improve our taxonomic and phylogenetic understanding of one of these species, namely Amphidinium semilunatum, we re‐investigated organisms from several distant sites around the world using light and scanning electron microscopy and molecular phylogenetic methods. Our results enabled us to describe this species within a new heterotrophic genus, Ankistrodinium. Cells of A. semilunatum were strongly laterally flattened, rounded‐quadrangular to oval in lateral view, and possessed a small asymmetrical epicone. The sulcus was wide and characteristically deeply incised on the hypocone running around the antapex and reaching the dorsal side. The straight acrobase with hook‐shaped end started at the sulcal extension and continued onto the epicone. The molecular phylogenetic results clearly showed that A. semilunatum is a distinct taxon and is only distantly related to species within the genus Amphidinium sensu stricto. The nearest sister group to Ankistrodinium could not be reliably determined.     

AMPHIDINIUM CRYOPHILUM SP. NOV. (DINOPHYCEAE) A NEW FRESHWATER DINOFLAGELLATE. I. SPECIES DESCRIPTION USING LIGHT AND SCANNING ELECTRON MICROSCOPY1

Amphidinium cryophilum sp. nov. was found in the fall of 1979 in a small pond near Madison, Wisconsin. During the ensuing winter, it became the dominant phytoplankter. Cell numbers remained high despite a thick layer of ice and snow. After the ice melted in the spring the organism disappeared from plankton samples. A successful culture of A. cryophilum was established only when isolates were incubated at 5–7° C. It is compared with two morphologically similar species, A. amphidinioides (Geitler) Schiller and Gymnodinium inversum Nygaard. Amphidinium cryophilum is distinguished from the former by its pigmentation (golden-yellow vs. blue-green), the location of the cingulum, and its lack of an eyespot. It differs from the latter in cell shape, the route of the sulcus and position of the nucleus.     

POLARELLA GLACIALIS, GEN. NOV., SP. NOV. (DINOPHYCEAE): SUESSIACEAE ARE STILL ALIVE!

The culture CCMP 1383, obtained from sea-ice brine collected in McMurdo Sound (Ross Sea, Antarctica), is a small gymnodinioid dinoflagellate. This species is very abundant in the upper land-fast sea ice, where it can both grow and overwinter as a spiny encysted stage. The motile vegetative stage and the cyst produced in the culture were studied by scanning electron microscopy (SEM) and transmission electron micrscopy (TEM). The amphiesma of the vegetative cells is constituted by thin vesicles that are organized into nine latitudinal series of plates: three in the epitheca, two in the cingulum, and four in the hypotheca. The same tabulation is reflected in the cyst wall by acicular processes arising from the center of paraplates, with the exception of the paracingulum, in which acicular processess are absent. On the basis of the peculiar plate pattern of this dinoflagellate, we establish the new genus Polarella and the new species Polarella glacialis (family Suessiaceae, order Suessiales). This species has a remarkable similarity with fossil Suessiaceae cysts dating back to the Triassic and Jurassic and represents, up to now, the only extant member of the subfamily Suessiaceae. Phylogenetic analysis based on the small-subunit ribosomal RNA gene confirmed the placement of this species in the order Suessiales and its close relationship with the genus Symbiodinium Freudenthal.     

ALEXANDRIUM SATOANUM SP. NOV. (DINOPHYCEAE) FROM MATOYA BAY,CENTRAL JAPAN1

The gonyaulacoid dinofiagellate Alexandrium satoanum Yuki et Fukuyo sp. nov. is described from Matoya Bay, Pacific coast of central Japan. The species is distinctive in its conical epitheca with almost straight sides and dorsal concavity of the hypotheca. The plate formula is Po, pc, 4′, 6″, 6c, 10s, 5″″, and 2″″, including two accessory plates inside the sulcus. The apical pore plate is triangular and possesses an anterior attachment pore at the right margin. The first apical plate does not make contact with the apical pore plate and lacks a ventral pore. A posterior attachment pore lies at the center of the posterior sulcal plate. In Matoya Bay, vegetative cells occur as solitary cells or sometimes in pairs during late spring and early summer in low concentrations. In connection with this study, the following new combination is proposed: Alexandrium pseudogonyaulax (Biecheler) Horiguchi ex Yuki et Fukuyo comb. nov.     

PHYLOGENETICS OF RHINODINIUM BROOMEENSE GEN. ET SP. NOV., A PERIDINIOID,SAND‐DWELLING DINOFLAGELLATE (DINOPHYCEAE)1

The phylogeny of Rhinodinium broomeense, a new genus and species of heterotrophic peridinioid dinoflagellates, has been studied based on morphological and molecular genetic data. The genus was found in tidal marine sand habitats in Broome, north‐western Australia, and from three marine sand habitats in Japan. The thecal plate formula is Po 3′ 1^a 5″ 4c ?s 5″′ 1″″. A large apical hook points toward the dorsal side. Its plate pattern is similar to species of the genus Roscoffia; however, it differs from that genus in its much larger epitheca, narrow cingulum, which could be interpreted as incomplete, the narrow sulcus without sulcal lists on both sides, and the strong oblique lateral compression. Phylogenetic analyses using partial LSU rDNA sequences, as well as plate pattern information, support the placement of this genus in the Peridiniales; however, it is sufficiently different from other genera that the family affinity remains unclear.     

MORPHOLOGY AND LIFE CYCLE EVENTS IN PYROPHACUS STEINII (SCHILLER) WALL ET DALE (DINOPHYCEAE)

Cells of Pyrophacus steinii (Schiller) Wall et Dale are round and lens shaped and have an anteroposteriorly compressed theca. The epitheca has a truncated, conical horn and a hexagonally shaped apical pore plate with two arched slits positioned off center. The cingulum is equatorial, narrow, and deep. The hypotheca is flat. The sulcus is narrow, slightly curved, and recessed and does not reach the cell's antapex. The plate formula in these specimens of P. steinii is Po, 8', Oa, 13", 13C, 12"', 3p, 3"", and 8S with a difference in the number of precingular (13") and postcingular (12"') plates. No additional posterior intercalary plates were present (Oap). Pregametic stages of P. steinii were observed during cell division via binary fission, with formation of two cells and multiple division with formation of four and eight cells. These newly formed cells were pale in color and were enclosed in double-layered hyaline membrane. Gametes with gymnodinoid morphology were observed within the parental cells. Planozygotes are large and round and enclosed in double-layered hyaline membrane. Mature cell forms are brown with a microgranular cytoplasm, storage bodies, and a red accumulation body. The hypnozygote exhibits triple-layered hyaline membrane, irregularly shaped and comparable with bulbous processes of Tuberculodinium vancampoae Rossigol resting cysts. Division within a hypnocyst of P. steinii involves shedding the parental theca and the development and emergence of two daughter cells with the size and morphology of pregametic cells.