Issue Information

A tidal pool dinoflagellate, Chiharadinium hexapraecingulum (T. Horiguchi & Chihara) Dawut & T. Horiguchi gen. & comb. nov. Upper row: Light micrographs of lateral view, ventral view, epifluorescence micrograph of motile cells and scanning electron micrograph of apical view of a cell. Lower row: Scanning electron micrographs of ventral view, left lateral view and antapical view of motile cells. Establishment of a new genus Chiharadinium is reported by Dawut et al. in this issue.

     

Ultrastructure of Stylodinium Iittorale (Dinophyceae) with special reference to the stalk and apical stalk complex

The ultrastructure of Stylodinium littorale Horiguchi et Chihara, a marine, sand-dwelling coccoid dinoflagel-late, was investigated with special emphasis on its stalk and the apical stalk complex. The dinoflagellate alternates between non-motile and motile cells in its life cycle. The non-motile cell possesses a long and distinct stalk. The stalk, consisting of a main cylindrical part and a holdfast, is firmly attached to a thecal plate (the apical pore plate). A part of its proximal portion is hollow and V-shaped in section. The V-shaped hollow space is underlain by a projection from the apical pore plate. An apical stalk complex is present in the motile cells and consists of a large apical pore plate and mucilaginous material. The apical pore plate is depressed into the cell, but has a narrow central tubular projection. The mucilaginous stalk-building material is stored between this plate and the outer plate membrane. The tubular projection of the apical pore plate corresponds to the apical pore of other dinoflagellates and its lumen is filled with electron-dense material. The structure of the apical stalk complex is compared with the homologous structure in Bysmatrum arenicola, the only other example of an apical stalk complex that has been investigated. A general ultrastructural survey revealed that S. littorale possesses a typical dinoflagellate cellular structure.     

Plagiodinium ballux sp. nov. (Dinophyceae), a deep (36 m) sand dwelling dinoflagellate from subtropical Japan

A new species of marine sand‐dwelling dinoflagellate, Plagiodinium ballux N. Yamada, Dawut, R. Terada & T. Horiguchi is described from a deep (36 m) seafloor off Takeshima Island, Kagoshima Prefecture, Japan in the subtropical region of the northwest Pacific. The species is thecate and superficially resembles species of Prorocentrum, but possesses an extremely small epitheca. The cell varies from ovoid to a rounded square, and is small (15.0–22.5 μm in length) and laterally compressed. The thecal plates are smooth and the thecal plate arrangement (Po, 1′, 0a, 5″, 5C, 2S, 5?, 0p, 1″″) is similar to that of Plagiodinium belizeanum, the type species of the genus. Molecular phylogenetic analyses based on SSU rDNA and partial LSU rDNA reveal that the dinoflagellate is closely related to P. belizeanum, but it can be clearly distinguished by its size and cell shape. This suite of morphological and molecular differences leads to the conclusion that this deep benthic dinoflagellate represents a new species of the genus Plagiodinium.     

Amphidiniella sedentaria gen. et sp. nov. (Dinophyceae), a new sand-dwelling dinoflagellate from Japan

A new sand-dwelling dinoflagellate is described from Sesoko Beach, Okinawa Island, subtropical Japan and its micromorphology is studied by means of light and electron microscopy. The cell consists of a small epitheca and a large hypothecs superficially resembling members of the unarmored genus Amphidinium. The cell is dorso-ventrally flattened and possesses a single chloroplast with a large conspicuous pyrenoid. Transmission electron microscopy revealed that the dinoflagellate possesses typical dinoflagellate cellular organization. Scanning electron microscopy demonstrated that the organism is thecate and the thecal plate arrangement is Po, 4′, 1a, 7″, 5c, 4s, 6″′, 2″″. Most of the characteristics suggest gonyaulacalean affinity of the new species. These are the presence of ventral pore, lack of canal plate, direct contact between the sulcal anterior plate and the flagellar pore, possession of six postcingular plates and asymmetrical arrangement of the antapical plates. Affinity to existing families of the order Gonyaulacales has not been determined. Based on the unique cell shape, thecal plate arrangement and the presence of ventral pore, a new genus, Amphidiniella, is established for this organism and the species is named A. sedentaria Horiguchi gen. et sp. nov.     

HALOSTYLODINIUM ARENARIUM, GEN ET SP. NOV. (DINOPHYCEAE), A COCCOID SAND-DWELLING DINOFLAGELLATE FROM SUBTROPICAL JAPAN

A new genus and species of marine coccoid dinoflagellate from subtropical Japan, Halostylodinium arenarium Horiguchi et Yoshizawa-Ebata, gen. et sp. nov., is described. The dominant stage of the dinoflagellate is a nonmotile ovoidal to spheroidal cell with a distinct stalk. The stalk consists of an upper thick tubule, a lower thin tubule, and a discoidal holdfast. The dinoflagellate possesses a yellowish-brown chloroplast with multiple lobes radiating from a central pyrenoid. It reproduces by the formation of two motile cells, which swim for a short period and then transform directly into the stalked nonmotile cell. The stalk is produced during transformation from the apical stalk complex present in the apex of the motile cell. The apical stalk complex consists of a double-folded apical pore plate and doughnut-shaped holdfast-building material. The ultrastructure of the apical stalk complex is compared with those of Bysmatrum arenicola and Stylodinium littorale. Halostylodinium arenarium possesses delicate thecal plates, and the thecal plate formula is Po, 5', 2a, 7", 7c, 6s, 5"', 1p, 2"". A phylogenetic study based on the 18S ribosomal RNA gene did not show any clear affinities between this organism and any species included in the analysis.     

Scrippsiella hexapraecingula sp. nov. (Dinophyceae), a tida pool dinoflagellate from the Northwest Pacific

Specimens of dinoflagellate collected in tide pools along the Pacific coast of central and southern Japan are described as a new species,Scrippsiella hexapraecingula Horiguchi et Chihara, of the Peridiniaceae (Class Dinophyceae). The plate formula is pp, x, 4′, 3a, 6″, 6c, 5‴, 2″" and, 5s, the same as that of other species ofScrippsiella, except in lacking one precingular plate. The genus must be emended, therefore, as having either six or seven precingular plates. This dinoflagellate migrates diurnally. In the morning motile cells are released from non-motile cells attached to the substrate and in the evening the motile cells swim down to settle on the bottom of the tide pool. Attached non-motile cells form either motile mono- or bispores. Sexual reproduction was not observed.     

ULTRASTRUCTURE OF A NEW SAND-DWELLING DINOFLAGELLATE,SCRIPPSIELLA ARENICOLA SP. NOV1

A new dinoflagellate, Scrippsiella arenicola Horiguchi et Pienaar sp. nov., is described from tidal pools with sandy substrates along the east coast of South Africa. S. arenicola exhibits a vertical migratory rhythm which is in synchrony with the tidal cycle. It is a medium-sized armoured dinoflagellate with many rod-shaped chloroplasts. Thecal plate arrangement is pp, x, 4′, 3a, 7′, 6c, 5′, 2″ and 4s. The 2a and 3a plates are separated from each other. S. arenicola has several unique ultrastructural features. Electron-dense fibres are found on the protruded part of the thecal plates, such as on the ornamental projections or extremities of the lists. In addition to the 9 + 2 axoneme, additional fibres are found in the free moving part of the longitudinal flagellum. The portion of the transverse flagellum covered by the left sulcal list possesses a dense array of mastigonemes which connect the flagellum and the cell. The flagellar pore platelets differ from ordinary thecal plates in their thickness and fibrous nature. The ultrastructure of the apical stalk and its associated structures is described. The vertical migration and mode of cell division is also described.     

Life cycle,behavior and morphology of a new tide pool dinoflagellate,Gymnodinium pyrenoidosum sp. nov. (Gymnodiniales,Pyrrhophyta)

A new tide pool dinoflagellate,Gymnodinium pyrenoidosum Horiguchi et Chihara sp. nov. is described from central Japan. It was found to form dense blooms with a characteristic greenish color from April to November. The species exhibits a characteristic diurnal vertical migration and an alternation of a motile with a nonmotile phase, which are dependent on light intensity and tidal movement. Cells of the motile phase are unarmored and relatively small. They have a single, reticulate chloroplast, orange stigma situated near the sulcus and conspicuous pyrenoid in epicone. The alga reproduces itself by means of zoospores which are produced by the bipartition of protoplasm within the parent cell wall during the nonmotile stage which occurs at night. The occurrence of another type of motile cell, termed a macroswarmer, which differs from normal zoospore in size and shape has also been demonstrated.     

Roscoffia minor sp. nov. (Peridiniales, Dinophyceae): A new, sand-dwelling, armored dinoflagellate from Hokkaido, Japan

A new, sand-dwelling, armored dinoflagellate, Roscoffia minor sp. nov., is described from Ishikari beach, Hokkaido, Japan. The dinoflagellate has been collected from sand samples taken both near the water's edge and further upshore (25 m from the water's edge at a depth of 1 m), indicating that it is a true sand-dwelling species. Roscoffia minor is heterotrophic and lacks both a chloroplast and an eye-spot. The cell consists of a flattened cap-shaped epitheca and a large hemispheroidal hypotheca, and it is quite different from cells of the typical armored dinoflagellates. The thecal plate formula is: Po, 3′, la, 5″, 3c, 3s, 5″, 1″″. Its distinct cell shape and the thecal plate arrangement indicate affinity to the monotypic genus Roscoffia. Roscoffia minor is distinguished from Roscoffia capitata, the type species, by its smaller size and the possession of a finger-like apical projection. The thecal arrangement of the epitheca is similar to those of the members of the family Podolampaceae, while the hypothecal arrangement is the same as that of members of the subfamily Diplopsalioideae (family Congruentidiaceae). The organism seems to be positioned somewhere intermediate between these two families, but the family to which this dinoflagellate should be affiliated could not be determined.     

Development of the cell covering in the dinoflagellate Scrippsiella hexapraecingula (Peridiniales, Dinophyceae)

The organization and development of cell coverings in two alternate phases of the life cycle in a marine dinoflagellate, Scrippsiella hexapraecingula Horiguchi et Chihara, were investigated by thin sectioning and freeze‐fracture electron microscopy. In one of these phases, the motile phase, cells have an outermost plasma membrane that is lined with flattened amphiesmal vesicles. Groups of microtubules lie beneath these vesicles. In mature motile cells, thecal plates are completely enclosed in individual amphiesmal vesicles. After settling, the cells enter the second, non‐motile phase. Here, ecdysis occurs, resulting in several steps including formation of the first pellicle layer (PI), fusion of the inner amphiesmal vesicle membranes to form the new plasma membrane, deposition of the second pellicle layer (PM) under PI, and the appearance and fusion of juvenile amphiesmal vesicles to form new territories, which eventually give rise to new thecal plates in the next motile phase. Thus, the pattern in which thecal plates are arranged in motile cells is determined at the time when the amphiesmal vesicles develop into non‐motile cells.     

Pyramidodinium atrofuscum gen. et sp. nov. (Dinophyceae), a new marine sand-dwelling coccoid dinoflagellate from tropical waters

A new marine sand‐dwelling coccoid dinoflagellate Pyramidodinium atrofuscum Horiguchi et Sukigara gen. et sp. nov. is described from Jellyfish Lake, Republic of Palau. The dinoflagellate alternates a non‐motile vegetative stage with a motile gymnodinioid stage within its life cycle. The non‐motile stage is dominant in the life cycle and the dinoflagellate reproduces itself by means of the production of two motile cells. The released motile cell swims only for a short period and is directly transformed into the non‐motile cell. The non‐motile cell is sessile, pyramidal in shape, with a single longitudinal ridge and a double transverse ridge. The surface of the cell wall is covered with many processes. The motile cell has a Gymnodinium‐like morphology, but no apical groove is present. An ultrastructural study revealed that the dinoflagellate possesses typical dinoflagellate organelles. Based on the unique morphology of the vegetative non‐motile stage, we propose a new genus Pyramidodinium for this dinoflagellate, with the type species Pyramidodinium atrofuscum Horiguchi et Sukigara, gen. et sp. nov.     

Bispinodinium angelaceum gen. et sp. nov. (Dinophyceae), a new sand‐dwelling dinoflagellate from the seafloor off Mageshima Island,Japan1

A new athecate dinoflagellate, Bispinodinium angelaceum N. Yamada et Horiguchi gen. et sp. nov., is described from a sand sample collected on the seafloor at a depth of 36 m off Mageshima Island, subtropical Japan. The dinoflagellate is dorsiventrally compressed and axi‐symmetric along the sulcus. The morphology resembles that of the genus Amphidinium sensu lato by having a small epicone that is less than one third of the total cell length. However, it has a new type of apical groove, the path of which traces the outline of a magnifying glass. The circular component of this path forms a complete circle in the center of the epicone and the straight “handle” runs from the sulcus to the circular component. Inside the cell, a pair of elongated fibrous structure termed here the “spinoid apparatus” extends from just beneath the circular apical groove to a point near the nucleus. Each of two paired structures consists of at least 10 hyaline fibers and this is a novel structure found in dinoflagellates. Phylogenetic analyses based on the SSU and LSU RNA genes did not show any high bootstrap affinities with currently known athecate dinoflagellates. On the basis of its novel morphological features and molecular signal, we conclude that this dinoflagellate should be described as a new species belonging to a new genus.     

Testudodinium gen. nov. (Dinophyceae), a new genus of sand‐dwelling dinoflagellates formerly classified in the genus Amphidinium

A new genus of sand‐dwelling photosynthetic dinoflagellate, Testudodinium Horiguchi, Tamura, Katsumata et A. Yamaguchi is proposed based on Testudodinium testudo (Herdman) Horiguchi, Tamura, Katsumata, et A. Yamaguchi comb. nov. (Basionym: Amphidinium testudo Herdman) and a new species in this new genus, Testudodinium maedaense Katsumata et Horiguchi sp. nov. is described. Amphidinium corrugatum is also transferred to this genus, making a new combination T. corrugatum (Larsen et Patterson) Horiguchi, Tamura et A. Yamaguchi. These three species are similar to the members of the genus Amphidinium in having an extremely small episome and a dorsoventrally flattened cell body. They are, however, distinguished from the genus Amphidinium seusu stricto by the possession of a distinct longitudinal furrow in the middle of ventral side of the episome. Phylogenetic trees based on small subunit (SSU) rDNA revealed that all three of these Testudodinium species formed a robust clade and, although statistical support is not high, the tree suggests Testudodinium clade is not closely related to Amphidinium seusu stricto clade. The morphological differences together with molecular data support the establishment of a new genus for A. testudo and its related species.     

Heterocapsa circularisquama sp. nov. (Peridiniales, Dinophyceae): A new marine dinoflagellate causing mass mortality of bivalves in Japan

Heterocapsa circularisquama Horiguchi sp. nov. is described from Ago Bay, central Japan. The dinoflagellate produced large-scale red tides in the bays of central and western Japan and caused mass mortality of bivalves, notably the pearl oysters. The cell is small and is composed of a conical epitheca and a hemi-spheroidal hypothecs. The chloroplast is single and is connected to the single pyrenoid. The nucleus is elongated and is located in the left side of the cell. Thecal plate arrangement has been determined as: Po, cp, 5′, 3a, 7″, 6c, 5s, 5″′, 2″″. Heterocapsa circularisquama is morphologically very similar to Heterocapsa illdefina and it is almost impossible to distinguish these two species at light microscopical level. The characteristics which can be used to distinguish these two species are the morphology of body scales and the ultrastructure of the pyrenoid matrix. The body scales of H. circularisquama possess six radiating ridges on the circular basal plate; no such ridges can be observed on the roughly triangular basal plate of the scales of H. illdefina. Furthermore, the scales of the latter species possess substantially shorter spines compared to those of H. circularisquama. The pyrenoid matrix of H. circularisquama is hardly perforated by cytoplasmic tubules, while in H. tlldefina the pyrenoid matrix is always penetrated by many cytoplasmic tubules. Based on the arrangement of thecal plates, morphology of body scales, and ultra-structure of the pyrenoid, I am placing H. circularisquama sp nov. into the genus Heterocapsa.     

Heterocapsa psammophila sp. nov. (Peridiniales, Dinophyceae), a new sand-dwelling marine dinoflagellate

A new armored dinoflagellate species, Heterocapsa psammophila Tamura, Iwataki et Horiguchi sp. nov. is described from Kenmin‐no‐hama beach, Hiroshima, Japan using light and electron microscopy. This dinoflagellate possesses the typical thecal plate arrangement of the genus Heterocapsa, Po, cp, 5′, 3a, 7′′, 6c, 5s, 5′′′, 2′′′′; and the 3‐D body scales of Heterocapsa on the plasma membrane. The cell shape is ovoidal. The spherical nucleus and the pyrenoid are situated in the hypotheca and the epitheca, respectively. The ultrastructure of H. psammophila is typical of dinoflagellates and the pyrenoid is invaginated by cytoplasmic tubules. H. psammophila is distinguished from all other hitherto‐described Heterocapsa species by the cell shape, the relative position of the nucleus and pyrenoid and the structure of the body scale. The habitat and behavior of this new species in culture suggest that the organism is truly a sand‐dwelling species.     

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.     

Phototactic responses of four marine dinoflagellates with different types of eyespot and chloroplast

Great structural variety is seen in the eyespot of dinoflagellates, a structure involved in phototaxis. Although there are several works on the phototactic responses in some species of dinoflagellates, none of the dinoflagellates used in these studies possessed an eyespot and, therefore, we have no knowledge of the relationship between eyespot type and phototactic response. In this study, we determined wavelength dependency curves for phototaxis in four marine dinoflagellates that possess a different type of either eyespot or chloroplast. These include: (i) a dinoflagellate possessing a peridinin-containing ohioroplast with an eyespot (Scrippsiella hexapraecingula Horiguchi et Chihara); (ii) a dinoflagellate containing a diatom endosymbiont and with the type B eyespot sensu Dodge (1984; (Peridinium foli-aceum (Stein) Biecheler); (iii) a dinoflagellate with peri-dinin-containing chloroplasts, but lacking an eyespot (Atexandrium hiranoi Kita et Fukuyo); and (iv) a dinoflagellate with fucoxanthin, 19′-hexanoyloxyfucoxanthin and 19′-butanoyloxyfucoxanthin, but lacking an eyespot (Gymnodinium mikimotoi Miyabe et Kominami ex Oda), Regardless of the eyespot or the chloroplast type, all four dinoflagellates showed similar wavelength dependency curves for phototaxis, with sensitivity between 380 and 520 nm, the highest peak at approximately 440 or 460 nm and smaller peaks or shoulders at 400–420 nm and 480–500 nm. Substantial peaks have also been noted in the ultraviolet range (260–280 nm). The ultrastructural study of the eye-spot of Scrippsiella hexapraecingula revealed that the eyespot consists of two layers of lipid globules and probably acts as a quarter-wave stack antenna.     

Ankistrodinium armigerum sp. nov. (Dinophyceae), a new species of heterotrophic marine sand‐dwelling dinoflagellate from Japan and Australia

A new heterotrophic sand‐dwelling dinoflagellate, Ankistrodinium armigerum K. Watanabe, Miyoshi, Kubo, Murray et Horiguchi sp. nov., is described from Ishikari Beach, Hokkaido, Japan and Port Botany, NSW, Australia. The dinoflagellate is laterally compressed, possessing a short triangular epicone and a large sac‐like hypocone. It possesses a right‐handed cingulum and a deeply‐incised sulcus. The sulcus descends towards the posterior of the cell where it becomes much deeper and wider, resulting in a bilobed ventral side to the hypocone, with a greater excavation of the left lobe than the right. In addition, the right lobe of the hypocone is shorter than the left lobe, which allows a partial view of the left sulcal wall when the cell is viewed from its right side. The sulcus ascends in the epicone to form an apical groove. The apical groove is linear but terminates in an ellipsoid fashion and its extremity approaches, but does not form a closed loop with the apical end of the linear portion. The dinoflagellate possesses two distinct size classes of trichocysts. The large trichocysts are located in the posterior part of the cell, while small trichocysts are distributed throughout the cell. The dinoflagellate shares morphological characteristics with the heterotrophic sand‐dwelling dinoflagellate, Ankistrodinium semilunatum, the type species of the genus. These include a laterally compressed cell, a right‐handed cingulum, a deeply‐incised sulcus and the same basic structure to the apical groove. Molecular phylogenetic analyses based on small and large subunits of rDNA showed that in both trees, A. semilunatum and A. armigerum formed a robust clade, suggesting that these two species are closely related. Because no organism with the characteristics of this species exists and because this species is closely related to A. semilunatum, we concluded that this species should be described as a second species of the genus Ankistrodinium.     

TAKAYAMA GEN. NOV. (GYMNODINIALES,DINOPHYCEAE), A NEW GENUS OF UNARMORED DINOFLAGELLATES WITH SIGMOID APICAL GROOVES,INCLUDING THE DESCRIPTION OF TWO NEW SPECIES1

A new potentially ichthyotoxic dinoflagellate genus, Takayama de Salas, Bolch, Botes et Hallegraeff gen. nov., is described with two new species isolated from Tasmanian (Australia) and South African coastal waters: T. tasmanica de Salas, Bolch et Hallegraeff, sp. nov. and T. helix, de Salas, Bolch, Botes et Hallegraeff, sp. nov. The genus and two species are characterized by LM and EM of field samples and laboratory cultures as well as large subunit rDNA sequences and HPLC pigment analyses of several cultured strains. The new Takayama species have sigmoid apical grooves and contain fucoxanthin and its derivatives as the main accessory pigments. Takayama tasmanica is similar to the previously described species Gymnodinium pulchellum Larsen, Gyrodinium acrotrochum Larsen, and G. cladochroma Larsen in its external morphology but differs from these in having two ventral pores, a large horseshoe‐shaped nucleus, and a central pyrenoid with radiating chloroplasts that pass through the nucleus. It contains gyroxanthin‐diester and a gyroxanthin‐like accessory pigment, both of which are missing in T. helix. Takayama helix has an apical groove that is nearly straight while still being clearly inflected. A ventral pore or slit is present. It has numerous peripheral, strap shaped, and spiraling chloroplasts with individual pyrenoids and a solid ellipsoidal nucleus. The genus Takayama has close affinities to the genera Karenia and Karlodinium.     

THE RECLASSIFICATION OF PFIESTERIA SHUMWAYAE (DINOPHYCEAE): PSEUDOPFIESTERIA,GEN. NOV.1

Pfiesteria shumwayae Glasgow et Burkholder is assigned to a new genus Pseudopfiesteria gen. nov. Plate tabulation differences between Pfiesteria and Pseudopfiesteria gen. nov. as well as a maximum likelihood phylogenetic analysis based on rDNA sequence data warrant creation of this new genus. The Kofoidian thecal plate formula for the new genus is Po, cp, X, 4′, 1a, 6′′, 6c, PC, 5+s, 5′′′, 0p, 2′′′′. In addition to having six precingular plates, P. shumwayae comb. nov. also has a distinctive diamond or rectangular‐shaped anterior intercalary plate. Both Pfiesteria and Pseudopfiesteria gen. nov. are reassigned to the order Peridiniales based on an apical pore complex (APC) with a canal (X) plate that contacts a symmetrical 1′, four to five sulcal plates, and the conservative hypothecal tabulation of 5′′′, 0p, and 2′′′′. These morphological characters and the life histories of Pfiesteria and Pseudopfiesteria are consistent with placement of both genera in the Peridiniales. Based on the plate tabulations for P. shumwayae, P. piscicida, and the closely related “cryptoperidiniopsoid” and “lucy” groups, the family Pfiesteriaceae is amended to include species with the following tabulation: 4‐5′, 0‐2a, 5‐6′′, 6c, PC, 5+s, 5′′′, 0p, and 2′′′′ as well as an APC containing a pore plate (Po), a closing plate (cp), and an X plate; the tabulation is expanded to increase the number of sulcal plates and to include a new plate, the peduncle cover (PC) plate. Members of the family have typical dinoflagellate life cycles characterized by a biflagellated free‐living motile stage, a varying number of cyst stages, and the absence of multiple amoeboid stages.