GYMNODINIUM COROLLARIUM SP. NOV. (DINOPHYCEAE)—A NEW COLD‐WATER DINOFLAGELLATE RESPONSIBLE FOR CYST SEDIMENTATION EVENTS IN THE BALTIC SEA1

A naked dinoflagellate with a unique arrangement of chloroplasts in the center of the cell was isolated from the northern Baltic proper during a spring dinoflagellate bloom (March 2005). Morphological, ultrastructural, and molecular analyses revealed this dinoflagellate to be undescribed and belonging to the genus Gymnodinium F. Stein. Gymnodinium corollarium A. M. Sundström, Kremp et Daugbjerg sp. nov. possesses features typical of Gymnodinium sensu stricto, such as nuclear chambers and an apical groove running in a counterclockwise direction around the apex. Phylogenetic analyses based on partial nuclear‐encoded LSU rDNA sequences place the species in close proximity to G. aureolum, but significant genetic distance, together with distinct morphological features, such as the position of chloroplasts, clearly justifies separation from this species. Temperature and salinity experiments revealed a preference of G. corollarium for low salinities and temperatures, confirming it to be a cold‐water species well adapted to the brackish water conditions in the Baltic Sea. At nitrogen‐deplete conditions, G. corollarium cultures produced small, slightly oval cysts resembling a previously unidentified cyst type commonly found in sediment trap samples collected from the northern and central open Baltic Sea. Based on LSU rDNA comparison, these cysts were assigned to G. corollarium. The cysts have been observed in many parts of the Baltic Sea, indicating the ecologic versatility of the species and its importance for the Baltic ecosystem.     

COOLIA CANARIENSIS SP. NOV. (DINOPHYCEAE), A NEW NONTOXIC EPIPHYTIC BENTHIC DINOFLAGELLATE FROM THE CANARY ISLANDS1

A new photosynthetic dinoflagellate species, Coolia canariensis S. Fraga sp. nov., is described based on samples taken from tidal ponds on the rocky shore of the Canary Islands, northeast Atlantic Ocean. Its morphology was studied by LM and SEM. It is almost spherical and has a thick smooth theca with many scattered pores. Plate 1′ is the biggest of the epithecal plates, and 7″ is twice as wide as it is long. Phylogeny inferred from the D1/D2 regions of the LSU nuclear rDNA of three strains of C. canariensis and several strains of other Coolia species, C. monotis, C. sp., showed that C. canariensis strains clustered in a well‐supported clade distinct from the other species. No toxins were detected using mouse bioassay, liquid chromatography with Fluorescence detection (LC‐FLD) or liquid chromatography‐mass spectrometry (LC‐MS). Its pigment composition is of the peridinin type of dinoflagellates. Together with this new species, many other strains of C. monotis from the Atlantic Ocean and Mediterranean Sea have been analyzed for toxin presence, and no evidence of toxin production related to yessotoxins (YTXs) was found, as was previously suggested for C. monotis from Australia.     

DESCRIPTION OF A NEW GENUS OF PFIESTERIA‐LIKE DINOFLAGELLATE,LUCIELLA GEN. NOV. (DINOPHYCEAE), INCLUDING TWO NEW SPECIES: LUCIELLA MASANENSIS SP. NOV. AND LUCIELLA ATLANTIS SP. NOV.1

A new genus of Pfiesteria‐like heterotrophic dinoflagellate, Luciella gen. nov., and two new species, Luciella masanensis sp. nov. and Luciella atlantis sp. nov., are described. These species commonly occur with other small (<20 μm) heterotrophic and mixotrophic dinoflagellates in estuaries from Florida to Maryland and the southern coast of Korea, suggesting a possible global distribution. An SEM analysis indicates that members of the genus Luciella have the enhanced Kofoidian plate formula of Po, cp, X, 4′, 2a, 6″, 6c, PC, 5+s, 5?, 0p, and 2″″. The two four‐sided anterior intercalary plates are diamond shaped. The genus Luciella differs from the other genera in the Pfiesteriaceae by a least one plate in the plate tabulation and in the configuration of the two anterior intercalary plates. An SSU rDNA phylogenetic analysis confirmed the genus as monophyletic and distinct from the other genera in the Pfiesteriaceae. The morphology of Luciella masanensis closely resembles Pfiesteria piscicida Steid. et J. M. Burkh. and other Pfiesteria‐like dinoflagellates in size and shape, making it easily misidentified using LM. Luciella atlantis, in contrast, has a more distinctive morphology. It can be distinguished from L. masanensis and other Pfiesteria‐like organisms by a larger cell size, a more conical‐shaped epitheca and hypotheca, larger rhombic‐shaped intercalary plates, and an asymmetrical hypotheca. The genus Luciella is assigned to the order Peridiniales and the family Pfiesteriaceae based on plate tabulation, plate pattern, general morphology, and phylogenetic analysis.     

SEASONAL VARIABILITY OF THE ORGANIC‐WALLED DINOFLAGELLATE CYST PRODUCTION IN THE COASTAL UPWELLING REGION OFF CAPE BLANC (MAURITANIA): A FIVE‐YEAR SURVEY1

A 5‐year sediment trap survey in the upwelling area off Cape Blanc (NW Africa) provides information on the seasonal and annual resting cyst production of dinoflagellates, their sinking characteristics and preservation potential. Strong annual variation in cyst production characterizes the region. Cyst production of generally all investigated species, including Alexandrium pseudogonyaulax (Biecheler) T. Horig. ex T. Kita et Fukuyo (cyst genus Impagidinium) and Gonyaulax spinifera (Clap. et J. Lachm.) Diesing (cyst genus Nematosphaeropsis) was enhanced with increasing upper water nutrient and trace‐element concentrations. Cyst production of Lingulodinium polyedrum (F. Stein) J. D. Dodge was the highest at the transition between upwelling and upwelling‐relaxation. Cyst production of Protoperidinium americanum (Gran et Braarud) Balech, Protoperidinium monospinum (Paulsen) K. A. F. Zonn. et B. Dale, and Protoperidinium stellatum (D. Wall) Balech, and heterotrophic dinoflagellates forming Brigantedinium spp. and Echinidinium aculeatum Zonn., increased most pronouncedly during upwelling episodes. Production of Protoperidinium conicum (Gran) Balech and Protoperidinium pentagonum (Gran) Balech cysts and total diatom valves were related, providing evidence of a predator–prey relationship. The export cyst‐flux of E. aculeatum, P. americanum, P. monospinum, and P. stellatum was strongly linked to the flux of total diatom valves and CaCO₃, whereas the export production of Echinidinium granulatum Zonn. and Protoperidinium subinerme (Paulsen) A. R. Loebl. correlated with total organic carbon, suggesting potential consumption of diatoms, prymnesiophytes, and organic matter, respectively. Sinking velocities were at least 274 m · d^?1, which is in range of the diatom‐ and coccolith‐based phytoplankton aggregates and “slower” fecal pellets. Species‐selective degradation did not occur in the water column, but on the ocean floor.     

GALEIDINIIUM RUGATUM GEN. ET SP. NOV. (DINOPHYCEAE), A NEW COCCOID DINOFLAGELLATE WITH A DIATOM ENDOSYMBIONT1

A new sand‐dwelling dinoflagellate from Palau, Galeidinium rugatum Tamura et Horiguchi gen. et sp. nov., is described. The life cycle of this new alga consists of a dominant nonmotile phase and a brief motile phase. The motile cell transforms itself directly into the nonmotile cell after swimming for a short period, and cell division takes place in the nonmotile phase. The nonmotile cell possesses a dome‐like cell covering, which is wrinkled and equipped with a transverse groove on the surface. The cell has 10–20 chloroplasts and a distinct eyespot. The motile cell is Gymnodinium‐like in shape. The dinoflagellate possesses an endosymbiotic alga to which the chloroplasts belong and which is separated from the host (dinoflagellate) cytoplasm by a unit membrane. The endosymbiont cytoplasm also possesses its own eukaryotic nucleus and mitochondria. The eyespot is surrounded by triple membranes and is located in the host cytoplasm. Photosynthetic pigment analysis, using HPLC, revealed that G. rugatum possesses fucoxanthin as the principal accessory pigment instead of peridinin. The rbcL tree showed that G. rugatum is monophyletic with Durinskia baltica (Levander) Carty et Cox and Kryptoperidinium foliaceum (Stein) Lindemann and that this clade is closely related to the pennate diatom, Cylindrotheca sp. The endosymbiont of G. rugatum is therefore shown to be a diatom. Phylogenetic analysis based on small subunit rDNA sequences demonstrated that G. rugatum, D. baltica, and K. foliaceum, all of which are known to harbor an endosymbiont of diatom origin, are closely related.     

AMPHIDINIUM CRYOPHILUM SP. NOV. (DINOPHYCEAE) A NEW FRESHWATER DINOFLAGELLATE. II. ULTRASTRUCTURE1

The dinoflagellate Amphidinium cryophilum sp. nov. is one of the few gymnodinians to be studied at the ultrastructural level. It resembles other dinoflagellates in the structure of the nucleus, trichocysts, storage materials, flagella, mitochondria, and microbodies. Other features of A. cryophilum less commonly observed in related organisms include a network of small interconnected vesicles, a system of large, peripheral vacuoles, chloroplasts bound by two rather than three membranes, an accumulation body, thylakoid-associated plastoglobuli, a vesiculated nuclear envelope, a complex tubular pusule, striated flagellar collars, collared pits, and a peduncle. The occurrence of a peduncle, a structure implicated in phagotrophy, in this autotrophic organism is noteworthy. The ultrastructure of the peduncle of A. cryophilum differs significantly from that reported in another dinoflagellate.     

DESCRIPTION OF THECADINIUM MUCOSUM SP. NOV. (DINOPHYCEAE), A NEW SAND‐DWELLING MARINE DINOFLAGELLATE,AND AN EMENDED DESCRIPTION OF THECADINIUM INCLINATUM BALECH1

A new thecate, phototrophic, marine, sand‐dwelling dinoflagellate, Thecadinium mucosum Hoppenrath et Taylor sp. nov., is described from a culture isolated from Boundary Bay, British Columbia, Canada. It was illustrated with LM as well as SEM and TEM, and its position in the phylogenetic tree of dinoflagellates was investigated using molecular methods. Cells are asymmetrical, oval, laterally flattened, and strongly pigmented, with the plate formula P 3′ 1a 6′′ 7/8c 5 s 6′′′ 2′′′′. Thecal plates are smooth with scattered pores, and there is a distinctive anterior intercalary plate that could be involved in mucus secretion. Thecadinium inclinatum Balech (=Sabulodinium inclinatum (Balech) Saunders et Dodge), a thecate, marine, sand‐dwelling species that has been previously confused with what we now call T. mucosum, was also examined and illustrated through LM and SEM. New information on T. inclinatum is provided, including its plate formula P 3′ 6′′ 7c ?s 5′′′ 1p 1′′′′; we consider T. inclinatum to be related to most other Thecadinium species and not to Sabulodinium. Molecular phylogenetic analyses based on the small subunit ribosomal gene of T. mucosum, T. kofoidii (the type species of the genus), and T. dragescoi weakly support earlier suspicions based on morphology that T. dragescoi is not a member of Thecadinium. Tabulational patterns of the species suggest a relationship to the genus Amphidiniopsis.     

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.     

PUNCTICULATA VERSIFORMIS SP. NOV. AND CYCLOTELLA KATHMANDUENSIS SP. NOV. (BACILLARIOPHYTA), NEW FOSSIL SPECIES FROM MIDDLE PLEISTOCENE LACUSTRINE SEDIMENTS,KATHMANDU, NEPAL HIMALAYA1

New fossil species, Puncticulata versiformis sp. nov. and Cyclotella kathmanduensis sp. nov., are described from lacustrine sediments in the Kathmandu Basin on the southern slope of the Nepal Himalaya. They were dominant in the Middle Pleistocene. Both LM and SEM observations reveal their unique morphological features. Puncticulata versiformis is characterized by (1) various valve outlines (circular, elliptical, or oval), (2) a tangentially undulate central area, (3) a complex alveolar structure composed of three kinds of costae (thick costae, thin costae, and thin and short costae), (4) well‐developed spines (Y‐shaped and tapering) located on one side of the valve face/mantle area junction, and (5) a valvocopula with an extremely undulate margin. The species‐specific feature of C. kathmanduensis is the presence of two kinds of alveolate zones in a single valve: type‐1 zone composed only of normal costae, and type‐2 zone composed of both normal costae and recessed costae bearing fultoportulae. Changes in valve ornamentation occur in these two species from initial valves to vegetative valves. In P. versiformis, the arrangement of areolae with internal domed cribra and fultoportulae in the central area changes from radial rows in the initial valve to groups in the vegetative valve. In the initial valve of C. kathmanduensis, the type‐1 alveolate zone is generally absent.     

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.     

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.     

SYMBIODINIUM NATANS SP. NOV.: A “FREE‐LIVING” DINOFLAGELLATE FROM TENERIFE (NORTHEAST‐ATLANTIC OCEAN)1

We examined a free‐living Symbiodinium species by light and electron microscopy and nuclear‐encoded partial LSU rDNA sequence data. The strain was isolated from a net plankton sample collected in near‐shore waters at Tenerife, the Canary Islands. Comparing the thecal plate tabulation of the free‐living Symbiodinium to that of S. microadriaticum Freud., it became clear that a few but significant differences could be noted. The isolate possessed two rather than three antapical plates, six rather than seven to eight postcingular plates, and finally four rather than five apical plates. The electron microscopic study also revealed the presence of an eyespot with brick‐shaped contents in the sulcal region and a narrow anterior plate with small knob‐like structures. Bayesian analysis revealed the free‐living Symbiodinium to be a member of the earliest diverging clade A. However, it did not group within subclade A_I (=temperate A) or any other subclades within clade A. Rather, it occupied an isolated position, and this was also supported by sequence divergence estimates. On the basis of comparative analysis of the thecal plate tabulation and the inferred phylogeny, we propose that the Symbiodinium isolate from Tenerife is a new species (viz. S. natans). To elucidate further the species diversity of Symbiodinium, particularly those inhabiting coral reefs, we suggest combining morphological features of the thecal plate pattern with gene sequence data. Indeed, future examination of motile stages originating from symbiont isolates will demonstrate if this proves a feasible way to identify and characterize additional species of Symbiodinium and thus match ribotypes or clusters of ribotypes to species.     

MESOPHYLLUM SPHAERICUM SP. NOV. (CORALLINALES,RHODOPHYTA): A NEW MAËRL‐FORMING SPECIES FROM THE NORTHEAST ATLANTIC1

Mesophyllum sphaericum sp. nov. is described based on spherical maërl individuals (up to 10 cm) collected in a shallow subtidal maërl bed in Galicia (NW Spain). The thalli of these specimens are radially organized, composed of arching tiers of compact medullary filaments. Epithallial cells have flattened to rounded outermost walls, and they occur in a single layer. Subepithallial initials are as long as, or longer than the daughter cells that subtend them. Cell fusions are abundant. Multiporate asexual conceptacles are protruding, mound‐like with a flattened pore plate, lacking a peripheral raised rim. Filaments lining the pore canal and the conceptacle roof are composed of five to six cells with straight elongate and narrow cells at their base. Carposporangial conceptacles are uniporate, protruding, and conical. Spermatangial conceptacles were not observed. Molecular results placed M. sphaericum near to M. erubescens, but M. sphaericum is anatomically close to M. canariense. The examination of the holotype and herbarium specimens of M. canariense indicated that both species have pore canal filaments with elongate basal cells, but they differ in number of cells (five to six in M. sphaericum vs. four in M. canariense). Based on the character of pore canal filaments, M. canariense shows similarities with M. erubescens (three to five celled). The outermost walls of epithallial cells of M. canariense are flared compared to the round to flattened ones of M. erubescens, the latter being widely accepted for the genus Mesophyllum. The addition of M. sphaericum as new maërl‐forming species suggests that European maërl beds are more biodiverse than previously understood.     

HETEROCAPSA ARCTICA SUBSP. FRIGIDA SUBSP. NOV. (PERIDINIALES,DINOPHYCEAE)—DESCRIPTION OF A NEW DINOFLAGELLATE AND ITS OCCURRENCE IN THE BALTIC SEA1

Characteristics important in identification of Heterocapsa species (i.e., thecal plate pattern, body scale structure, and shape and position of the nucleus and pyrenoid) are practically identical in the dinoflagellate investigated here and in Heterocapsa arctica T. Horig. described from the Canadian Arctic. Analysis of internal transcribed spacer (ITS) sequences confirms that the two dinoflagellates are very closely related; however, there is a clear difference in their size and shape. Our experiments show that the low‐salinity Baltic Sea brackish water does not reduce the size of the marine H. arctica to match that of the Baltic Sea morphotype. On the basis of these dissimilarities in general morphology and its geographic isolation in the Baltic Sea, we consider our material sufficiently differentiated from the typical H. arctica to warrant the status of a new subspecies, H. arctica subsp. frigida subsp. nov. Being of a distinct cell shape, the occurrence of subsp. frigida has been recorded in Algaline phytoplankton monitoring data collected since 1993. Although it has never been responsible for high biomass blooms, it commonly occurs in spring in the Northern Baltic Proper and in the western Gulf of Finland, when the water temperatures are <5°C.     

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.