Taxonomic studies on the Chlorarachniophyta. II. Generic delimitation of the chlorarachniophytes and description of Gymnochlora stellata gen. et sp. nov. and Lotharella gen. nov.

A new chlorarachniophytan alga, Gymnochlora stellata Ishida et Y. Hara gen. et sp. nov., has been isolated from Anae Island in Guam. It is a green, star-shaped, unicellular, amoeboid organism with several filopodia that do not form a reticulopodial network. Neither zoospores nor walled coccoid cells have been observed throughout the life cycle. The chloroplast ultrastructure is similar to those of described species; however, the pyrenoid matrix, which is invaded by many tubular structures originating from the inner membrane of the chloroplast envelope, is unique. A classification system is proposed for the Chlorarachniophyta. In this system, the ultrastructural features of the pyrenoid and the location of the nucleomorph in the periplastidial compartment are used as generic criteria, while the morphological features of the vegetative cells and life cycle patterns are used for species criteria. The described species, except for Cryptochlora perforans Calderon-Saenz et Schnetter, are also reassessed under the new system, and consequent nomenclatural requirements for the genus Chlorarachnion are dealt with in this paper. The taxonomic rank of a previously described species, Chlorarachnion globosum Ishida et Y. Hara, is elevated and Lotharella globosa (Ishida et Y. Hara) Ishida et Y. Hara gen. nov. et comb. nov. is proposed.     

Pleurosigma hinzianum sp. nov. and P. frenguellianum sp. nov. (Pleurosigmataceae,Bacillariophyta) from Argentinean coastal waters,in comparison with P. amara Stidolph and P. elongatum W. Smith

Two new marine diatom species from Argentinean coastal waters, Pleurosigma hinzianum Sterrenburg, Sunesen & Sar, sp. nov. and P. frenguellianum Sunesen, Sterrenburg & Sar, sp. nov., are described. The characters permitting their identification are specified, based on comparison with type material of the morphologically similar species P. amara Stidolph and P. elongatum W. Smith in the light (LM) and scanning electron microscope (SEM). New information on the type material of P. elongatum is presented and its taxonomic concept is emended. The main criteria for separation of the species discussed here are: valve outline, path of the raphe-sternum, raphe angle and angle of intersection of the oblique striae in LM; and external central and terminal raphe fissures, internal details of the central area, external and internal apical structure, and morphology of the hymen-occluded internal pores in SEM. The occurrence of these (and probably other) species in the genus Pleurosigma in net samples is adventitious and not indicative of a true planktonic mode of life.     

Grimmia serrana (Bryopsida,Grimmiaceae), a new species from California,U.S.A.

Abstract

Grimmia serrana, a new endemic species for California, is described and illustrated. The new species is readily distinguished by the presence of multi-stratose marginal bands that are rounded in cross-section. The leaf morphology displayed by Grimmia serrana is unique within the genus. The distribution of Grimmia serrana is along the western foothills of the Sierra Nevada and in the northern Coast Range.     

Phylogeny,species diversity and biogeographic patterns of the genus Tricleocarpa (Galaxauraceae,Rhodophyta) from the Indo-Pacific region,including T. confertus sp. nov. from Taiwan

Among four different genera in Galaxauraceae, species diversity of the genus Tricleocarpa Huisman & Borowitzka is underestimated and requires further examination. In this study, we explored a molecular phylogeny of Tricleocarpa with an emphasis on Taiwan species and reassessed species diversity of the genus from the Indo-Pacific region based on analyses of rbcL and COI-5P sequences and morphological observations. The molecular analyses showed that species of Tricleocarpa examined are grouped into seven clades: six clades in the T. cylindrica group (the T. cylindrica complex, T. confertus, T. jejuensis, T. natalensis and two uncharacterized T. ‘cylindrica’ clades) and one clade as the T. fragilis group. Overall, at least 12 species in Tricleocarpa were detected from the Indo-Pacific region, including a new species, T. confertus S.-L. Liu & S.-M. Lin, from Taiwan. Among them, T. confertus can be separated from other species of Tricleocarpa by a thallus consisting of dichotomous or trichotomous, articulated and moniliform branches with smooth surfaces, constrictions at nodes and slightly anti-triangular in shape. The cystocarp morphology of T. confertus is similar to the T. cylindrica group by having paraphyses intermixing with gonimoblast filaments. Recognition of this new species from other species is also supported by the DNA-based, algorithmic species delimitation approaches. With the combination of molecular phylogeny and species diversity reassessment, our phylogeographic analysis supported a trend that species in the T. fragilis group are restricted in their distribution to subtropical and tropical areas whereas species in the T. cylindrica group have a wide distribution, ranging from temperate through to tropical areas.     

MORPHOLOGY AND MOLECULAR ANALYSES OF THREE TOXIC SPECIES OF GAMBIERDISCUS (DINOPHYCEAE): G. PACIFICUS, SP. NOV., G. AUSTRALES, SP. NOV., AND G. POLYNESIENSIS, SP. NOV.

Three new dinoflagellate species, Gambierdiscus polynesiensis, sp. nov., Gambierdiscus australes, sp. nov., and Gambierdiscus pacificus, sp. nov., are described from scanning electron micrographs. The morphology of the three new Gambierdiscus species is compared with the type species Gambierdiscus toxicus Adachi et Fukuyo 1979, and two other species: Gambierdiscus belizeanus Faust 1995 and Gambierdiscus yasumotoi Holmes 1998. The plate formula is: Po, 3′, 7", 6C, 8S, 5‴, 1p, 2". Culture extracts of these three new species displayed both ciguatoxin- and maitotoxin-like toxicities. The following morphological characteristics differentiated each species. 1) Cells of G. polynesiensis are 68–85 μm long and 64–75 μm wide, and the cell’s surface is smooth. They are identified by a large triangular apical pore plate (Po), a narrow fish-hook opening surrounded by 38 round pores, and a large, broad posterior intercalary plate (1p) wedged between narrow postcingular plates 2‴ and 4‴. Plate 1p occupies 60% of the width of the hypotheca. 2) Cells of G. australes also have a smooth surface and are 76–93 μm long and 65–85 μm wide in dorsoventral depth. They are identified by the broad ellipsoid apical pore plate (Po) surrounded by 31 round pores and a long and narrow 1p plate wedged between postcingular plates 2‴ and 4‴. Plate 1p occupies 30% of the width of the hypotheca. 3) Cells of G. pacificus are 67–77 μm long and 60–76 μm wide in dorsoventral depth, and its surface is smooth. They are identified by the four-sided apical pore plate (Po) surrounded by 30 round pores. A short narrow 1p plate is wedged between the wide postcingular plates 2‴ and 4‴. Plate 1p occupies 20% of the width of the hypotheca. These three newly described species were also characterized by isozyme electrophoresis and DNA sequencing of the D8–D10 region of their large subunit (LSU) rRNA genes. The consistency between species designations based on SEM microscopy and classification inferred from biochemical and genetic heterogeneities was examined among seven isolates of Gambierdiscus. Their classification into four morphospecies was not consistent with groupings inferred from isozyme patterns. Three molecular types could be distinguished based on the comparison of their LSU rDNA sequences. Although G. toxicus TUR was found to be more closely related to G. pacificus, sp. nov. than to other G. toxicus strains, the molecular classification was able to discriminate G. polynesiensis, sp. nov. and G. australes, sp. nov. from G. toxicus. These results suggest the usefulness of the D8–D10 portion of the Gambierdiscus LSU rDNA as a valuable taxonomic marker.     

Haramonas viridissp. nov. (Raphidophyceae, Heterokontophyta), a new sand-dwelling raphidophyte from cold temperate waters

A new, marine, sand‐dwelling raphidophyte from Sylt, Germany, Haramonas viridis Horiguchi et Hoppenrath sp. nov. is described. This represents a second species in the previously monotypic genus Haramonas, which was originally described from a sand sample from a mangrove river mouth in tropical Australia, based on the type species, H. dimorpha. This new species from a cold temperate region: (i) possesses a tubular invagi‐nation in the posterior part of the cell; (ii) produces copious amounts of mucilage in culture; (iii) possesses both motile and non‐motile stages in its life cycle; and (iv) has overlapping discoidal chloroplasts, all of which are diagnostic features of the genus Haramonas. Therefore, it is indisputable that this species belongs to this genus. However, the species from Sylt differs from the type species of the genus in: (i) having a larger cell size; (ii) possessing a larger number of chloroplasts; and (iii) being greenish in color. The ultrastructural study revealed that the structure of the tubular invagi‐nation was the same as that of the type species.     

A revision of the genus Cryptonemia (Halymeniaceae,Rhodophyta) in Bermuda,western Atlantic Ocean,including five new species and C. bermudensis (Collins & M. Howe) comb. nov.

Cryptonemia specimens collected in Bermuda over the past two decades were analysed using gene sequences encoding the large subunit of the nuclear ribosomal DNA and the large subunit of RuBisCO as genetic markers to elucidate their phylogenetic positions. They were additionally subjected to morphological assessment and compared with historical collections from the islands. Six species are presently found in the flora including C. bermudensis comb. nov., based on Halymenia bermudensis, and the following five new species: C. abyssalis, C. antricola, C. atrocostalis, C. lacunicola and C. perparva. Of the eight species known in the western Atlantic flora prior to this study, none is found in Bermuda. Specimens reported in the islands in the 1900s attributed to C. crenulata and C. luxurians are representative of the new species, C. antricola and C. atrocostalis, respectively.     

Sargassum boreale sp. nov. (Fucales, Phaeophyceae) from Hokkaido, Japan

A new species, Sargassum boreale Yoshida et Horiguchi is described. It belongs to the subgenus Bactrophycus section Teretia, with cylindrical receptacles and is distinct from Sargassum confusum C. Agardh, S. pallidum (Turner) C. Agardh and Sargassum microceratium (Turner) C. Agardh in having a rather elongated stem with smooth surface and distantly issuing main branches, with narrow leaves. The distinction between S. boreale and these species is also revealed by a difference in internal transcribed spacer 2 (ITS‐2) sequences. In addition to the base substitutions, the existence of a large gap in S. boreale distinguishes this species from others. Sargassum boreale is distributed around Hokkaido and Saghalien to 50°N latitude. A key to the species of section Teretia is provided.     

NORTH CAROLINA MARINE ALGAE. IX. ONSLOWIA ENDOPHYTICA GEN. ET SP. NOV. (PHAEOPHYTA,SPHACELARIALES) AND NOTES ON OTHER NEW RECORDS FOR NORTH CAROLINA1

Four benthic algae are reported here for the first time in the North Carolina flora. The new brown algal genus and species, Onslowia endophytica Searles, is described as an endophyte of Halymenia floridana from the North Carolina continental shelf. New records of Boodleopsis pusilla and Naccaria corymbosa from North Carolina constitute range extensions of these tropical species on the American coast north from Florida. Blastophysa rhizopus, an endophyte and epiphyte known from the North Atlantic coast of Europe and America as well as the Caribbean is reported from North Carolina for the first time and in a new host, Predaea feldmannii.     

Cladosiphon takenoensis sp. nov. (Ectocarpales s.l., Phaeophyceae) from Japan

The new brown algal species Cladosiphon takenoensis H. Kawai (Chordariaceae, Ectocarpales s.l.) is described from Takeno, Hyogo, Japan based on morphology and DNA sequences. The species is a spring annual, growing on subtidal rocks at more or less exposed sites. It resembles C. umezakii in its gross morphology, and the two often grow together, but is distinguishable from C. umezakii in having a more hairy appearance. Cladosiphon takenoensis has a slimy, cylindrical, multiaxial and sympodial erect thallus, branching once to twice, and is provided with long assimilatory filaments (up to 1.8 mm long, composed of up to 100 cells). Unilocular zoidangia are formed on the basal part of assimilatory filaments. The species is genetically most related to C. umezakii and has the same basic thallus structures, but differs from C. umezakii and other Cladosiphon species in lacking phaeophycean hairs and plurilocular zoidangia of the assimilatory filaments. DNA sequences of the mitochondrial cox1 and cox3, chloroplast atpB, psaA, psbA and rbcL genes and the nuclear rDNA ITS2 region support the distinctness of the species. The genus Cladosiphon was paraphyletic in our analyses because the clades of C. okamuranus/C. zosterae and C. takenoensis/C. umezakii were split by Mesogloia vermiculata. However, since the genus‐level taxonomy of Chordariaceae needs considerable revision, we suspend the genus‐level taxonomy of the new species, and tentatively describe it as C. takenoensis.     

Facieplatycauda pratti gen. n., sp. n. and Two New Species of Myxobolus (Myxosporida)*

Synopsis. Facieplatycauda pratti gen. n., sp. n. and Myxobolus kozloffi sp. n., from the Klamath Lake sucker, Catostomus luxatus (Cope), are described. The new genus is placed in the family Myxobolidae between Henneguya and Myxobolus. The spore valves beyond the sporoplasmic space are broad and long but not attenuated as an inverted triangle as in certain species of Myxobolus with tails, or of Henneguya with sometimes bifurcated tails. An organism resembling Myxobolus insidiosus (Wyatt & Pratt) was found in the muscle of the body of Salmo clarki (Richardson). Differences in spore structure and site of infection suggest this is a subspecies of the previous form which should be designated Myxobolus insidiosus clarki. A previously unreported species of Myxobolus from Cottus aleuticus (Gilbert) is also described.     

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.     

Systematic revision of the genus Phycodrys (Delesseriaceae,Rhodophyta) from New Zealand,with the descriptions of three new species,P. novae-zelandiae sp. nov., P. franiae sp. nov. and P. adamsiae sp. nov.

Two species of Phycodrys, Phycodrys quercifolia (Bory) Skottsberg and Phycodrys profunda E.Y.Dawson were previously recorded from New Zealand. However, an examination of Phycodrys collections from the New Zealand region showed that all were morphologically different from P. quercifolia (Type locality: the Falkland Islands) and P. profunda (Type locality: CA, USA). RbcL sequence analyses established that the New Zealand Phycodrys species formed a natural assemblage within the genus, consisting of three new species: P. novae-zelandiae sp. nov., P. franiae sp. nov. and P. adamsiae sp. nov. Phycodrys novae-zelandiae is the largest of the three, up to 20 cm in height, with a distinct midrib and multicellular, opposite to subopposite lateral macroscopic veins. It has entirely monostromatic blades except near the midrib and veins, and its procarp contains a three-celled sterile group one (st1) and a one-celled sterile group two (st2). Phycodrys franiae was previously treated as a cryptic species among herbarium collections of P. ‘quercifolia’. It is smaller (4–11 cm high) with weakly developed midribs and veins, the blade is tristromatic throughout, except at the growing margins, and the procarp consists of a four-celled st1 and a two–three-celled st2. Phycodrys adamsiae, previously reported as P. profunda, is a small decumbent or prostrate plant, 1–8 cm long, with a midrib and inconspicuous lateral veins. The blades are tristromatic with serrated margins, two–four-celled surface spines and multicellular marginal holdfasts that differ from those of Californian specimens. The tetrasporangia are borne on marginal bladelets. Phylogenetic analyses place the New Zealand species in a separate group that is distantly removed from most other Phycodrys species.     

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.     

Research note: Amorphochlora amoebiformis gen. et comb. nov. (Chlorarachniophyceae)

A chlorarachniophycean alga, Lotharella amoebiformis, which has been classified in the genus Lotharella is placed into a new genus Amorphochlora gen. nov., based on its phylogenetic position, which has been clarified by the recently accumulated molecular phylogenetic information, and the morphological difference between the vegetative cells of the Lotharella species. Following this taxonomic treatment, a new combination Amorphochlora amoebiformis comb. nov., is proposed.     

CRYPTOPERIDINIOPSIS BRODYI GEN. ET SP. NOV. (DINOPHYCEAE), A SMALL LIGHTLY ARMORED DINOFLAGELLATE IN THE PFIESTERIACEAE1

A new genus and species of heterotrophic dinoflagellate, Cryptoperidiniopsis brodyi gen. et sp. nov., are described. This new species commonly occurs in estuaries from Florida to Maryland, and is often associated with Pfiesteria piscicida Steidinger et Burkholder, Pseudopfiesteria shumwayae (Glasgow et Burkholder) Litaker et al., and Karlodinium veneficum (Ballantine) J. Larsen, as well as other small (<20 μm) heterotrophic and mixotrophic dinoflagellates. C. brodyi gen. et sp. nov. feeds myzocytotically on pigmented microalgae and other microorganisms. The genus and species have the enhanced Kofoidian plate formula of Po, cp, X, 5′, 0a, 6″, 6c, PC, 5+s, 5″′, 0p, and 2″″ and are assigned to the order Peridiniales and the family Pfiesteriaceae. Because the Pfiesteriaceae comprise small species and are difficult to differentiate by light microscopy, C. brodyi gen. et sp. nov. can be easily misidentified.     

An ultrastructural study of Marsupiomonas pelliculata gen. et sp. nov., a new member of the Pedinophyceae

A new species, Marsupiomonas pelliculata gen. et sp. nov. (Pedinophyceae), is described. A single flagellum emerges from a deep pit with a distinctive thickened margin. The flagellum has rigid fibrillar hairs which are probably formed in the perinuclear space. A short second flagellar basal body lies within the cell close to the basal body of the emergent flagellum and the flagellar root system consists of striated and microtubular roots. There is a distinctive theca covering all but the anterior end of the cell and also a single large bright green chloroplast with an immersed pyrenoid surrounded by a starch shell. The wide salinity tolerance of the species is discussed in relation to its distribution in estuarine and salt marsh habitats. The salient features of the new species—the insertion of the emergent flagellum into a deep pit and the possession of a theca—are also seen in Pedinomonas tenuis, and it is suggested that P. tenuis could be transferred to the new genus Marsupiomonas. The class Pedinophyceae now includes three genera (Pedinomonas, Resultor and Marsupiomonas) and the distinguishing features are discussed.