MOLECULAR PHYLOGENY,ULTRASTRUCTURE, AND TAXONOMIC REVISION OF CHLOROGONIUM (CHLOROPHYTA): EMENDATION OF CHLOROGONIUM AND DESCRIPTION OF GUNGNIR GEN. NOV. AND RUSALKA GEN. NOV.1

We examined the molecular phylogeny and ultrastructure of Chlorogonium and related species to establish the natural taxonomy at the generic level. Phylogenetic analyses of 18S rRNA and RUBISCO LSU (rbcL) gene sequences revealed two separate clades of Chlorogonium from which Chlorogonium (Cg.) fusiforme Matv. was robustly separated. One clade comprised Cg. neglectum Pascher and Cg. kasakii Nozaki, whereas the other clade included the type species Cg. euchlorum (Ehrenb.) Ehrenb., Cg. elongatum (P. A. Dang.) Francé, and Cg. capillatum Nozaki, M. Watanabe et Aizawa. On the basis of unique ultrastructural characteristics, we described Gungnir Nakada gen. nov. comprising three species: G. neglectum (Pascher) Nakada comb. nov., G. mantoniae (H. Ettl) Nakada comb. nov., and G. kasakii (Nozaki) Nakada comb. nov. We also emended Chlorogonium as a monophyletic genus composed of Cg. euchlorum, Cg. elongatum, and Cg. capillatum. Because Cg. fusiforme was distinguished from the redefined Chlorogonium and Gungnir by the structure of its starch plate, which is associated with pyrenoids, we reclassified this species as Rusalka fusiformis (Matv.) Nakada gen. et comb. nov.     

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.     

THE FINE STRUCTURE AND SCALE FORMATION OF CHRYSOLEPIDOMONAS DENDROLEPIDOTA GEN. ET. SP. NOV.(CHRYSOLEPIDOMONADACEAE FAM. NOV., CHRYSOPHYCEAE)1

Chrysolepidomonas gen. nov. is described for single-celled monads with two flagella, a single chloroplast, and distinctive canistrate and dendritic scales. The type species, Chrysolepidomonas dendrolepidota sp. nov., is described for the first time. The canistrate scales bear eight “bumps” on the top surface, and the dendriticscales have a tapered base with a quatrifid tip. These organic scales are formed in the Golgi apparatus and storred in a scale reservoir. The scale reservoir is bounded on two sides by the R₁ and R₂ in microtubular roots of the basal apparatus. The cyst (=stomatocyst, statospore) forms endogenously by means of a silica deposition vesicle. The outer cyst surface is smooth, and the pore region is unornamented. Two other organisms bearing canistrate and dendritic scales, previously assigned to the genus Sphaleromants, are transferred to the genus Chrysolepidomonas. They are C.angalica sp. nov. and C. marine(Pienaar) comb. nov. The distinguishing features of Chrysolepidomonas and Sphaleromantis are discussed. A new family, Chrysolepidomonadceae fam. noc., is described for flagellates covered with organic scales.     

GAYLIELLA GEN. NOV. IN THE TRIBE CERAMIEAE (CERAMIACEAE,RHODOPHYTA) BASED ON MOLECULAR AND MORPHOLOGICAL EVIDENCE1

On the basis of comparative morphology and phylogenetic analyses of rbcL and LSU rDNA sequence data, a new genus, Gayliella gen. nov., is proposed to accommodate the Ceramium flaccidum complex (C. flaccidum, C. byssoideum, C. gracillimum var. byssoideum, and C. taylorii), C. fimbriatum, and a previously undescribed species from Australia. C. transversale is reinstated and recognized as a distinct species. Through this study, G. flaccida (Kützing) comb. nov., G. transversalis (Collins et Hervey) comb. nov., G. fimbriata (Setchell et N. L. Gardner) comb. nov., G. taylorii comb. nov., G. mazoyerae sp. nov., and G. womersleyi sp. nov. are based on detailed comparative morphology. The species referred to as C. flaccidum and C. dawsonii from Brazil also belong to the new genus. Comparison of Gayliella with Ceramium shows that it differs from the latter by having an alternate branching pattern; three cortical initials per periaxial cell, of which the third is directed basipetally and divides horizontally; and unicellular rhizoids produced from periaxial cells. Our phylogenetic analyses of rbcL and LSU rDNA gene sequence data confirm that Gayliella gen. nov. represents a monophyletic clade distinct from most Ceramium species including the type species, C. virgatum. We also transfer C. recticorticum to the new genus Gayliella.     

Morphology and phylogeny of a new wall‐less freshwater volvocalean flagellate,Hapalochloris nozakii gen. et sp. nov. (Volvocales,Chlorophyceae)

New strains of a wall‐less unicellular volvocalean flagellate were isolated from a freshwater environment in Japan. Observations of the alga, described here as Hapalochloris nozakii Nakada, gen. et sp. nov., were made using light, fluorescence, and electron microscopy. Each vegetative cell had two flagella, four contractile vacuoles, and a spirally furrowed cup‐shaped chloroplast with an axial pyrenoid, and mitochondria located in the furrows. Based on the morphology, H. nozakii was distinguished from other known wall‐less volvocalean flagellates. Under electron microscopy, fibrous material, instead of a cell wall and dense cortical microtubules, was observed outside and inside the cell membrane, respectively. Based on the phylogenetic analyses of 18S rRNA gene sequences, H. nozakii was found to be closely related to Asterococcus, Oogamochlamys, Rhysamphichloris, and “Dunaliella” lateralis and was separated from other known wall‐less flagellate volvocaleans, indicating independent secondary loss of the cell wall in H. nozakii. In the combined 18S rRNA and chloroplast gene tree, H. nozakii was sister to Lobochlamys.     

DEUCALION GEN. NOV. AND ANISOSCHIZUS GEN. NOV. (CERAMIACEAE,CERAMIALES), TWO NEW PROPAGULE-FORMING RED ALGAE FROM SOUTHERN AUSTRALIA1

Two new propagule-farming red algae from southern Australia, Deucalion levringii (Lindauer) gen. et comb. nov. and Anisoschizus propaguli gen. et sp. nov., are described and defined largely on their development in laboratory culture. Deucalion is included in the tribe Compsothamnieae on the basis of its subapical procarp and alternate distichous branching. It differs from the other genera included in that tribe in that it produces 3-celled propagules, polysporangia, a subapical cell of the fertile axis which bears 3 pericentral cells, and an apparently post-fertilization involucre which develops from the hypogenous and sub-hypogenous cells of the fertile axis. Its gametophyte morphology has been elucidated in culture, as only sporophytes are known from the field. Gametophytes do not appear to produce propagules. Anisoschizus is provisionally included in the tribe Spermothamnieae on the basis of its subdichotomous branching, possession of a prostrate system and the production of polysporangia. It differs from the other genera of the tribe in the production of 2-celled propagules. Observations on the germination of the “monosporangia” of Mazoyerella arachnoidea and Monosporus spp. indicate that they are analagous to the propagules of Deucalion and Anisoschizus. The nature of these propagules and their role in recycling the parent plant are discussed and contrasted with true monosporangia. It is recommended that Monosporus be maintained as a form genus containing representatives from more than one tribe, as exemplified by plants from Lord Howe I. provisionally identified as M. indicus Boergesen which have both prostrate and erect, as opposed to only erect, axes.     

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.     

LIFE HISTORY AND TAXONOMY OF RHIZOOCHROMONAS ENDOLORICATA GEN. ET SP. NOV., A NEW FRESHWATER CHRYSOPHYTE INHABITING DINOBRYON LORICAE1

The life cycle of a previously undescribed chrysophyte, assigned to the new genus Rhizoochromonas, is described. It includes a small motile stage with heterokont flagellation which invades a Dinobryon lorica. Reproduction by cell division of the nearly spherical rhizopodial vegetative stage frequently leads to expulsion of the host protoplast through overcrowding of the lorica. Endogenous cysts (stomatocysts) are also formed within the Dinobryon lorica. The new family, Brehmiellaceae, is established to accommodate pseudopodial/rhizopodial chrysophytes with heterokont flagellation in the motile stage. Rhizoochromonas endoloricata gen. et sp. nov. has been found at two widely separated softwater locations in Ontario, and at one it constituted a major component of the planktonic flora during the autumns of six successive years.     

MORPHOLOGY OF THE SUBANTARCTIC RED ALGA CENACRUM SUBSUTUM GEN. ET SP. NOV. (RHODYMENIALES) FROM MACQUARIE ISLAND1

The red alga Cenacrum subsutum gen. et sp. nov. is described from material collected at Macquarie Island in the subantarctic between November 1977 and February 1978. The habit and carposporophyte development are similar to members of the family Rhodymeniaceae (Rhodymeniales), but certain vegetative features are unique. The frond is a variously incised or lobed foliose blade with hollow apices above and a medulla which becomes progressively filled basipetally with ingrowing rhizoidal filaments. Details of carpogonial branch, auxiliary cell, connecting cell and gonimoblast anatomy are given, as well as observations on the habitats and distribution of the species.     

ATRACTOMORPHA ECHINATA GEN. ET SP. NOV., A NEW ANISOGAMOUS MEMBER OF THE SPHAEROPLEACEAE (CHLOROPHYCEAE)1,2

Atractomorpha echinata gen. et sp. nov. is described from isolates derived from zygotes present in a dry soil sample obtained from Texas. The new genus is distinguished from Sphaeroplea primarily by its pattern of vegetative growth. While Sphaeroplea is distinctly filamentous with numerous coenocytic cells uniseriately arranged, Atractomorpha grows as individual, multinucleate, spindle-shaped cells with sharply pointed extremities. Such cells may vary considerably in length (25–6000 μm, or more) and normally lack septa. In young, rapidly growing cultures the cells often attain lengths of 300–500 μm, but rarely exceed 1800 μm. The new species is further characterized by: (1) the regular formation of biflagellate zoospores in asexual reproduction, (2)anisogamy (occasionally oogamy) and (3) the size and ornamentation of its zygotes. Variations in vegetative morphology are discussed as are conditions for obtaining gametogenesis.     

Chlamydomonas neoplanoconvexa nom. nov. and its unique phylogenetic position within Volvocales (Chlorophyceae)

Chlamydomonas Ehrenb. is a unicellular volvocalean genus consisting of 400–600 species, most of which are known solely based on microscopy. In this study, a newly isolated strain of Chlamydomonas neoplanoconvexa (M. O. P. Iyengar) Nakada nom. nov. (≡Chlamydomonas planoconvexa M. O. P. Iyengar non J. W. G. Lund) was examined using light microscopy and combined 18S rRNA, rbcL and psaB gene phylogeny. The C. neoplanoconvexa strain was quite similar to Chlamydomonas perpusilla (Korshikov) Gerloff in terms of its small fusiform vegetative cells (～10 µm in length), parietal chloroplast containing a single pyrenoid, and nucleus posterior to the pyrenoid. However, C. neoplanoconvexa was distinguished from C. perpusilla based on the morphology of the papillae and pyrenoids. The two species belong to the clade Caudivolvoxa in the order Volvocales, but are distantly related to each other. Chlamydomonas perpusilla was shown to be sister to Chlorogonium Ehrenb. (clade Chlorogonia, within Caudivolvoxa), while C. neoplanoconvexa represented a second basal lineage within Caudivolvoxa, next to the clade Characiosiphonia. Although the morphology of C. neoplanoconvexa was not particularly outstanding, its unique phylogenetic position will encourage further investigation of this species and its uncultivated relatives.     

PROPOSAL OF THE EUPTILOTEAE HOMMERSAND ET FREDERICQ,TRIB. NOV. AND TRANSFER OF SOME SOUTHERN HEMISPHERE PTILOTEAE TO THE CALLITHAMNIEAE (CERAMIACEAE,RHODOPHYTA)1

Morphological and molecular studies demonstrate that the tribe Ptiloteae (Ceramiaceae, Ceramiales) is polyphyletic. The Ptiloteae, sensu stricto, occur only in the Northern Hemisphere and all Southern Hemisphere representatives belong in other tribes. Three genera (Euptilota, Seirospora, and Sciurothamnion) are transferred to the Euptiloteae Hommersand et Fredericq, trib. nov., and the Callithamnieae is revised to include three Ptilota‐like genera, Georgiella, Falklandiella, and Diapse, and two new genera. Heteroptilon Hommersand, gen. nov. is erected to receive Euptilota pappeana Kützing 1849 and Aglaothamnion rigidulum De Clerck, Bolton, Anderson et Coppejans 2004 from South Africa, and Aristoptilon Hommersand et W. A. Nelson, gen. nov. is established to receive Euptilota mooreana Lindauer 1949 from New Zealand. The principal difference between the Euptiloteae and the Callithamnieae is seen in the earliest stages after fertilization. The fertilized carpogonium enlarges and forms a pair of tube‐like protuberances directed toward the auxiliary cells that are cut off as connecting cells in the Euptiloteae, whereas in the Callithamnieae the carpogonium usually divides into two cells, each of which cuts off a small connecting cell that fuses with an adjacent enlarging auxiliary cell. Nuclei are terminal in spermatangia of the Euptiloteae, subtended by mucilaginous vesicles, and are medial in the Callithamnieae situated between apical and basal vesicles. The Euptiloteae and Callithamnieae (including the Ptilota‐like members) are each strongly supported in maximum‐likelihood tree topologies resulting from analyses of combined 18S rDNA, 28S rDNA, 16S rDNA, and rbcL data sets. Their sister relationship is also well supported.     

STARRIA ZIMBABWEËNSIS (CYANOPHYCEAE) GEN. NOV. ET SP. NOV.: A FILAMENT TRIRADIATE IN TRANVERSE SECTION1

A new filamentous bluegreen alga Starria zimbabweënsis gen. nov. et sp. nov. isolated from a soil sample collected near Zimbabwe, Southern Rhodesia, has a unique triradiate morphology. In transverse section each narrow cell possesses three arm-like projections, separated by 120° and in which pigment is concentrated. Filaments may be straight or twisted and various clonable biradiate forms have originated in cultures initiated from the triradiate type. Cell ultra-structure is typical of the Oscillatoriaceae except that 70 nm pit-like pores occur throughout the L-II layer of the longitudinal walls. Wall structure of deviant forms is identical to that of the triradiate “wild type.” The organism is assigned to the Oscillatoriaceae rather than to the Gomontiellaceae or to a new family.     

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.     

FURTHER STUDIES OF PRESUMEDLY PRIMITIVE GREEN ALGAE,INCLUDING THE DESCRIPTION OF PEDINOPHYCEAE CLASS. NOV. AND RESULTOR GEN. NOV.1

The tiny jumping flagellate originally described as Pedinomonas mikron Throndsen was isolated into pure culture from Australian waters and its ultrastructure critically examined. Pedinomonas mikron differs in behavior and in features of the flagellar apparatus from P. minor, the type species from freshwater, and is referred to the new genus Resultor. The two genera are closely related and form the new class Pedinophyceae, which is characterized by features of the flagellar apparatus, mitosis, and cytokinesis. The flagella show the 11/5 orientation otherwise characteristic of Ulvophyceae and Pleurastrophyceae, but they are arranged end to end as in the Chlorophyceae. The flagellar root system is asymmetric and includes a rhizoplast that emerges from the base of one flagellum but subsequently associates with a microtubular root from the second basal body. Mitosis studied previously by Pickett-Heaps and Ott in Pedinomonas is closed, unlike in other green algae, and the spindle is persistent. No phycoplast or phragmoplast is formed during cytokinesis. The eyespot of the Pedinophyceae is located at the opposite end of the cell from the flagella and adjacent to the pyrenoid, as in the most primitive members of the Prasinophyceae. Members of the Pedinophyceae lack prasinoxanthin and Mg 2,4D, characteristic of certain other primitive green algae. The primitive green algae include the classes Prasinophyceae and Pedinophyceae. Micromonadophyceae Mattox et Stewart is considered a synonym of Prasinophyceae. Two new orders are established, Pedinomonadales, containing all known members of the Pedinophyceae, and Scourfieldiales, with the single family Scourfieldiaceae fam. nov. and the single genus Scourfieldia.     

MORPHOLOGY AND SYSTEMATICS OF SCIUROTHAMNION STEGENGAE GEN. ET SP. NOV. (CERAMIACEAE,RHODOPHYTA) FROM THE INDO‐PACIFIC1

A new ceramiaceous alga, Sciurothamnion stegengae De Clerck et Kraft, gen. et sp. nov., is described from the western Indian Ocean and the Philippines. Sciurothamnion appears related to the tribe Callithamnieae on the basis of the position and composition of its procarps and by the majority of post‐fertilization events. It differs, however, from all current members of the tribe by the presence of two periaxial cells bearing determinate laterals per axial cell. Additionally, unlike any present representative of the subfamily Callithamnioideae, no intercalary foot cell is formed after diploidization of the paired auxiliary cells. The genus is characterized by a terminal foot cell (“disposal cell”), which segregates the haploid nuclei of the diploidized auxiliary cell from the diploid zygote nucleus. The nature of three types of foot cells reported in the Ceramiaceae (intercalary foot cells containing only haploid nuclei, intercalary foot cells containing haploid nuclei and a diploid nucleus, and terminal foot cells containing only haploid nuclei) is discussed.     

STRUCTURE AND REPRODUCTION OF AMANSIA AND MELANAMANSIA GEN. NOV. (RHODOPHYTA,RHODOMELACEAE) ON THE SOUTHEASTERN AFRICAN COAST1

Four species of Amansia Lamouroux were initially found in Natal. More Complete studies on these species revealed a new genus, Melanamansia, Which is described on the basis of presence of two dorsal pseudopericentral cells in two new species from Natal (M. seagriefii sp. nov. & M. fimbrifolia sp. nov.) in addition to other structural characters and features of pigmentation and reproduction. Pseudopericentral cells are not present in the type species of Amansia, A. multifida Lamouroux. The other two species of Amansia occurring in Natal, A. glomerata C. Agardh & A. loriformis sp. nov., have characters similar to the type species. Comparison of species from other regions of the world has shown that eight additional species, previously assigned to Amansia, belong to the new genus.     

CHLOROPELTA GEN. NOV., AN ULVACEOUS GREEN ALGA WITH A DIFFERENT TYPE.OF DEVELOPMENT1

A diminutive, distromatic ulvaceous green alga was collected in southern California and studied in culture. The initial stages of development resemble those found in the Ulvaceae sensu Bliding. Germlings pass through a uniseriate filamentous stage, a multiseriate stage and a monostromatic saccate stage. At this stage the development departs from the developmental patterns found in the Ulvaceae. Each cell of the monostromatic upright portion of the germling undergoes a single division in a plane parallel to the surface of the germling to form a distromatic saccate germling. Rupture of the apical end of the germling and continued growth eventually results in a peltate distromatic alga superficially resembling Ulva. Based on the developmental pattern, which is unique to the green algae, the new genus Chloropelta gen. nov. and new species Chloropelta caespitosa sp. nov. are proposed for this alga.     

Description of a new diatom genus Dorofeyukea gen. nov. with remarks on phylogeny of the family Stauroneidaceae

Type material of Navicula kotschyi was studied, and this species was transferred to Dorofeyukea gen. nov. as D. kotschyi comb. nov. Dorofeyukea was described on the basis of DNA sequence and morphological data. Additional species assigned to this genus that were previously included in Navicula include: D. ancisa comb. nov., D. grimmei comb. nov., D. ivatoensis comb. nov., D. orangiana comb. nov., D. rostellata comb. nov. & stat. nov., D. savannahiana comb. nov., D. tenuipunctata comb. nov., and D. texana comb. nov. All Dorofeyukea species share the same morphological features, including having a narrow stauroid fascia surrounded by 1–3 irregularly shortened striae, uniseriate, and weakly radiate striae, circular, or rectangular puncta in the striae that are covered internally by dome‐shaped hymenes, presence of a pseudoseptum at each apex and absence of septa. Partial DNA sequences of SSU and rbcL loci show Dorofeuykae belongs to the clade of stauroneioid diatoms together with Stauroneis, Prestauroneis, Craticula, Karayevia, Madinithidium, Fistulifera, Parlibellus, and, possibly, Schizostauron. A new species from the monoraphid genus Madinithidium, M. vietnamica sp. nov., was described based on valve and chloroplast morphology as well as DNA sequence data.     

PYCNOCOCCUS PROVASOLII GEN. ET SP. NOV., A COCCOID PRASINOXANTHIN-CONTAINING PHYTOPLANKTER FROM THE WESTERN NORTH ATLANTIC AND GULF OF MEXICO1

The new genus Pycnococcus Guillard is based on several clones from the western North Atlantic and Gulf of Mexico. The type and only described species, Pycnococcus provasolii Guillard, sp. nov., is typified by clone Ω48-23 from the North Atlantic. Cells of Pycnococcus provasolii are solitary, spherical, 1.5–4.0 μm in diameter, have a resistant cell wall lacking sporopollenin, and have the ultrastructural characteristics of green algae. With the light microscope they are scarcely distinguishable from cells of other coccoid planktonic organisms. In pigmentation P. provasolii resembles Micromonas pusilla, Mantoniella squamata, and Mamiella gilva in having chl a, much chl b, Mg 2,4-divinylphaeoporphyrin a₅ monomethyl ester (presumably), and prasinoxanthin as a major xanthophyll. The pyrenoid of P. provasolii has a cytoplasmic channel, which is unique among species closely related to it. Flagellates, occurring rarely in culture, are similar to but distinguishable from known Pedinomonas species by size and shape. Pycnococcus provasolii is referred to the new family Pycnococcaceae Guillard, in the order Mamiellales of the class Micromonadophyceae (Chlorophyta). Clones of Pycnococcus provasolii are oceanic in nutritional characteristics, require only vitamin B₁₂ in culture, and are well adapted to growth under blue or blue-violet light of low intensity.