Characterization of the edible red alga Meristotheca papulosa (Solieriaceae, Gigartinales) from Japan

The vegetative and reproductive morphology of the edible red alga Meristotheca papulosa (Montagne) J. Agardh (Solieriaceae) was reexamined based on material collected from various localities in Japan. Although the habit of the blades is variable according to the length and width of the axes, the frequency of branching and the abundance of proliferations, rbcL sequence analyses indicate their conspecificity. M. papulosa displays four distinctive reproductive features (presence of an auxiliary cell complex, occurrence of cystocarps on marginal proliferations and the blade surface (although very rare) in addition to the margins of axes, frequent production of spinose outgrowths on the pericarp and tetrasporangial initials typically basally attached to their parental cells) that have not been reported for M. papulosa from other areas. Although these features might warrant recognition of the Japanese entity as a separate species, a better understanding of their possible taxonomic value requires comparisons with M. papulosa from other geographic regions, including the type locality.     

TWO SOLIERIACEAN ALGAE NEW TO THE MEDITERRANEAN: AGARDHIELLA SUBULATA AND SOLIERIA FILIFORMIS (RHODOPHYTA,GIGARTINALES)1

Two Atlantic species found in the Mediterranean Sea were ascribed to the family Solieriaceae on the basis of morphological features. In the extensive drifting beds of the Mar Piccolo basin (Taranto, Ionian Sea), Agardhiella subulata (C. Agardh) Kraft et Wynne and Solieria filiformis (Kützing) Gabrielson were the dominant species in summer. There were no attached populations, and perennation was ensured only by vegetative propagation. Characteristics of the outer cortex and chromatophores were useful for distinguishing the taxa at the species level and are recommended for recognition of sterile specimens.     

Consideration of the order Cryptonemiales and the families Nemastomataceae and Furcellariaceae (Gigartinales,Rhodophyta) in light of the morphology of Adelophyton corneum (J. Agardh) gen. et comb. nov. from southern Australia

A new red algal genus is described, based on the southern Australian Chaetangium corneum J. Agardh. It is reproductively unique in that while the auxiliary cell is intercalary in an adventitious filament, a defining character of the order Cryptonemiales, the vegetative structure, carpogonial branches, connecting filaments and gonimoblast development seem strongly allied to lower families of the Gigartinales. Although its predominantly thallus-inward gonimoblast development is characteristic of the gigartinalian family Furcellariaceae, it is suggested that the new alga should be placed in the Nemastomataceae and that this family exhibits features which, in an ancient algal stock, could theoretically have given rise to the lower cryptonemialian and higher gigartinalian lines. The possible relationships between the Nemastomataceae, Furcellariaceae and Solieriaceae are discussed, and some seemingly primitive features of the Nemastomataceae are enumcrated.     

THE ATLANTIC SPECIES OF SOLIERIA (GIGARTINALES,RHODOPHYTA): THEIR MORPHOLOGY,DISTRIBUTION AND AFFINITIES1

Solieria chordalis (C. Agardh) J. Agardh and S. tenera (J. Agardh) Wynne et Taylor exhibit multiaxial growth from a cluster of four to eight obconical apical cells. A single periaxial cell is cut off from each axial cell and successive periaxial cells are rotated 120° in a zig-zag pattern along each axial filament. Periaxial cells produce branched, laterally diverging filaments which form the cortex. The medulla is composed of axial cells, elongate cells of lateral filaments, stretched interconnecting cells, and secondary rhizoids. The two species are nonprocarpic. Carpogonial branches are 3-celled, inwardly directed, with a reflexed trichogyne. The auxiliary cell together with associated darkly-staining inner cortical cells form an association, the auxiliary cell complex, that is recognizable prior to diploidization. A single, unbranched, non-septate connecting filament issues from the fertilized carpogonium and fuses with the inner, lateral side of an auxiliary cell. Production of an involucre from surrounding vegetative cells is stimulated and a gonimoblast initial is cut off toward the interior of the thallus which divides to form a compact cluster of gonimoblast cells. A fusion cell is produced through fusion of inner gonimoblast cells with the auxiliary cell that, in turn, fuses progressively with cells of the lateral file bearing the auxiliary cell. Mature cystocarps have terminal carposporangia cut off from gonimoblast cells at the periphery of the fusion cell and are surrounded by an involucre with a distinct ostiole. Tetrasporangia are cut off laterally from surface cortical cells which then cut off one or two additional derivatives toward the outside. A lectotype is designated for Solieria chordalis, but the lectotypification of S. tenera is questioned. We conclude that Solieria is closely related to Rhabdonia and place the Rhabdoniaceae in synonomy with the Solieriaceae.     

MORPHOLOGY OF PLACENTOPHORA (SOLIERIACEAE,GIGARTINALES: RHODOPHYTA), A NEW GENUS BASED ON SARCODIOTHECA COLENSOI FROM NEW ZEALAND1

The only member of the red algal family Solieriaceae known from New Zealand is the endemic Sarcodiotheca colensoi (Hook. & Harv.) Kylin. This study shows that it differs in several respects from the type S. furcata (Setch. & Gard.) Kylin; thus a new genus Placentophora is created for the New Zealand alga. Although P. colensoi nov. comb. is retained in the Solieriaceae on the basis of vegetative, spermatangial, tetrasporangial, carpogonial-branch and early gonimoblast features, it differs from typical members of that family in its pattern of later carposporophyte development. After a single gonimoblast initial is cut off from the auxiliary cell towards the center of the thallus, further gonimoblasts develop from the initial as ramifying, radiating filaments. These filaments enter an extensive “nutritive-cell” region surrounding the auxiliary cell, form, numerous connections to the “nutritive” cells, and incorporate most of them into a central placenta of interconnected, and variously-fused vegetative and gonimoblast cells. Carpo-sporangia then form in short chains around the periphery of the placenta. The cystocarp lacks both a central fusion cell and a sterile-celled investment, or “Faserhülle.” The distinctive carposporophyte of Placentophora is compared to patterns of gonimoblast development, known in other members of the Solieriaceae.     

MORPHOLOGY AND TAXONOMY OF MERISTIELLA GEN. NOV. (SOLIERIACEAE,RHODOPHYTA) 1

Eucheuma acanthocladum (Harvey) J. Agardh, E. gelidium (J Agardh) J. Agardh, E. echinocarpum Areschoug and E. schrammii(P. et H. Crouan) J. Agardh from the tropical and warm temperate waters of the western Atlantic Ocean and Caribbean Sea are transferred to a new genus, Meristiella. Meristiella exhibits the following Unique combination of characters among genera in the Solieriaceae: (1) rotated periaxial cells, (2) a loosely filamentous medulla. (3) an auxiliary cell complex, (4) Single and twin connecting filaments and (5) spinose cystocarps composed of a central, small-celled placentum, based on its reproductive features, Meristiella. is assigned to the tribe Agardhielleae. Culture experiments and herbarium studies provide evidence that E, gelidium and E. acantghocladum are conspecific. Lectotypes are designated for the included species.     

Morphology of Laurencia clavata and L. elata (Ceramiales,Rhodophyta) in relation to generic circumscription in the Laurencia complex

Laurencia Lamouroux (Rhodophyta) was recently separated into three genera–Laurencia, Chondrophycus (Tokida et Saito) Garbary et Harper and Osmundea Stackhouse – each of which was newly defined based on vegetative and reproductive structures. In this study, the previously unknown vegetative and reproductive morphology of two Australian endemic species of Laurencia, L. clavata Sonder and L. elata (C. Agardh) Harvey, was studied, particularly in the context of the revised generic delineation. These species exhibit vegetative axes with four pericentral cells and trichoblast-type spermatangial development. Tetrasporangia are abaxially produced from the existing third and fourth pericentral cells. L. clavata has terete thalli with distinctive verticillate branching and is similar to Chondria C. Agardh, rather than to Laurencia, in having an unusually marked constriction at the base of the branches and starch accumulation in subcortical and medullary cells. Compared to Laurencia, apical cells of this species exhibit a less oblique division ; the resulting recognizable axial cell rows extend somewhat below the branches, and particularly at a young stage they are also clear throughout branchlets. However, other vegetative and spermatangial structures show that L. clavata is more closely allied to Laurencia than to Chondria, and it is placed in Laurencia. By contrast, L. elata exhibits morphology typical of Laurencia and is characterized by large, robust, compressed thalli with fastigiately distichous branching and an extensive secondary cortex. Furthermore, it appears to be distinct from similar species in sometimes having a parasitic species of Janczewskia Solms-Laubach (Rhodophyta). Taxonomy of Laurencia is discussed on the basis of these and previous studies.     

A morphological and molecular assessment of the genus Prionitis J. Agardh (Halymeniaceae, Rhodophyta)

A critical reassessment of the morphological features of two closely related red algal genera, Grateloupia C. Agardh and Prionitis J. Agardh (Halymeniaceae), shows that members of the two genera share very similar reproductive (including the Grateloupia‐type auxiliary‐cell ampullae) and vegetative characters. Diagnostic features hitherto used for distinguishing these two genera, the texture of blades (lubricous to leathery in Grateloupia vs cartilaginous in Prionitis) and the position of reproductive structures (scattered over the entire blade in Grateloupia vs confined to particular portions of the blade in Prionitis), are continuous across some 75 species of both genera, thus making it difficult to draw a clear‐cut distinction between the two genera. In ribulose‐1,5‐bisphosphate carboxylase/oxygenase gene (rbcL) sequence analyses, the species of Grateloupia and Prionitis, including the two generitypes, constitute a large monophyletic clade in the Halymeniaceae. It is therefore proposed that Prionitis be included in the synonymy under Grateloupia and the appropriate combinations are proposed.     

REPRODUCTIVE DEVELOPMENT OF THE PARASITIC RED ALGA GARDNERIELLA TUBERIFERA (SOLIERIACEAE,GIGARTINALES)1

Examination of the reproductive morphology of the adelphoparasitic red alga Gardneriella tuberifera Kylin reveals that this monotypic genus is correctly placed in the family Solieriaceae (Gigartinales), to which its host Agardhiella gaudichaudii (Montagne) Silva et Papenfuss also belongs. Gardneriella is multiaxial, nonprocarpic and has an inwardly directed, three-celled carpogonial branch. The large, reniform uninucleate auxiliary cell is distinct prior to and after fertilization. It is diploidized by an unbranched, multicellular connecting filament which lacks pit connections. One or two connecting filaments arise from each fertilized carpogonium. From the diploidized auxiliary cell, the gonimoblast initial is cut off obliquely toward the interior of the thallus. The cells of the gonimoblast fuse with adjacent unpigmented vegetative cells of Gardneriella and pigmented cells of the host. These cells become incorporated into the developing cystocarp and, from those of Gardneriella, additional short chains of gonimoblast cells arise. The mature cystocarp is placentate, radiately lobed, and lacks a surrounding involucre. Carposporangia are borne in short chains and the unpigmented carpospores are released upon the dissolution of outer vegetative cells. No ostiole is present. Gardneriella appears to be most closely related to the placentate solieriacean genera Agardhiella, Sarcodiotheca, and Meristiella and therefore this genus should be placed in the tribe recently erected for these taxa, the Agardhielleae.     

A new species of Laurencia, L. omaezakiana (Ceramiales, Rhodophyta), from Japan

Laurencia omaezakiana Masuda, sp. nov. (Ceramiales, Rhodophyta) is described from Japan. It is characterized by the following set of features: (i) the production of four periaxial cells from each vegetative axial cell; (ii) a shift in branching from distichous to spiral; (iii) the presence of projecting superficial cortical cells near the apices of branches; (iv) the presence of longitudinally oriented secondary pit-connections between contiguous superficial cortical cells; (v) the presence of lenticular thickenings in the walls of medullary cells; (vi) the occurrence of 1–2 corps en cerise in each superficial cortical cell and a single corps en cerise in each trichoblast cell; and (vii) a parallel arrangement of tetrasporangia. Furthermore, it produces a characteristic triterpenoid (enshuol), which has not been detected in other species of Laurencia, as a major halogenated secondary metabolite. A synoptical key to the 23 species of Laurencia growing in Japan is given. Laurencia ceytanica J, Agardh and Laurencia heteroclada Harvey are excluded from the Japanese marine algalflora. The latter is a distinct species from Laurencia filiformis (C. Agardh) Montagne.     

Observations on Japanese species of the genus Martensia (Delesseriaceae, Rhodophyta), with the description of Neomartensia gen. nov.

The morphology of vegetative and reproductive structures of Martensia fragilis Harvey, Martensia australis Harvey and Martensia flabelliformis Harvey ex J. Agardh was studied based on material collected from Japan. M. fragilis and M. australis have cystocarps with terminal carposporangia, and their membranous portions are composed of cell layers with regularly stacked flat cells of similar size. M. flabelliformis has the following combination of characters: (i) carposporangia are formed in short chains; and (ii) the membranous portion is composed of irregularly arranged flat cells of various sizes. These characters are distinctly different from those of Martensia and warrant the establishment of Neomartensia gen. nov., with the type species Neomartensia flabelliformis comb. nov.     

Carrageenan of <Emphasis Type="Italic">Eucheuma isiforme</Emphasis> (Solieriaceae,Rhodophyta) from Nicaragua

The yield and physicochemical properties of native and alkali treated carrageenan from Eucheuma isiforme harvested from the Nicaraguan coast were investigated. The native carrageenan yield was 57.2% of dry weight and decreased to 43.5% when the alga was alkali treated. Native carrageenan viscosities showed significant differences between native (144.6 ± 3.3 cPs) and treated carrageenan (113.9 ± 2.6 cPs) (p < 0.01). Alkali treatment reduced carrageenan sulphate content by 19.3% and increased 3,6 AG content by 13%. Alkali-treated carrageenan formed very weak gels in 1.5% solutions (<50 g cm⁻²). Chemical analysis and FTIR spectra revealed that Eucheuma isiforme from Nicaragua is a good source of relatively pure iota-carrageenan with sufficient quality to serve as a substitute for traditional iota-carrageenan sources.     

Halarachnion (Furcellariaceae,Rhodophyta), a newly reported genus for the Gulf of Mexico,with the description of Halarachnion louisianensis sp. nov

Halarachnion is a newly reported genus for the Gulf of Mexico where it is represented by an undescribed species growing attached to rhodoliths at 52–73 m depth offshore Louisiana in the northwestern Gulf, USA. Aspects of the vegetative and female reproductive morphology of the Gulf species are documented, and compared to of Halarachnion latissimun from Japan. Halarachnion louisianensis sp. nov. is a monoecious, presumably heteromorphic species as inferred from the absence of tetrasporophytic specimens in our collections. Halarachnion latissimum, in contrast, is a dioecious species that is isomorphic. A distinct reproductive character that differentiates both species is the behavior of a connecting filament of which a lateral process fuses with an auxiliary cell lobe and effects more than one diploidization in H. louisianensis versus directly fusing with an auxiliary cell and effecting only a single diploidization in H. latissimum. Comparative chloroplast‐encoded rbcL sequence analysis indicates that the closest species relationship of isolates from the Gulf of Mexico remains equivocal, whereas reproductive morphology points to a closer affinity with H. ligulatum. This is the first report of the Furcellariaceae, a family in the Solieriaceae‐complex, for the Gulf of Mexico.     

Diluted seawater promoted the green tide of Ulva prolifera (Chlorophyta,Ulvales)

Ulva prolifera (Müller) J. Agardh is the main causative species of the 2008 Yellow Sea green tide incident. We investigated the influences of diluted seawater on the vegetative growth and reproductive cell formation of the alga. The thalli that were cultivated under low salinities (10‰ and 20‰) and low pH values (pH around 7.0) showed obvious and steady biomass gain, while those cultivated under high salinities (40‰) and relatively higher pH conditions (pH around 8.0) manifested significant biomass loss. The trend was, however, completely the opposite for reproductive cell formation and there were indications that enrichment, over very wide concentration ranges of both nitrogen and phosphate, could significantly promote vegetative growth. Results also indicated that relatively low salinity and low pH regimes boosted vegetative growth but were unfavorable for reproductive cell formation and vice versa. Based on these results, the possible origin and development mechanisms of the green tide event are discussed. Eutrophication in the Changjiang River estuary and adjacent sea areas, as well as extremely high freshwater inflows before, and during, the flood of 2007 – due to the full operation of large‐scale water facilities in the area – may have both played an important role in the formation and development of the green tide event.     

ULTRASTRUCTURE DURING DEVELOPMENT OF THE NUCLEUS OF BATOPHORA OERSTEDII (CHLOROPHYTA; DASYCLADACEAE)1

The primary nucleus of Batophora oerstedii J. Agardh like that of the related Acetabularia, undergoes a great increase in size throughout vegetative development. Formation of small secondary nuclei represents an irreversible stage in the development of reproductive structures in the gametangia. Changes observed in the course of the life cycle include: i) an increase from 3 to 200 μm diam, of the nucleus; ii) increase in number of nucleoli; iii) development of the perinuclear region; iv) increased pore density in the nuclear membrane; v) development of chromosomes in the nucleoplasm; vi) formation of secondary nuclei; and, vii) division of secondary nuclei.     

Taxonomic notes on Laurencia brongniartii (Rhodomelaceae, Rhodophyta)

The red alga Laurencia brongniartii J. Agardh (Rhodomelaceae, Ceramiales) is characterized by: (i) the production of four periaxial cells from each vegetative axial segment; (ii) the presence of two or three corps en cerise per superficial cortical cell and one per trichoblast cell; (iii) the production of a single tetrasporangium-bearing periaxial (fourth) cell per fertile segment; (iv) a tetrasporangial arrangement that is intermediate between perpendicular and parallel types; (v) procarps produced from the last-formed (fifth) periaxial cell of the terminal segment of a two-celled female trichoblast; and (vi) distally positioned spermatangial nuclei, in addition to known features. The production of a single tet-rasporangium-bearing periaxial cell per fertile segment allies this species to Laurencia similis Nam et Saito.     

FINE CHEMICAL STRUCTURE OF CARRAGEENAN FROM THE COMMERCIALLY CULTIVATED KAPPAPHYCUS STRIATUM (SACOL VARIETY) (SOLIERIACEAE,GIGARTINALES, RHODOPHYTA)1

The polysaccharide extracted from Kappaphycus striatum (Schmitz) Doty (sacol variety), a recently farmed eucheumoid seaweed, was characterized by Fourier transform infrared and NMR spectroscopy. The extract is composed mainly of 3‐linked β‐d ‐galactopyranosyl‐4‐sulfate residues alternating with 4‐linked 3,6‐anhydro‐α‐d ‐galactopyranosyl residues (κ‐carrageenan). Minor components include 3,6‐anhydrogalactopyranosyl‐2‐sulfate residues (ι‐carrageenan) and 6‐O‐methylgalactopyranosyl‐4‐sulfate residues (methylated carrageenan). The substitution pattern of carrageenan in the tribe Eucheumatoideae and the family Solieriaceae is revisited in terms of chemotaxonomic affiliation.     

CENTROCERAS INTERNITENS N. SP. (RHODOPHYCEAE,CERAMIACEAE) FROM THE WESTERN TROPICAL NORTH ATLANTIC OCEAN1

Centroceras internitens n. sp. (Rhodophyceae, Ceramiaceae) was found in the Florida Keys and Bermuda only as an epiphyte on Ceramium nitens (C. Agardh) J. Agardh. It is described on the basis of many collections, including an abundance of reproductive material representing all stages of the life history. Laboratory culture has not been successful. The type locality is a well-known limestone platform on the Atlantic Ocean side of West Summerland Key, Florida Keys. In addition to living material, C. internitens was found on eight herbarium specimens of Ceramium nitens at the United States National Museum, thereby establishing records of its occurrence throughout the Caribbean Sea and dating back to 1887.