Erythroglossum latum sp. nov. (Delesseriaceae, Rhodophyta) from the east coast of Honshu, Japan

A new species, Erythroglossum latum, is described based on specimens from Fukushima Prefecture along the east coast of Honshu, Japan. This species differs from Erythroglossum pinnatum Okamura in having a broader main blade, a clear midrib and lateral branches that are smaller in size and not clearly constricted at the base.     

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.     

Drachiella liaoii sp. nov., a new member of the Schizoserideae (Delesseriaceae,Rhodophyta) from Taiwan and the Philippines

A new member of Delesseriaceae (Ceramiales, Rhodophyta) is described from Southern Taiwan and the Philippines. On the basis of comparative vegetative and reproductive morphology, and phylogenetic analysis inferred from nuclear-encoded large-subunit ribosomal DNA sequences (LSU rDNA), we conclude that it belongs in the genus Drachiella, tribe Schizoserideae, subfamily Phycodryoideae. The new taxon shares with other Drachiella species the absence of macro- and microscopic veins; diffuse growth by marginal and intercalary meristematic cells; a polystromatic, lobed thallus; abundance of rhizoidal marginal proliferations used for attachment; convoluted plastids in surface cells; abundant secondary pit connections among adjacent vegetative cells; large intercellular spaces between surface cells; procarps confined to the upper side of the thallus, circular in outline, consisting of a supporting cell bearing a strongly curved carpogonial branch and two sterile groups that remain undivided; vertical division of gonimoblast initial from auxiliary cell, and unilateral, monopodial branching of gonimoblasts; and mature cystocarps with a massive candelabrum-like fusion cell of fused gonimoblasts bearing carposporangia in branched chains. It is distinguished from the other members of the genus by thalli that consist of extensive tangled mats of prostrate and overlapping decumbent blades, procarps confined to the upper side of the thallus, and the lack of basal stalks or stipes. Whereas the Schizoserideae is predominantly a Southern Ocean tribe, one of the tribe's four genera, Drachiella, was known only from the eastern Atlantic and Mediterranean. We herein report the first record of the genus for the Indo-Pacific Ocean, and describe Drachiella liaoii, sp. nov., as a fourth species in the genus.     

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.     

Phycodrys valentinae sp. nov. (Delesseriaceae, Rhodophyta) with Discussion of Other Species of the Phycodrys Genus from the North Pacific

A new species Phycodrys valentinae Seliv. et Zhigad. from the northwest of the Bering Sea, distinct from other species of the genus Phycodrys by the presence of a distromatic lamina and by the arrangement of basal generative prolifications, was described. Based on the analysis of the morphological and anatomic traits of species of the genus Phycodrys, P. serratiloba (Rupr.) A. Zin., which was considered as synonym of P. riggii Gardn. to the present [27, 28, 30], was reinstated as an independent species. The taxonomic status of other representatives of the genus Phycodrys of the studied region is considered. In our opinion, five species of the genus Phycodrys (P. serratiloba, P. riggii, P. amchitkensis, P. vinogradovae, and P. valentinae) grow in the seas of the Russian Far East.     

Polysiphonia freshwateri sp. nov. and Polysiphonia koreana sp. nov.: two new species of Polysiphonia (Rhodomelaceae,Rhodophyta) from Korea

Polysiphonia sensu lato comprises approximately 200 species, which are currently assigned to several different genera. To date, one of these genera, namely, Polysiphonia, has been reported to have 17 species. Here, we describe for the first time P. freshwateri sp. nov. and P. koreana sp. nov. from Uljin and Ulleung Island, Korea, based on morphological and molecular evidence. Polysiphonia freshwateri sp. nov. and P. koreana sp. nov. are characterized by having the typical Polysiphonia features. Polysiphonia freshwateri sp. nov. is further characterized by having abundant trichoblasts, conspicuous scar cells, and tetrasporangia arranged in spiral series. Polysiphonia koreana sp. nov. is further characterized by having very scarce scar cells placed between two pericentral cells, from which cicatrigenous branches arise. The results of our rbcL sequence analyses support the taxonomic placement of P. freshwateri sp. nov. and P. koreana sp. nov. within Polysiphonia.     

Taxonomic revision of the Agaraceae with a description of Neoagarum gen. nov. and reinstatement of Thalassiophyllum

We confirmed the monophyly of the Agaraceae based on phylogenetic analyses of six mitochondrial and six chloroplast gene sequences from Agarum, Costaria, Dictyoneurum, and Thalassiophyllum species, as well as representative species from other laminarialean families. However, the genus Agarum was paraphyletic, comprising two independent clades, A. clathratum/A. turneri and A. fimbriatum/A. oharaense. The latter clade was genetically most closely related to Dictyoneurum spp., and morphologically, the species shared a flattened stipe bearing fimbriae (potential secondary haptera) in the mid‐ to upper portion. The phylogenetic position of Thalassiophyllum differed between the two datasets: in the chloroplast gene phylogeny, Thalassiophyllum was included in the A. clathratum/A. turneri clade, but in the mitochondrial gene phylogeny, it formed an independent clade at the base of the Agaraceae, the same position it took in the phylogeny when the data from both genomes were combined despite a larger number of bp being contributed by the chloroplast gene sequences. Considering the remarkable morphological differences between Thalassiophyllum and other Agaraceae, and the molecular support, we conclude that Thalassiophyllum should be reinstated as an independent genus. Dictyoneurum reticulatum was morphologically distinguishable from D. californicum due to its midrib, but because of their close genetic relationship, further investigations are needed to clarify species‐level taxonomy. In summary, we propose the establishment of a new genus Neoagarum to accommodate A. fimbriatum and A. oharanese and the reinstatement of the genus Thalassiophyllum.     

Two new species of Martensia (Delesseriaceae, Rhodophyta) from Jeju Island, Korea

Two new species of Martensia (Delesseriaceae, Rhodophyta) are described from Jeju Island, Korea. Martensia jejuensis is characterized by the thallus of long linear axes and a proximal membranous part (initial blade), the linear axes that are derived from the longitudinal lamellae of the latticework, the flabellate bladelets arising from the linear axes terminally and laterally, the leading margins of the latticework giving rise to spine‐like or spatulate projections at intervals, and tetrasporangia borne on the whole thallus except for the spatulate projections. Martensia bibarii is characterized by the fimbriate aspect, the ribbon‐like blade with lobes, the flabellate bladelets derived from the lobes of the blade, the bladelets bearing the fringe or the latticework distally, the fringe consisting of many fine and strap‐shaped lamellae, and tetrasporangia borne only on the blade.     

Comparative morphology and systematics of Chondrymenia lobata from the Mediterranean Sea and a phylogeny of the Chondrymeniaceae fam. nov. (Rhodophyta) based on rbcL sequence analyses

The Chondrymeniaceae Rodríguez-Prieto, G. Sartoni, S.-M. Lin & Hommersand, fam. nov., is proposed for Chondrymenia lobata. Analyses of rbcL sequences place the new family in a large gigartinalean assemblage that comprises the Cystocloniaceae–Solieriaceae complex. Plants are decumbent and growth takes place by division of multiple apical cells at the margin of the blade. Thalli consist of an outer cortex of subspherical to elongate cortical cells arranged in anticlinal rows, a subcortex of cells cross-linked by lateral arms, and a large central medulla composed of primary medullary filaments intermixed with numerous rhizoidal filaments. Male stages are reported in monoecious individuals. Inactive carpogonial branches consist of a two-celled filament that is directed inwards from the supporting cell. Functional carpogonial branches are oriented outwardly, with the carpogonia and trichogynes pointed towards the thallus surface. After presumed fertilization, the carpogonium fuses with the hypogynous cell and transfers the zygote nucleus. The hypogynous cell, in turn, fuses with the supporting cell which contains many haploid nuclei. The resulting fusion cell functions as an auxiliary cell that cuts off a single gonimoblast initial, which produces the gonimoblast filaments. Gametophytic cells close to the auxiliary cell unite with it to form a placental fusion network of variable size and outline, and a placental fusion cell. Proximal gonimoblast cells fuse with the placental fusion cell, while the distal cells differentiate into branched chains of subspherical carposporangia. The superficial similarity of the outwardly developed osteolate cystocarp is responsible for Kylin's (1956) placement of Chondrymenia in his family Sarcodiaceae; however, the manner in which the placenta is formed is more like that seen in the Cystocloniaceae–Solieriaceae complex.     

CHARACTERIZATION OF MYHOGRAMME LNIDA, MYRIOGRAMMEAE TRIB. NOV (DELESSEIUACEAE, FWODOPHYTA)

The Myriogramme group of Kylin was found to contain two distinct clusters of genera that merit recognition at the tribal level. In this paper, we establish the tribe Myriogrammae based on a study of the type species of Myriogramme, M. livida, from the Southern Hemisphere. The Myriogrammae is characterized by 1) marginal and diffuse intercalary meristems; 2) nuclei arranged in a ring bordering the side walls of vegetative cells; 3) microscopic veins absent; 4) procarps scattered, formed opposite one another on both sides of the blade posterior to one or more vegetative pericentral cells (cover cells) and consisting of a carpogonial branch, a one-/to two-celled lateral sterile group and a one-celled basal sterile group; 5) auxiliary cell diploidized by a connecting cell cut off posteriolaterally from the fertilized carpogonium; 6) gonimoblast initial cut off distally from the auxiliary cell, generating one distal and one to two lateral gonimoblast filaments that branch in the plane of the expanding cystocarp cavity and later fuse to from an extensive, branched fusion cell; 7) spermatangial and tatrasporangial sori formed inside the margin on both sides of the blade by resumption of meristematic activity; and 8) tetrasporangia produced primarily from the central cells. The Myriogrammae currently includes Myriogramme Kylin , Gonimocolax Kylin , Haraldiophyllum A. Zinova , Hideophyllum A. Zinova, and a possible undescribed genus from Pacific North and South America. Genera are separated based primarily on features of gonimoblast and carposporangial development .     

Comparative Genome Analysis Reveals Cyanidiococcus gen. nov., A New Extremophilic Red Algal Genus Sister to Cyanidioschyzon (Cyanidioschyzonaceae,Rhodophyta)

The taxonomic placement of strains belonging to the extremophilic red alga Galdieria maxima has been controversial due to the inconsistent phylogenetic position inferred from molecular phylogenetic analyses. Galdieria maxima nom. inval. was classified in this genus based on morphology and molecular data in the early work, but some subsequent molecular phylogenetic analyses have inferred strains of G. maxima to be closely related to the genus Cyanidioschyzon. To address this controversy, an isolated strain identified as G. maxima using the rbcL gene sequence as the genetic barcode was examined using a comprehensive analysis across morphological, physiological, and genomic traits. Herein are reported the chloroplast-, mitochondrion-, and chromosome-level nuclear genome assemblies. Comparative analysis of orthologous gene clusters and genome arrangements suggested that the genome structure of this strain was more similar to that of the generitype of Cyanidioschyzon, C. merolae than to the generitype of Galdieria, G. sulphuraria. While the ability to uptake various forms of organic carbon for growth is an important physiological trait of Galdieria, this strain was identified as an ecologically obligate photoautotroph (i.e., the inability to utilize the natural concentrations of organic carbons) and lacked various gene models predicted as sugar transporters. Based on the genomic, morphological, and physiological traits, we propose this strain to be a new genus and species, Cyanidiococcus yangmingshanensis. Re-evaluation of the 18S rRNA and rbcL gene sequences of the authentic strain of G. maxima, IPPAS-P507, with those of C. yangmingshanensis suggests that the rbcL sequences of “G. maxima” deposited in GenBank correspond to misidentified isolates.     

AUGOPHYLLUM,A NEW GENUS OF THE DELESSERIACEAE (RHODOPHYTA) BASED ON rbcL SEQUENCE ANALYSIS AND CYSTOCARP DEVELOPMENT1

A new genus, Augophyllum Lin, Fredericq et Hommersand gen. nov. related to Nitophyllum, tribe Nitophylleae, subfam. Nitophylloideae of the Delesseriaceae, is established to contain the type species Augophyllum wysorii Lin, Fredericq et Hommersand sp. nov. from Caribbean Panama; Augophyllum kentingii Lin, Fredericq et Hommersand sp. nov. from Taiwan; Augophyllum marginifructum (R. E. Norris et Wynne) Lin, Fredericq et Hommersand comb. nov. (Myriogramme marginifructa R. E. Norris et Wynne 1987) from South Africa, Tanzania, and the Sultanate of Oman; and Augophyllum delicatum (Millar) Lin, Fredericq et Hommersand comb. nov. (Nitophyllum delicatum Millar 1990 ) from southeastern Australia. Like Nitophyllum, Augophyllum is characterized by a diffuse meristematic region, the absence of macro‐ and microscopic veins, procarps consisting of a supporting cell bearing a slightly curved four‐celled carpogonial branch flanked laterally by a cover cell and a sterile cell, a branched multicellular sterile group after fertilization, absence of cell fusions between gonimoblast cells, and tetrasporangia transformed from multinucleate surface cells. Augophyllum differs from Nitophyllum by the blades becoming polystromatic inside the margins, often with a stipitate cylindrical base, the possession of aggregated discoid plastids neither linked by fine strands nor forming bead‐like branched chains, spermatangia and procarps initiated at the margins of blades, not diffuse, and a cystocarp composed of densely branched gonimoblast filaments borne on a conspicuous persistent auxiliary cell with an enlarged nucleus. Analyses of the rbcL gene support the separation of Augophyllum from Nitophyllum. An investigation of species attributed to Nitophyllum around the world is expected to reveal other taxa referable to Augophyllum.     

A taxonomic revision of a Hypnea charoides-valentiae complex (Rhodophyta, Gigartinales) in Japan, with a description of Hypnea flexicaulis sp. nov.

Morphology of a Hypnea charoides‐valentiae complex (Rhodophyta, Gigartinales) in Japan is investigated on the basis of field‐collected and cultured plants. We can recognize two species, Hypnea charoides Lamouroux and H. flexicaulis Yamagishi et Masuda, sp. nov., which are distinguished by: (i) conspicuous percurrent main axes growing straight in the former versus less conspicuous percurrent main axes growing flexuosely in the latter; (ii) narrow branching angles (less than 90°) in the former versus wide branching angles (up to 150°) in the latter; (iii) the presence of branches and branchlets showing abrupt adaxial bending in the former versus those showing pronounced abaxial bending, some growing into hooked branchlets in the latter; and (iv) the absence (in the former) or presence (in the latter) of antler‐like branches. Nucleotide sequences of the plastid‐encoded large subunit of the ribulose‐1,5‐bisphosphate carboxylase/oxygenase gene (rbcL) support the differentiation of these two entities: the pairwise distance between them was 83 b.p. (6.2%).     

Diversity of Chroothece (Rhodophyta,Stylonematales) including two new species

Chroothece has been reported from a range of freshwater environments, including streams, shallow ponds, trickling water on cliffs and moist soils, mostly in Europe and North America. The identification of genera and species by morphology is difficult because of overlaps in critical characters. To help clarify diversity within the genus, samples from Spain and from other regions (UK and Guam, western Pacific) were compared. Ecological and morphological data from field and cultured material were correlated with molecular data (rbcL gene sequences) that differentiate two new species: Chroothece thermalis I. Chapuis, P. Sánchez, M. Aboal & O. Necchi Jr., sp. nov. in a thermal spring and Chroothece lobata M. Aboal, B. A. Whitton, I. Chapuis, P. Sánchez & O. Necchi Jr., sp. nov. in a semi-arid stream. The results suggest recognition of four species, C. thermalis, C. lobata, C. richteriana Hansgirg and C. rupestris Hansgirg, from Spain. Morphology and ecology are useful to help distinguish these species, but the genus needs further study for possible cryptic diversity.     

Sorellocolax stellaris gen. et sp. nov., a hemiparasitic alga (Delesseriaceae, Rhodophyta) from the east coast of Honshu, Japan

A hemiparasitic alga, Sorellocotax stellaris sp. nov. is described growing on plants of Sorella repens collected from Onagawa, Miyagi Prefecture, east coast of Honshu, Japan. The thallus is small, up to 2 mm high, once or twice branched from the margin. The growing apex has a transversely dividing apical cell, and intercalary cell divisions occur in the cells of first-order rows. Tetrasporangia are cut off from the cells of the inner cortex, The procarp is composed of a supporting cell, one group of sterile cells and two carpogonial branches. Carposporangia are borne in short chains.     

Recent developments in the systematics of the Gigartinaceae (Gigartinales, Rhodophyta) based on rbcL sequence analysis and morphological evidence

The phylogenetic systematics of the Gigartinaceae is discussed for seven genera and three undescribed generic lineages and 65 taxa representing 62 species based on an analysis of rbcL sequences and morphological evidence. An examination of rbcL trees resulting from analyses of these taxa identifies seven lineages: (i) ‘Gigartina’ alveata; (ii) Rhodoglossum/Gigartina; (iii) Chondracanthus; (iv) Ostiophyllum; (v) Sarcothalia; (vi) ‘Gigartina’ skottsbergii; and (vii) a large clade containing Iridaea/‘Sarcothalia’, Mazzaella and Chondrus. These lineages and Chondrus are strongly supported; however, two groups, Iridaea/‘Sarcothalia’ and Mazzaella, receive no bootstrap support. The morphology of the female reproductive system is investigated with the aid of computer-generated, color-coded tracings of photographs of cystocarps seen in cross section at different developmental stages. Seven basic cystocarp types were found which corresponded to species groups seen in rbcL trees. These were: (i) a ‘Gigartina’ alveata group in which the carposporangia-bearing filaments develop apomictically from gametophytic cells; (ii) a Rhodoglossum/Gigartina group in which gonimoblast filaments penetrate the surrounding envelope fusing progessively with envelope cells; (iii) a Chondracanthus group in which gonimoblast filaments penetrate the envelope but fuse with envelope cells only at late developmental stages; (iv) a Sarcothalia group in which the gonimoblast filaments displace an envelope composed mainly of secondary gametophytic filaments and link to envelope cells by terminal tubular gonimoblast cells; (v) an Iridaea group similar to the Sarcothalia group, but with an envelope composed of a mixture of medullary cells and secondary gametophytic filaments; (vi) a Mazzaella group that lacks a true envelope and in which gonimoblast filaments connect to modified gametophytic cells by means of terminal tubular cells; (vii) a Chondrus group in which gonimoblast filaments penetrate the medulla and link to modified medullary cells by means of conjunctor cells forming secondary pit connections. The further separation of these groups into genera is based largely on tetrasporangial characters.     

CHARACTERIZATION OF SCHIZOSERIS CONDENSATA,SCHIZOSERIDEAE TRIB. NOV. (DELESSERIACEAE,RHODOPHYTA)1

The Myriogramme group of Kylin contains two distinct clusters of genera that merit recognition at the tribal level. We previously established the tribe Myriogrammeae, and in this paper we erect the Schizoserideae based on a study of the type species of Schizoseris, S. laciniata (=S. condensata), from the southern hemisphere. The Schizoserideae is characterized by 1) marginal and diffuse intercalary meristems; 2) nuclei initially arranged in a plate in the median plane in meristematic and mature cells; 3) chloroplasts one to few, lobed or dissected; 4) microscopic veins absent; 5) procarps scattered, formed singly on either side of the blade with cover cells absent and consisting of a one- to two-celled lateral sterile group, a one- to two-celled basal sterile group, and a four-celled carpogonial branch in which the trichogyne passes beneath the lateral sterile group and emerges anterior to it; 6) auxiliary cell diploidized by a connecting cell cut off posteriolaterally from the fertilized carpogonium; 7) gonimoblast initial cut off laterally from one side of the auxiliary cell and giving rise to unilaterally branched gonimoblast filaments bearing carposporangia in branched chains; 8) gonimoblast fusion cell highly branched, candelabra-like, incorporating all but the basalmost cells of the carposporangial chains and radiating through the central cells in the floor of the cystocarp; 9) spermatangial and tetrasporangial sori formed from surface cells in both monostromatic and polystromatic portions on both sides of the blade; and 10) tetrasporangia formed primarily from cortical rather than from central cells. The Schizoserideae presently includes Schizoseris Kylin, Neuroglossum Kützing, Abroteia J. Agardh, and Polycoryne Skottsberg in Kylin and Skottsberg.