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.     

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.     

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 .     

The morphology and ecology of Nemastoma and Predaea species (Nemastomataceae,Rhodophyta) from Ghana

Two members of the family Nemastomataceae (Gigartinales, Rhodophyta) are described from subtidal habitats in Ghana. Nemastoma confusum sp. nov. is a plant of irregularly lobed, thick gelatinous blades with subacute marginal projections and surface proliferations. It is composed of a lax medulla and submoniliform cortical filaments with prominent intercalary gland cells. Carposporophytes are one to three spherical lobes of carposporangia borne on gonimoblast initials arising directly from auxiliary cells contacted by connecting filaments. A rudimentary involucre is formed around the gonimoblast by elongating vegetative cortical cells borne on the auxiliary cell. The genus Predaea is recorded for the first time from Africa, and P. feldmannii Boerg. is described in morphological detail together with some observations on its ecology in Ghana. Distinctive features of connecting filament formation, nutritive cell production and gonimoblast initiation and development are illustrated and compared to other species of the genus. A second species, P. masonii (Setch. & Gardn.) De Toni fil., is represented by a single specimen in the collections and appears to be distinct from P. feldmannii on cortical and gland cell features.     

Molecular phylogeny and developmental studies of Apoglossum and Paraglossum (Delesseriaceae,Rhodophyta) with a description of Apoglosseae trib. nov.

Our morphological and molecular studies indicate that species from the southern hemisphere previously placed in Delesseria belong in Paraglossum and that Paraglossum and Apoglossum comprise a separate tribe, the Apoglosseae, S.-W. Lin, Fredericq & Hommersand, trib. nov., within the family Delesseriaceae. From a vegetative perspective the Apoglosseae is readily recognized because some or all fourth-order cell rows are formed on the inner sides of third-order cell rows. All fourth-order cell rows grow adaxially in Apoglossum, whereas both adaxial and abaxial cell rows are present in Paraglossum. Periaxial cells do not divide in Apoglossum, whereas they divide transversely in Paraglossum in the same way as in Delesseria. Major branches are formed mainly from the margins of midribs in the Apoglosseae. The procarp consists of a straight carpogonial branch and two sterile cells, with the second formed on the same side as the first. The carpogonium cuts off two connecting cells in tandem from its apical end, the terminal cell being nonfunctional and the subterminal cell typically fusing with the auxiliary cell. Gonimoblast filaments radiate in all directions from the gonimoblast initials and produce carposporangia terminally in branched chains, with pit connections between the inner gonimoblast cells broadening and enlarging. The auxiliary cell, supporting cell, and sterile cells unite into a fusion cell, which remains small in Apoglossum but incorporates the branched inner gonimoblast filaments and cells in the floor of the cystocarp in Paraglossum. Elongated inner cortical cells seen in mature cystocarps in the Delesserieae are absent in the Apoglosseae. Phylogenetic studies based on rbcL (RuBisCO large subunit gene) sequence analyses strongly support the recognition of the Apoglosseae within the subfamily Delesserioideae of the Delesseriaceae, in agreement with our previous observations based primarily on analyses of large subunit ribosomal DNA (LSU).     

PROPOSAL OF THE GRACILARIALES ORD. NOV. (RHODOPHYTA) BASED ON AN ANALYSIS OF THE REPRODUCTIVE DEVELOPMENT OF GRACILARIA VERRUCOSA1

The mode of division of vegetative cells, formation of spermatangial parent cells, initiation of the carpogonial branch apparatus, and formation of tetrasporangial initials are homologous developmental processes that are documented for the first time in the type species of the economically important family Gracilariaceae, Gracilaria verrucosa (Hudson) Papenfuss from the British Isles. G. verrucosa is characterized by a supporting cell of intercalary origin that bears a 2-celled carpogonial branch flanked by two sterile branches, direct fusion of cells of sterile branches onto the carpogonium, formation of an extensive carpogonial fusion cell through the incorporation of additional gametophytic cells prior to gonimoblast initiation, gonimoblast initials produced from fusion cell lobes, schizogenous development of the cytocarp cavity, inner gonimoblast cells producing tubular nutritive cells that fuse with cells of the pericarp or floor of the cystocarp, absence of cytologically modified tissue in the floor of the cystocarp, and carposporangial initials produced in clusters or irregular chains. Spermatangial parent cells are generated in flaments from intercalary cortical cells that line an intercellular space forming a ‘pit’ or ‘conceptacle’. Tetrasporangial initials are transformed from terminal cells derived through division of an outer cortical cell. Tetrasporangia are cruciately divided. The Gracilariaceae is removed from Gigartinales and transferred to the new order Gracilariales. Their closest living relatives appear to be agarophytes belonging to the Gelidiales and Ahnfeltiales.     

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.     

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.     

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.     

Morphology and molecular relationships of Leptofauchea rhodymenioides (Rhodymeniales,Rhodophyta), a new record for Japan

Leptofauchea rhodymenioides Taylor (Faucheaceae, Rhodymeniales) is reported from Japan for the first time, based on detailed morphological studies and molecular phylogenetic analyses of nuclear‐encoded small subunit ribosomal RNA (SSU rRNA) and plastid‐encoded rbcL gene sequences. This is the first report of male gametophytes and detailed carposporophyte development in the genus Leptofauchea. This species is characterized as follows: (i) flat, membranous, and regularly and dichotomously branched thalli; (ii) the older blades are constricted below the apices; (iii) the cortex is composed of a continuous layer with an irregularly arranged outer layer, and the medulla of two to three incomplete layers; (iv) gametophytes are dioecious; (v) in males, the cortical cells cut off two to three spermatangial mother cells, which produce terminal spermatangia; (vi) in females, the procarp is composed of a three‐celled carpogonial branch and a two‐celled auxiliary cell branch; (vii) upon fertilization, the carpogonium directly contacts the auxiliary cell; (viii) the auxiliary mother cell fuses with vegetative cells, and forms a large trunk‐like fusion cell; (ix) gonimoblast filaments develop outwardly, and transform completely into carposporangia; (x) the carposporophyte is covered with a pericarp with a well‐defined tela arachnoidea; (xi) the mature cystocarp is spherical, has an ostiole, and protrudes from the blade margins; and (xii) the cruciately divided tetrasporangia are formed in nemathecia, produced laterally from paraphyses or terminally on short filaments. Molecular analyses suggest that Leptofauchea forms a strong sister alliance with the genus Webervanbossea. The families Faucheaceae and Lomentariaceae, and the genera Leptofauchea and Webervanbossea are monophyletic, but the latter two genera are not included in the Faucheaceae.     

VEGETATIVE AND REPRODUCTIVE MORPHOLOGY OF EUCHEUMA ISIFORME (SOLIERIACEAE,GIGARTINALES, RHODOPHYTA)1

Eucheuma isiforme (C. Agardh) J. Agardh exhibits a combination of vegetative and reproductive features that distinguish it from other critically studied genera in the Solieriaceae. The development of the multiaxial thallus, emphasizing the arrangement of periaxial cells around each axial file; presence of reproductive nemathecia that contain carpogonial branches and auxiliary cells; and post-diploidization stages, including gonimoblast and pericarp initiation, stages of fusion cell formation, and carposporophyte development are described and illustrated for the first time in this species. The vegetative and reproductive features observed in E. isiforme are not diagnostic of any of the recently erected tribes in the Solieriaceae. Eucheuma appears most closely related to the Indian Ocean genus, Sarconema.     

Studies of the Liagoraceae (rhodophyta) of Western Australia: Gloiotrichus fractalis gen. et sp. nov. and Ganonema helminthaxis sp. nov.

Two new taxa of Liagoraceae (Nemaliales) are described from Western Australia. Gloiotrichus fractalis gen. et sp. nov. has been collected from 3–20 m depths at the Houtman Abrolhos, Western Australia. Plants are calcified, extremely lubricous, and grow to 17 cm in length. Carpogonial branches are straight, 6 or 7 cells in length, arise from the basal or lower cells of cortical fascicles, and are occasionally compound. Branched sterile filaments of narrow elongate cells arise on the lower cells of the carpogonial branch prior to gonimoblast initiation, at first on the basal cells, then on progressively more distal cells. Following presumed fertilisation the carpogonium divides transversely, with both cells giving rise to gonimoblast filaments. The distal cells of the carpogonial branch then begin to fuse, with fusion progressing proximally until most of the cells of the carpogonial branch are included. As fusion extends, the filaments on the carpogonial branch are reduced to the basal 2 or 3 cells. The gonimoblast is compact and bears terminal carposporangia. Spermatangial clusters arise on subterminal cells of the cortex, eventually displacing the terminal cells. The sequence of pre- and post-fertilisation events occurring in the new genus separates it from all others included in the Liagoraceae, although it appears to have close affinities with the uncalcified genus Nemalion. Ganonema helminthaxis sp. nov. was collected from 12 m depths at Rottnest Island, Western Australia. Plants are uncalcified and mucilaginous, the axes consisting of a few (< 10) primary medullary filaments, each cell of which gives rise to a cortical fascicle at alternate forks of the pseudodichotomies borne on successive medullary cells. Subsidiary (adventitious) filaments and rhizoids comprise the bulk of the thallus. Carpogonial branches are straight, (3-)4(-6) cells in length, arise on the basal 1–4 cells of the cortical fascicles, and are frequently compound. Carposporophytes develop from the upper of two daughter cells formed by a transverse division of the fertilised carpogonium. Ascending and descending sterile filaments girdle the carpogonial branch cells and arise mostly on the supporting cell prior to fertilisation. Ganonema helminthaxis is the first completely non-calcified member of the genus, and its reproductive and vegetative morphology supports the recognition of Ganonema as a genus independent from Liagora. Liagora codii Womersley is a southern Australian species displaying features of Ganonema, to which it is transferred.     

A Light Microscope Study of Carposporophyte Development in Gracilaria verrucosa (Hudson) Papenfuss

Gracilaria verrucosa is a very common marine red alga of Greekcoasts. The diploid carposporophyte which develops attachedto the female gametophyte of Gracilaria is described. The immaturecystocarps are very small while the mature ones are globose,ostiolate and are borne profusely all over the surface of thethallus. The earliest observed fusion cell is small and fusesprogressively with adjacent vegetative cells to form a largemultinucleate cell. From this fusion cell gonimoblast initialsoriginate, dividing further and giving rise to a large numberof gonimoblast cells. The resultant carposporophyte consistsof a basal-central, multinucleate cell surrounded by a conicalor hemispherical mass of gonimoblast cells. Chains or clustersof successively maturing carpospores are borne from the terminalgonimoblast cells. The liberation of mature carpospores takesplace through the ostiole of the cystocarp. The liberated carposporeslack cell walls and are naked in a mucilage mass. Gracilaria verrucosa (Hudson) Papenfuss, Gigartinales, Gracilariaceae, Rhodophyta, carposporophyte, development     

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.     

New Insights in the Cryptonemiales–Rhodymeniales Complex and Resurrection of the Allied Red Algal Order Nemastomatales Kylin 1925 as Inferred From rbcL Sequence Analysis and Comparative Reproductive Morphology

In a global molecular phylogeny of florideophycean red algae inferred from chloroplast‐encoded rbcL sequence analysis, a major monophyletic assemblage comprises the Cryptonemiales (=Halymeniales), the Rhodymeniales, the Schizymeniaceae (Schizymenia, Titanophora, Platoma) and the Nemastomataceae (Nemastoma, Predaea). The phylogenetic significance of the auxiliary cell and its interaction with the fertilized egg cell in this assemblage is discussed in relation to established and newly proposed classification schemes. The order Nemastomatales Kylin 1925 is reinstated and emended to contain the nonprocarpic Schizymeniaceae and Nemastomataceae. Unifying characters of the Nemastomatales include fertilized carpogonia that may establish fusions with carpogonial nutritive cells prior to the formation of septate connecting filaments, and simple gonimoblasts developing outwardly from auxiliary cells or from connecting filaments in their vicinity. The auxiliary cell is a transformed vegetative intercalary cell (Sebdeniaceae), that becomes surrounded by either clusters of nutritive cells (Nemastomataceae), involucral filaments (Schizymeniaceae) or by three‐dimensional ampullary filaments (Halymeniaceae including the Corynomorphaceae), or is part of a procarp (Rhodymeniales). The homology of outward gonimoblast initiation and maturation into a simple ball of carposporangia in the Cryptonemiales, Rhodymeniales and Nemastomatales will be illustrated.     

A NEW METHOD OF CYSTOCARP DEVELOPMENT IN THE RED ALGAL GENUS CALLOPHYLLIS (KALLYMENIACEAE) FROM CHILE1

Traditional studies suggest that the Kallymeniaceae can be divided into two major groups, a nonprocarpic Kallymenia group, in which carposporophyte formation involves an auxiliary cell branch system separate from the carpogonial branch system, and a procarpic Callophyllis group, in which the carpogonial branch system gives rise to the carposporophyte directly after fertilization. Based on our phylogenetic studies and unpublished observations, the two groups each contain both procarpic and nonprocarpic genera. Here, we describe a new method of reproductive development in Callophyllis concepcionensis Arakaki, Alveal et Ramírez from Chile. The carpogonial branch system consists of a supporting cell bearing both a three‐celled carpogonial branch with trichogyne and two‐lobed “subsidiary” cells. After fertilization, large numbers of secondary subcortical and medullary cells are produced. Lobes of the carpogonial branch system cut off connecting cells containing enlarged, presumably diploid nuclei that fuse with these secondary vegetative cells and deposit their nuclei. Derivative enlarged nuclei are transferred from one vegetative cell to another, which ultimately cut off gonimoblast initials that form filaments that surround the central primary medullary cells and produce carposporangia. The repeated involvement of vegetative cells in gonimoblast formation is a new observation, not only in Callophyllis, but in red algae generally. These results call for a revised classification of the Kallymeniaceae based on new morphological and molecular studies.     

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.     

Studies on Metamastophora (Corallinaceae,Rhodophyta). I. M. flabellata (Sonder) Setchell: Morphology and anatomy

A morphological-anatomical study of Australian populations of Metamastophora flabellata (Sonder) Setchell, the type species of Metamastophora (Corallinaceae, Rhodophyta), has revealed that the primarily erect or ascending non-geniculate thallus possesses a dorsi-ventral organization of tissues. All conceptacles are uniporate and arise dorsally. Two distinct vegetative meristems occur: an apical primary meristem from which hypothallial cells are produced basipetally and a sub-epithallial secondary meristem which generates perithallial cells basipetally and secondary epithallial cells acropetally. Primary epithallial cells arise from divisions of subapical hypothallial cells. In younger parts, tissues are produced only dorsal to the hypothallium; in veins and stipes, tissue production occurs both dorsal and ventral to the hypothallium. Mature tetrasporic conceptacles contain peripheral tetrasporangia with zonately divided contents and a central sterile columella. Gametic conceptacles produce fertile tissue across the entire conceptacle chamber floor. After fertilization, the zygotic nucleus or a derivative is transferred (presumably) to an auxiliary cell through cells of the carpogonial branch; no tubular transfer siphon develops. Mature fusion cells are composed of the amalgamated supporting cells of carpogonial branches and are initiated from a single supporting cell which functions as an auxiliary cell. Unbranched 3–4 celled gonimoblast filaments arise from the fusion cell, do not become connected to other cells, and produce terminal carposporangia. Results from this study have led to a redefinition of hypothallium and perithallium in relation to meristems rather than substrate. In addition, carposporophyte ontogeny in the Corallinaceae is considered in terms of the presumed mode of transfer of the zygotic nucleus to the fusion cell, the extent of fusion cell development, and gonimoblast filament production in relation to auxiliary cells and fusion cells.     

Reproductive morphology of Gastroclonium pacificum (Champiaceae,Rhodymeniales) from Japan

Morphological studies were undertaken on Gastroclonium pacificum (E.Y. Dawson) C.F. Chang et B.M. Xia (Champiaceae, Rhodymeniales) from Japan. We describe the details of male reproductive structures, the procarp and early post‐fertilization stages. This species has a solid axis, with both tetrasporangia and polysporangia, spermatangia are cut off from spermatangial parent cells, and a procarp is composed of a 4‐celled carpogonial branch and two 2‐celled auxiliary cell branches. The mature cystocarp lacks a conspicuous ostiole, a characteristic of the genus Gastroclonium. The most distinctive characteristic of the species is the tela arachnoidea, which is lacking in other species of Gastroclonium.     

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.