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.     

LEACHIELLA PACIFICA,GEN. ET SP. NOV., A NEW PARASITIC RED ALGA FROM WASHINGTON AND CALIFORNIA

Leachiella pacifica, gen. et sp. nov., a marine alloparasitic red alga is described from Washington and California. Several species of Polysiphonia and Pterosiphonia are hosts for this parasite. The thallus is a white, multiaxial, unbranched pustule with rhizoidal filaments that ramify between host cells, forming numerous secondary pit connections with host cells. All reproductive structures develop from outer cortical cells. Tetrasporocytes, situated on stalk cells, undergo simultaneous, tetrahedral cleavage to form tetraspores. Spermatia are formed continuously by oblique cleavages of the elongate spermatial generating cells. This results in spermatial clusters consisting of 4–8 spermatia in an alternate arrangement. Carposporophyte development is procarpial. The carpogonium is part of a six-celled branch including a sterile cell that is formed by the basal cell. The carpogonial branch is attached laterally to an obovate supporting cell that also forms an auxiliary cell, presumably formed prior to fertilization. After fertilization the carpogonium temporarily fuses with the auxiliary cell apparently to transfer the diploid nucleus and initiate further fusion with the subtending supporting cell to form an incipient fusion cell. The auxiliary cell portion of this fusion cell divides to form gonimoblast initials that continue to divide, forming gonimoblast filaments whose terminal cells differentiate into carpospores. The remainder of the fusion cell enlarges by continual fusion with adjacent vegetative cells. The resultant carposporophyte consists of a basal, multinucleate fusion cell supporting a hemispherical cluster of gonimoblast filaments with terminally borne carpospores. Vegetatively, Leachiella resembles several other parasitic red algae but it is clearly separated by the procarp, carposporophyte development and structure, and tetrasporocyte cleavage.     

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.     

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.     

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.     

Amphithallia,a genus with four-celled carpogonial branches and connecting filaments in the Corallinales (Rhodophyta)

The South African marine alga Amphithallia crassiuscula, previously subsumed in the widely reported Synarthrophyton patena, is here re-described as a distinct species and genus. Thalli grow as obligate epiphytes on Gelidium capense in the upper sublittoral zone (while S. patena grows on Ballia callitricha). Gametophytes are monoecious with four-celled carpogonial branches and sterile cells are borne on supporting cells (dioecious or hermaphroditic with two or three-celled carpogonial branches and sterile cells borne on hypogynous cells in Synarthrophyton). Postfertilization stages involve a connecting filament linking the carpogonium to several putative auxiliary cells, demonstrating a non-procarpic condition with apparent absence of a fusion cell. Gonimoblast filaments develop at the level of basal cells of carpogonial branches. Spermatangial mother cells remain either unbranched (cutting off spermatangia only) or develop dendroid (branched) filaments with terminal spermatangia (as in Synarthrophyton). Multiporate conceptacles develop straight pore canals lined by non-differentiated cells (conical canals with differentiated pore cells along the base in Synarthrophyton). The here described pre- and post-fertilization characters are new for the order Corallinales motivating the establishment of the new genus Amphithallia.     

OBSERVATIONS ON THE DEVELOPMENT OF THE CARPOSPOROPHYTE OF SCINAIA PSEUDOJAPONICA YAMADA ET TANAKA (NEMALIALES,CHAETANGIACEAE)1

The development of the carposporophyte of Scinaia pseudojaponica Yamada et Tanaka is described for the first time. The carpogonial branch is 3-celled. Before fertilization the hypogynous cell divides into a group of 4 cells. Concurrently the cell beneath the hypogynous cells also produces initials which, following fertilization, develop into branched filaments that envelop the carposporophyte. After fertilization the gonimoblast initial is produced laterally from the basal part of the carpogonium. Carposporangia are produced in chains from the free ends of the gonimoblast filaments which grow toward the surface of the thyllus. A very thick pericarp surrounds the mature carposporophyte.     

DEVELOPMENT OF THE CARPOSPOROPHYTE OF KALLYMENIA RENIFORMIS (TURNER) J. AGARDH1

The development of the carposporophyte in Kallymenia reniformis involves an elaborate series of interactions between reproductive and vegetative tissues. Following fertilization, the inner cells of the carpogonial branches form processes that unite with the supporting cell and with each other, giving rise to a large fusion cell. A number of medullary filaments are enveloped and incorporated within the developing fusion cell. Secondary filaments may be produced from medullary cells outside the fusion cell after connecting filaments have been initiated. Connecting filaments are nonseptale and wind their way through the medulla. The presence of a connecting filament in the vicinity of an auxiliary cell appears to initiate a complex sequence of responses. Vegetative filaments are produced in the medulla and inner cortex that grow centripetally toward the auxiliary cell and fuse with one another, forming a network of secondary tissue. One of the vegetative cells may penetrate the auxiliary cell, while others intrude into subsidiary cells connecting them with the network. An ostiolate pericarp is initiated in the cortex above the auxiliary cell. The connecting filament fuses with the auxiliary cell and also with some of the surrounding vegetative cells, forming an irregular lobed structure. Gonimoblast is initiated in scattered pockets from vegetative cells in the vicinity of the auxiliary cell apparatus.     

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.     

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.     

COMPARATIVE DEVELOPMENT OF THE RED ALGAL PARASITES BOSTRYCHIOCOLAX AUSTRALIS GEN. ET SP. NOV. AND DAWSONIOCOLAX BOSTRYCHIAE (CHOREOCOLACACEAE,RHODOPHYTA)1

The development of two red algal parasites was examined in laboratory culture. The red algal parasite Bostrychiocolax australis gen. et sp. nov., from Australia, originally misidentified as Dawsoniocolax bostrychiae (Joly et Yamaguishi-Tomita) Joly et Yamaguishi-Tomita, completes its life history in 6 weeks on its host Bostrychia radicans (Montagne) Montagne. Initially the spores divide to form a small lenticular cell, and then a germ tube grows from the opposite pole. Upon contact with the host cuticle, the germ tube penetrates the host cell wall. The tip of the germ tube expands, and the spore cytoplasm moves into this expanded tip. The expanded germ tube tip becomes the first endophytic cell from which a parasite cell is cut off that fuses with a host tier cell. The nuclei of this infected host cell enlarge. As parasite development continues, other host-parasite cell fusions are formed, transferring more parasite nuclei into host cells. The erumpent colorless multicellular parasite develops externally on the host, and reproductive structures are visible within 2 weeks. Tetrasporangia are superficial and cruciately or tetra-hedrally divided. Spermatia are formed in clusters. The carpogonial branches are four-celled, and the carpogonium fuses directly with the auxiliary (support) cell. The mature carposporophyte has a large central fusion cell and sympodially branched gonimoblast filaments. Early stages of development differ markedly in Dawsoniocolax bostrychiae from Brazil. Upon contact with the host, the spore undergoes a nearly equal division, and a germ tube elongates from the more basal of the two spore cells, penetrates the host cell wall, and fuses with a host tier cell. Subsequent development involves enlargement of the original spore body and division to form a multicellular cushion, from which descending rhizoidal filaments form that fuse with underlying host cells. This radically different development is in marked contrast to the final reproductive morphology, which is similar to B. australis and has lead to taxonomic confusion between these two entities. The different spore germination patterns and early germ-ling development of B. australis and D. bostrychiae warrant the formation of a new genus for the Australian parasite.     

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.     

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).     

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.     

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.     

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.     

Ultrastructure of post-fertilization development in the red alga Scinaia articulata (Galaxauraceae,Nemaliales, Rhodophyta)

The ultrastructure of the carposporophyte and carposporogenesis is described for the red alga Scinaia articulata Setch. After fertilization, the trichogyne disappears, and the pericarp develops to form a thick protective tissue that surrounds the carposporophyte. The hypogynous cell cuts off both one-celled and two-celled sterile branches. Patches of chromatin are frequently observed in evaginations of the nuclear envelope, which appear to produce vesicles in the cytoplasm of the cell of the sterile branch. Large gonimoblast lobes extend from the carpogonium and cleave to form gonimoblast initials. Subsequently, a fusion cell is formed from fusions of the carpogonium, the hypogynous cell and the basal cell of the carpogonial branch. The mature carposporophyte comprises the fusion cell that is connected to the sterile branch cells, gonimoblast cells and carpospores and is surrounded by extensive mucilage. Young carpospores possess a large nucleus and proplastids with a peripheral thylakoid, but they have few dictyosomes and starch granules and are indistinguishable from gonimoblast cells. Subsequently, dictyosomes are formed, which produce vesicles with an electron-dense granule, which indicates an initiation of wall deposition. Thylakoid formation coincides with incipient starch granule deposition. The nuclear envelope produces fibrous vacuoles and concentric membrane bodies. Carpospores are interconnected by pit connections with two cap layers. Dictyosome activity increases, resulting in the production of vesicles, which either continue to deposit wall material or coalesce to form fibrous vacuoles. The final stage of carposporogenesis is characterized by the massive production of cored vesicles from curved dictyosomes. Mature carpospores are uninucleate and contain fully developed chloroplasts, numerous cored vesicles, numerous starch granules and fibrous vacuoles. The mature carpospore is surrounded by a wall layer and a separating layer, but a carposporangial wall is lacking.     

STRUCTURE AND DEVELOPMENT OF THE CARPOSPOROPHYTE OF NEMALION HELMINTHOIDES (NEMALIONALES,RHODOPHYTA)1

Following fertilization, the carposporophyte of Nemalion helminthoides (Velley in With.) Batters differentiates into four distinct regions: the fusion cell, the sterile gonimoblast cells, the carposporangial mother cells and the carposporangia. The gonimoblast is formed by apically dividing, monopodial filaments of limited growth which may later become pseudodichotomous. Upon differentiation of a terminal carposporangium, a gonimoblast filament may continue to grow sympodially. A single carposporangial mother cell may produce carposporangia in several different directions as well as proliferate successive carposporangia within the sporangial walls that remain after carpospore liberation. As the carposporophyte matures, the gonimoblast initial, the stalk cell, the hypogynous and subhypogynous cells fuse. Except for the fusion cell, all cells of the carposporophyte show organelle polarity and contain a distally located, lobed chloroplast and proximal nucleus.